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Cover. An adult burying beetle, Nicrophorus marginatus, tending her larvae in a brood ball, which she has 

helped to fashion with her mate, from a dead field mouse. Painting by David Reiser.

Bulletin 

of the 

UniveRsity of nebRaska state MUseUM 

volUMe 13 

________________________ 

The Carrion BeeTles (ColeopTera: silphidae) 

of neBraska 

by 

brett C. Ratcliffe 

Published by the University of nebraska state Museum 

lincoln, nebraska 

1996

Bulletin of the University of Nebraska State Museum 

volume 13 

issue Date: april 1996 

editor: brett C. Ratcliffe 

Production secretary: Gail littrell 

bulletins may be purchased from the Museum. 

address orders to: Publications secretary 

W436 nebraska Hall 

University of nebraska state Museum 

lincoln, ne 68588-0514 U.s.a. 

Price: $18.00 

Copyright © by the University of nebraska state Museum, 1996 

All rights reserved. Apart from citations for the purposes of research or review, no part 

of this Bulletin may be reproduced in any form, mechanical or electronic, including 

photocopying and recording, without permission in writing from the publisher. 

issn 0093-6812 

library of Congress Catalog Card number 

Printed in the United states of america 

the Bulletin is a peer-refereed journal.

C o n T e n T s 

introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

nebraska . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

Physical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

Climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 

vegetation of nebraska . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 

short and tallgrass Prairie . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 

sand Hills Prairie . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 

eastern Deciduous forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 

Rocky Mountain forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 

the silphidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 

Collecting silphids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 

subfamily silphinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 

Genus Aclypea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 

Genus Heterosilpha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 

Genus Necrodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 

Genus Necrophila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 

Genus Oiceoptoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 

Genus Thanatophilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 

subfamily nicrophorinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 

Genus Nicrophorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 

nicrophorine biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 

searching behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 

burial and Preparation of the Carcass . . . . . . . . . . . . . . . . 45 

Parental Care . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 

agonistic behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 

Mite Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 

nematode Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 

stridulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 

sociality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 

acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 

literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 

Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 

Checklist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 

about the author . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100

Bulletin of the 

University of Nebraska State Museum 

Volume 13 

The Carrion Beetles (Coleoptera: Silphidae) 

of Nebraska 

by 

Brett C. Ratcliffe 

Systematics Research Collections 

W436 Nebraska Hall 

University of Nebraska State Museum 

Lincoln, NE 68588-0514, U.S.A 

Email: bcr@unlinfo.unl.edu 

Abstract. A faunal study of the carrion beetles (Coleoptera: Silphidae) of Nebraska is 

presented. An overview of the family and its two subfamilies is given as well as keys to the 

adults and, when known, the larvae. Each of the six genera and 18 species is reviewed. 

The treatment for each species consists of synonymy, a brief diagnosis, distribution in 

general, Nebraska locality records, temporal distribution, and remarks. The remarks 

include commentary on how to best distinguish the adults, references to the larval stages, 

and a discussion of biology and ecology. 

Distribution maps, showing Nebraska county records, are given for all the species. A 

habitus drawing for each species is provided along with line drawings of particular characters to 

supplement the taxonomic keys. An extensive bibliography and a glossary are also included.

2 

BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM 

INTRODUCTION 

The Silphidae is a relatively small 

family of beetles, but interest in them has 

always been substantial because of the semisocial 

behavior displayed by species in the 

genus Nicrophorus. As currently defined, 

there are 13 genera and about 208 species 

found worldwide. North America has eight 

genera and 30 species. As a result of this 

study, silphids in the mid-continental state 

of Nebraska are now known to number six 

genera with 18 species (75% of the North 

American genera and 60% of the species). 

Those species are found in two subfamilies: 

the Silphinae with seven species and the 

Nicrophorinae with 11 species. 

This study was begun in 1990 in order 

to produce an identification manual that 

could be used by professional entomologists, 

students, interested amateurs, and 

an increasingly large number of biologists 

and ecologists who are conducting surveys 

for the endangered American burying beetle, 

Nicrophorus americanus Olivier. The intent 

has been to include as much information as 

possible about each species as well as illustrations 

and keys for identification of both 

adults and larvae. 

While other synoptic works have partially 

reviewed the North American fauna 

(e.g., Anderson and Peck 1985), this study 

marks the first time that an in-depth review 

of the Silphidae of any state has been provided. 

Meserve (1936) compiled a checklist 

of Nebraska silphids, but otherwise nothing 

of a comprehensive nature has been written 

for the state. 

MeThODS 

The results of this study were based 

on the examination and records of 29,719 

specimens. Most of the specimens are housed 

in the Systematics Research Collections 

(Division of Entomology) of the University of 

Nebraska State Museum. These collections, 

now numbering nearly two million specimens, 

are recognized as one of the top 20 collections 

in North America (Anonymous 1971, 

Fischer et al. 1975). They represent over 

a century of collecting and data gathering 

in the prairie biome. Additional data were 

gathered from the collections at Hastings 

College, Chadron State College, University 

of Nebraska at Kearney, and the University 

of Nebraska’s Cedar Point Biological Station 

in western Nebraska. 

Fig. 1. Number of silphid species recorded from each Nebraska county. Counties with no records or few records 

are clearly in need of additional collecting.

Extensive collecting was conducted 

across the state by myself and others (see 

Acknowledgments). Collecting techniques 

consisted of baited pitfall traps, light traps, 

whole animal bait stations, and examination 

of road-killed animals. While this collecting 

effort has been substantial, there are 

still areas of Nebraska that remain poorly 

known entomologically. Figure 1, showing 

the number of silphid species recorded from 

each Nebraska county, will give a general 

idea of where additional collecting could 

be done. Some counties have never been 

sampled for silphids, which is, of course, very 

different than silphids not occurring there. 

Conventional, artificial keys to all the 

silphids found in Nebraska are presented. I 

have attempted to use key characters that 

are consistently expressed, low in intrinsic 

variability, and easily observed with reasonable 

procedures (Figs. 2-3). The keys 

and descriptions are all accompanied by 

illustrations to aid the reader in correctly 

identifying specimens. The illustrations consist 

of line drawings and habitus drawings 

on pebble board. Dot maps showing county 

distributions are included to show where 

in the state beetles occur as exemplified by 

label data. 

Each genus and species-level taxon is 

introduced with its chronological, nomenclatural 

history. An abbreviated, descriptive 

diagnosis for each species then follows. 

This consists of range of length (from tip of 

clypeus to apex of elytra) followed by distinguishing 

characteristics of the head, thorax, 

elytra, and legs. 

Remarks on the overall distribution, 

and then the Nebraska distribution, are 

presented following the description. Fitzpatrick 

(1960) was the source for Nebraska 

place names. The locality data (accompanied 

by a reference to a map figure) is next 

and is followed by the temporal distribution, 

both rangewide and in Nebraska. The “Remarks” 

section is divided into distinguishing 

features of the adult, reference to larval 

descriptions, and life history and ecological 

information. 

THE CARRION BEETLES OF NEBRASKA 3 

By necessity, some technical terms 

(largely those dealing with body structure) 

have been used. A brief glossary is provided 

in the back of this work for those unfamiliar 

with these words. Definitions used are 

primarily those of Torre-Bueno (1937). A 

total of 242 references have been referred 

to in this work. 

Ph y s i c a l 

NeBRASKA 

Nebraska, one of the richest agricultural 

states in the nation, is located just north of 

the geographic center of the United States. 

Nebraska is found 40°-43° north of the equator 

and 95°25'-104° west of Greenwich. It 

occupies an area of 77,510 sq. miles (200,673 

sq. km.) and extends 462 miles (743 km.) 

east to west and 205 miles (330 km.) north to 

south (USGS data). The lowest elevation is 

825 ft. (251 m.) in Richardson County in the 

southeast. Elevation gradually rises to 5,340 

ft. (1,623 m.) in Banner and Kimball counties 

in the west near the Wyoming line. 

The chief rivers in the state are the 

Missouri (along the eastern border), Platte 

(its largest tributaries being the Loup and 

Elkhorn), Niobrara, Republican, and Big 

Blue. Surface drainage is generally from 

west to east. Aside from streams and 

rivers, Nebraska has about 2,500 lakes, 

marshes, and artificial reservoirs containing 

15 acres or more of water. The largest 

lake in the state (26 miles in length) 

is located in Keith and Garden counties. 

Most of the natural lakes and marshes are 

in the Sand Hills where, counting bodies 

of standing water of all sizes, the number 

exceeds 3,000 (Jones 1964). 

As is typical for all of the Great Plains, 

Nebraska’s geologic formations consist of 

deep granite or granite-like rocks, sedimentary 

bedrock in the form of shale, mudstone, 

sandstone, and limestone, and unconsolidated 

sediments (mantle rock) shaped by 

glaciers, water, and wind. The mantlerock 

is primarily Pleistocene in age whereas the

4 


Fig. 2. Dorsal aspect of adult Nicrophorus species showing morphological features.


Fig. 3. Ventral aspect of adult Nicrophorus species showing morphological features.

6 


bedrock ranges in age from the early Paleozoic 

to late Tertiary (Condra and Reed 1943). 

Three of the major soil divisions of North 

America are found in Nebraska in the form of 

six general soil associations. Three of these 

associations (brunizem, chernozem, and 

chestnut) are among the dark-colored soils 

that are found under prairie vegetation. The 

soils of Nebraska were described in detail by 

Elder (1969). 

The Nebraska Sand Hills is a remarkable 

physiographic region covering approximately 

19,300 square miles and stretching 

265 miles across Nebraska and into South 

Dakota. It is the largest sand dune area in 

the Western Hemisphere and is one of the 

largest grass-stabilized dune regions in the 

world (Bleed and Flowerday 1989). The 

Sand Hills are, for the most part, a treeless 

landscape of grass-covered sand dunes in 

the western two-thirds of Nebraska. Wright 

(1970) suggested that the sand that formed 

these dunes was probably deposited by Pleistocene 

periglacial winds in an environment 

that was characterized by Smith (1965) as 

first a desert (with large, transverse dunes 

built by northerly winds) and later as an 

area of sparse vegetation (with longitudinal 

dunes superimposed on the older dunes by 

northwesterly winds). The origin of the 

sand was probably the poorly consolidated 

Tertiary sediments of southwestern South 

Dakota and eastern Wyoming or from alluvium 

derived from these rocks about 10,000 

years ago (Watts and Wright 1966). Today, 

some dunes are as high as 400 feet and as 

long as 20 miles and have slopes as steep as 

25% (Bleed and Flowerday 1989). 

cl i m a t e 

The climate of the Great Plains is controlled 

largely by the rain shadow created 

by the Rocky Mountains as they intercept 

the easterly flow of moist Pacific air (Baker 

and Waln 1985). The plains are drier along 

the western edge near the rain shadow and 

become progressively moister to the east 

as air masses from the Gulf of Mexico play 

increasingly important roles in causing precipitation. 

Maximum precipitation in the 

Great Plains occurs in early summer (Fig. 4) 

due to the combined influences of moisture 

from the Gulf of Mexico and the penetration 

of modified Pacific air masses (Barry 1983). 

Fig. 4. Monthly precipitation for North Platte (Lincoln 

Co.) in the west and Lincoln (Lancaster Co.) in the east. 

A major characteristic of subhumid to 

semiarid climates (such as are found in Nebraska) 

is the high year to year variability in 

precipitation (Barry 1983). From ecological 

and historical perspectives, drought is possibly 

the most significant climatic element of 

the Great Plains environment because it determines 

the carrying capacity of the region. 

Nebraska has a continental, temperature 

zone climate characterized by hot summers, 

cold winters, moderate precipitation 

that is markedly seasonal or periodic, and 

occasional drastic changes in weather from 

day to day. It is often said in Nebraska, for 

example, that if you don’t like the weather, 

then wait an hour because it will change 

significantly. Rapid changes in weather are 

influenced by invasions of warm moist air 

from the Gulf of Mexico; hot, dry air from 

the Southwest; cool, usually dry air from the 

northern Pacific ocean; or cold, dry air from 

interior Canada. Masses of air over Nebraska 

are generally associated with the eastward 

movement of high and low pressure systems 

(Jones 1964). Precipitation and temperature 

are inversely related during the summer, 

and this relationship is less pronounced in

the spring and fall. Drought frequency is 

greater in the western part of the state, but 

there are more consecutive dry months in the 

east. Nebraska is subhumid in the east and 

gradually becomes semiarid in the west. 

The mean annual precipitation in the 

southeast is 33 inches (838 mm), 23 inches 

(584 mm) in the northeast, and decreases to 

15 inches (381 mm) in the west. Approximately 

three-fourths of this precipitation 

occurs between April and September and 

originates in warm, moist air from the Gulf 

of Mexico. However, much of the air moving 

north from the Gulf is deflected to the east 

by the eastward movement of air over the 

Rockies with the result that the easternmost 

part of the state receives more than 

twice as much precipitation as the drier 

westernmost part (Jones 1964). 

The mean annual temperature in 

Nebraska is 49.3°F (27.4°C). The mean 

annual summer temperature is 72.7°F 

(40.4°C). The mean annual winter temperature 

is 25.3°F (14.1°C). Winter temperatures 

in the northern Great Plains are 

low and often associated with strong winds 


Fig. 5. Nebraska climatological information. 

and occasional severe blizzards. The lowest 

temperature ever recorded was -47°F 

(-46.4°C) in February of 1899, and the 

highest temperature was 118°F (65.6°C) in 

July 1934. Figure 5 shows climatological 

information for the state. 

The prevailing winds are predominantly 

from the north and northwest in winter 

and from the south from May to November. 

Spring winds are usually the strongest and 

most variable in direction. It is of interest 

to note that greater climatic variation exists 

west to east across Nebraska (462 miles) 

than from eastern Nebraska to the Atlantic 

coast (approximately 1,110 miles). 

VeGeTATION OF NeBRASKA 

Although primarily a prairie state, 

Nebraska has many diverse habitats 

(Fig. 6) that range from eastern deciduous 

forests to short and tall grass 

prairies (actually six different grassland 

types), a large section of Sand Hills prairie 

(19,000 sq. mi.), and a small western 

component of Rocky Mountain forest.

8 


Weaver (1965) provided an excellent summary 

of the native vegetation of the state, 

and the distributions of all vascular plant species 

in the central grasslands were mapped in 

1977 (Great Plains Flora Association 1977). 

The vegetation in Nebraska has undergone 

considerable change since the pioneers 

first began settling here in the 1800s. Probably 

the three most significant changes are 

loss of many native prairies to agriculture, 

the introduction of trees in urban areas 

where once there were few, and the growth 

of woody vegetation in eastern Nebraska’s 

gullies and draws. These changes in the flora 

have affected the fauna because vegetation 

is a limiting factor for animals both as food 

and shelter. In some cases, floristic changes 

have been mirrored by the loss of animals 

to a particular habitat while in others it 

has resulted in a net gain in diversity. The 

interplay between plant and animal distribution 

is dynamic, and the human factor has 

substantially changed this relationship. 

The sense of living in a prairie environment 

or being in a prairie state has been 

largely lost because of the almost complete 

destruction of the original prairie by modern 

agriculture. This is especially true in 

eastern Nebraska where the once-dominant 

tallgrass prairie, stretching as far as one 

could see, has been eliminated by intense 

cultivation and urbanization. What follows 

is a very brief overview of Nebraska’s several 

floristic associations. 

Sh o r t a n d ta l l g r a s s Pr a i r i e 

The prairie is a land of waving grasses 

and broad-leaved forbs. Compositae and 

Leguminosae, along with the many grasses, 

are the dominant plant families found in 

the prairie. John Weaver, a noted Nebraska 

scholar of the prairie habitat, observed (1954) 

that the prairie appears almost monotonous 

in the general uniformity of its plant cover 

. . . but that it also has a special grandeur 

in its open expanses and in the abundance 

of its varicolored flowers. The dominance 

of perennial grasses, the paucity of shrubs, 

the absence of trees (except along rivers 

and streams), and a characteristic droughtenduring 

flora constitute its main features. 

Prairie is the name given to the vast expanse 

of grasslands in central North America. 

Similar grasslands in Eurasia are called 

“steppe” whereas in southern South America 

Fig. 6. Vegetation of Nebraska, circa 1850 (map modified from that by R. Kaul 1975).

they are referred to as “pampas,” and in 

southern Africa they are known as “veld.” 

The dominant grasses of North American 

prairies change along an east-west 

gradient as precipitation changes. This 

can be seen clearly in the vegetation map of 

Nebraska (Fig. 6). The eastern fourth of the 

state is (or was) tallgrass prairie, (Fig. 7), so 

called because the grasses there may reach 

a height of seven feet during moist years. 

The principal grasses are big bluestem 

(Andropogon gerardi Vitman), switchgrass 

(Panicum virgatum L.), and Indian grass 

(Sorghastrum avenaceum Michaux). The 

western border of tallgrass prairie approximates 

the line of 23 inches of annual precipitation 

(Jones 1964). 

Tallgrass prairie grades into mixed 

prairie (Fig. 8) where the dominant grasses 

are bluestems (Andropogon spp.), grama 

grasses (Bouteloua spp.), and buffalo grass 

(Buchloe dactyloides Nuttal). According 

to Weaver and Clements (1938), this association 

owes its name to the fact that its 

climax vegetation is composed of short and 

longer grasses in almost equal diversity. 


Jones (1964) observed that mixed prairie 

covers the largest area of any of the grassland 

associations in North America; it is 

the grassland of the Great Plains. Jones 

(citing Weaver and Clements) indicated 

the so-called shortgrass prairie of western 

Nebraska and adjacent regions was once 

considered a distinct grassland association, 

but was now known to represent only a “disclimax” 

of the mixed prairie that resulted 

primarily from overgrazing. There remains 

today no consensus of opinion as to whether 

this association is true shortgrass or “disclimax” 

mixed prairie. Based upon my own 

observations in western Nebraska, I prefer 

the designation of shortgrass prairie. 

With declining rainfall, changing soil 

profiles, and increasing evaporation in the 

western half of Nebraska, mixed prairie is 

replaced by shortgrass prairie, sand hills prairie, 

and sandsage prairie. The shortgrass prairie 

(Fig. 9) is dominated by blue grama grass 

(Bouteloua gracilis Humboldt, Bonpland, and 

Kunth) and buffalo grass (Buchleo dactyloides 

Nuttal). Shortgrass prairie is found in much 

of Nebraska’s western panhandle region. 

Fig. 7. Tallgrass prairie remnant at Roe Sanctuary near Gibbon in Buffalo Co. in 

eastern Nebraska. Photo courtesy of NEBRASKAland Magazine/Nebraska Game 

and Parks Commission.

10 


Fig. 8. Mixed prairie in Sarpy Co. in eastern Nebraska. Photo courtesy of 

NEBRASKAland Magazine/Nebraska Game and Parks Commission. 

Fig. 9. Shortgrass prairie at Jail and Courthouse Rocks in Morrill Co. in western 

Nebraska. Photo courtesy of NEBRASKAland Magazine/Nebraska Game and 

Parks Commission.


Fig. 10. Sandsage prairie near Parks in Dundy Co. in southwestern Nebraska. 

Photo courtesy of NEBRASKAland Magazine/Nebraska Game and Parks Commission. 

Fig. 11. Sand Hills prairie at “Arapaho Prairie” in Arthur Co. in western Nebraska. 

Photo by the author.

12 


Sandsage prairie (Fig. 10) is characterized by 

several species of sage (Artemisia spp.) as well 

as by sandreed grass (Calamovilfa longifolia 

Hooker) and bluestems (Andropogon spp.). 

This prairie type is found in the southwestern 

corner of Nebraska. 

Sa n d hi l l s Pr a i r i e 

The vegetation of the Sand Hills (Fig. 

11) is surprisingly diverse. It is also unique. 

. . not because it consists of many unusual 

species, but because it is a mixture of so 

many different types of vegetation. It is a 

“borrowed” vegetation in that most plants 

probably moved into the area from elsewhere 

during and after retreat of the glaciers (Kaul 

1989). Kaul noted that there is only one spe- 

cies of plant, Hayden’s, or blowout penstemon 

(Penstemon haydenii S. Watson), that 

is endemic to the Nebraska Sand Hills. It is 

the only endemic plant in Nebraska and one 

of only a few endemics in the Great Plains. 

Some of the most characteristic plants 

of the prairie region are bluestem grasses 

(Andropogon spp.), sandreed grass (Calamovilfa 

longifolia Hooker), needle grass (Stipa 

spp.), and yucca (Yucca spp.). The grass 

cover of the sand dunes comprising this area 

is fragile and susceptible to erosion. Excessive 

cultivation during the drought years 

of the 1930s caused erosion and some sand 

movement. Although the dunes are stabilized 

by plant cover today, local blowouts 

remain common. This region is one of the 

richest grazing areas in the United States. 

Fig. 12. Eastern deciduous forest adjacent to the Missouri River 

in Richardson Co. in southeastern Nebraska. Photo courtesy of 

NEBRASKAland Magazine/Nebraska Game and Parks Commission.

ea s t e r n de c i d u o u s Fo r e s t s 

The majority of Nebraska’s native trees 

entered the state (especially as they are 

represented in the Ohio Valley) following 

the Missouri River and its tributaries from 

the eastern forests (Pool 1929). 

Nebraska’s eastern deciduous, hardwood 

forests (Fig. 12) are largely restricted 

to the southeast corner of the state, the 

west bank of the Missouri River, and the 

Niobrara River in its eastern third. Within 

these riverine strips of forest, steep and 

undulating ridges contain dense upland 

forests dominated by the drought resistant 

bur oak (Quercus macrocarpa Michaux), 

shagbark hickory (Carya ovata [P. Miller]), 

and basswoods (Tilia americana L.). The 

deep ravines provide shelter from drying 

prairie winds and permit many species of 

broadleaf trees to survive. About 44 species 

of deciduous trees are native to southeastern 

Nebraska (Pool 1929). These hardwood 

forests extend west along rivers well out into 

the grasslands where they become impoverished 

in species (Kaul 1986). The eastern 


red cedar (Juniperus virginiana L.) is one 

of Nebraska’s four native coniferous trees. 

It is found widely scattered over the eastern 

half of the state on dry, gravelly slopes and 

limestone ridges (Pool 1929). 

Nebraska’s numerous floodplains that 

border rivers are usually at least partially 

covered by trees (Fig. 13) such as cottonwood 

(Populus deltoides Marshall), willow (Salix 

spp.), ash (Fraxinus pennsylvanica Marshall), 

elm (Ulmus americana L.), box elder 

(Acer negundo L.), and sycamore (Platanus 

occidentalis L.) (Kaul 1986). 

ro c k y mo u n t a i n Fo r e s t s 

Elements of the coniferous, evergreen 

forests of the Rocky Mountains (Fig. 14) are 

also found in Nebraska’s panhandle, primarily 

in the northwestern corner along the Pine 

Ridge escarpment. These forests extend 

eastward to approximately the 100th meridian 

on the north-facing slopes of the Niobrara 

River valley and its spring-fed tributaries. 

They meet the westward extensions of the 

eastern deciduous forests only in this region 

Fig. 13. Seasonally dry riverbed with gallery forest along the Platte River in 

Hall Co. in central Nebraska. Photo by the author.

14 


of north-central Nebraska. Ponderosa pine 

(Pinus ponderosa Lawson) and narrow-leaf 

cottonwood (Populus angustifolia James) 

are common Rocky Mountain trees that 

are found in this region of the state. Quaking 

aspen (Populus tremuloides Michaux) 

and western black birch (Betula fontinalis 

Sargent) also indicate the montane floral 

affinities of the Pine Ridge. These species 

probably occurred widely over much of the 

western part of the state in post-Wisconsin 

times, and the areas that remained when 

Europeans first reached western Nebraska 

were relics of this former widespread distribution 

(Jones 1964). The large influx of 

settlers since that time has altered considerably 

those remaining forest relics which, 

today, are disturbed remnants of the former 

plant associations. 

Fig. 14. Rocky Mountain forest near Chadron in 

Dawes Co. in northwestern Nebraska. Photo courtesy 

of NEBRASKAland Magazine/Nebraska Game and 

Parks Commission. 

The SIlPhIDAe 

The spectacle of nature is always new, 

for she is always renewing the spectors. Life 

is her most exquisite invention, and death 

is her expert contrivance to get plenty of it. 

— Goethe 

Carrion beetles is the term applied, in 

a strict sense, to a single family of beetles, 

the Silphidae. Silphids are also generally 

referred to as burying beetles or sexton 

beetles because of the behavioral adaptations 

of Nicrophorus species to inter small 

vertebrates in the ground. 

Silphids are relatively large beetles, 

ranging in size from 10 to 35 mm. The majority 

are usually a dull black or grey in color, 

but most species in the genus Nicrophorus 

have bright orange markings on the elytra 

that may serve as warning coloration. 

Most silphids occur in north temperate 

regions, which is where they probably originated 

(Peck and Anderson 1985, in part). 

The majority of silphids are scavengers on 

carrion, and a few are found on dung or fungi, 

are phytophagous, or prey on fly larvae. 

Carrion beetles are a conspicuous 

element of that vast host of scavengers 

that are responsible for breaking down 

and recycling back into the ecosystem the 

basic elements found inside of each organism. 

The decay process is an efficient and 

natural system whereby the raw materials 

of dead organisms are returned directly 

into the energy budgets of living organisms 

when they consume the dead ones. 

Once an animal dies, its remains are 

ravenously set upon by a diverse array of 

food-seeking scavengers and predators that 

are attracted by the odors of decay. Assuming 

that vertebrate scavengers do not find 

and consume the remains first while they 

are still fresh, the remains will become a 

valuable food resource for a reasonably orderly 

progression of bacteria and fungi (the 

microconsumers) and insects (the macroconsumers) 

(Ratcliffe 1980). The progression of 

insects is fairly predictable because specific

insects are attracted to a cadaver only after 

certain levels of decay have occurred. These 

stages of decay (and associated fauna) are 

influenced by season, weather, and the size 

and exposure of the remains. The net effect 

of this food partitioning is to reduce competition 

among the different guilds of insect scavengers 

by spacing them out through time and 

enabling increased use of a patchy, limited 

resource by more organisms. Arthropod succession 

at carrion has been thoroughly studied 

by Fuller (1934), Bornemissza (1957), 

Reed (1958), Payne (1965), and Early and 

Goff (1986), among others. 

Many silphids are active at night, which 

may be a strategy to reduce competition from 

flies that are primarily diurnal. If flies do 

manage to lay eggs on a carcass, that carcass 

can become unfit for use by Nicrophorus species 

because the fly larvae consume nearly all 

of the fleshy remains that would otherwise 

be used by the beetles. 

Nicrophorus species are renown for 

their habit of burying small vertebrate carcasses 

beneath the surface of the soil. Usually 

a male/female pair will process these 

remains to provision their developing larvae. 

The burial of the food source is important 

to these beetles and their young because it 

effectively removes the food from the arena 

of intense competition by maggots and other 

carrion-feeding insects. Nicrophorus species 

are unique among silphids because they are 

the only ones attempting to break the cycle 

of competition at a food source. At the same 

time, they provide their larvae with a safer 

underground environment that is relatively 

free from predators in which to develop. 

Species in the Silphinae do not inter 

remains like Nicrophorus species. Instead, 

adults arrive at a carcass during the early to 

middle stages of decay (Payne 1965, Johnson 

1974). Most seem to lay eggs just beneath 

the surface of the soil near the carcass, and 

the eggs hatch after four or five days (Anderson 

1982c). The larvae then feed on the 

remains at the same time as all the other 

carrion-frequenting insects. The larvae pass 

through three instars, and they pupate in 


earthen cells beneath the soil. Details of the 

life history for most of these species remain 

poorly known. Young (1983) assembled an 

extensive bibliography on the biology of the 

Silphidae. 

According to Lawrence and Newton 

(1982, 1995), the Silphidae, a once vaguely 

defined group, is now restricted to the larger 

carrion and burying beetles. The family 

is clearly monophyletic and related to the 

Staphylinidae. The Agyrtidae were formerly 

included in the Silphidae (e.g., Arnett 1968, 

Madge 1980, Cho and Lee 1986) as were 

parts of the Leiodidae (e.g., Hatch 1928), but 

these are now considered to be valid families 

unto themselves. 

The Silphidae, then, consists of two 

subfamilies: the Silphinae and Nicrophorinae. 

Between them, there are 13 genera 

with about 208 species worldwide. North 

America has eight genera and 30 species, 

and in Nebraska there are six genera and 

18 species. 

Early synoptic treatments of the North 

American silphids were provided by LeConte 

(1853) and Horn (1880), both of whom recognized 

only the genera Silpha and Nicrophorus. 

Portevin (1926) split Silpha into many of 

the genera that we use today although there 

was not wide acceptance of these genera until 

the works of Miller and Peck (1979) and 

Anderson and Peck (1985) appeared. 

Hatch (1927) and Arnett (1944) compiled 

relatively comprehensive works for the 

U.S. fauna, and they were among the first to 

actually use some character analysis. Portevin 

(1926) monographed the world fauna, 

and Hatch (1928) provided a checklist of the 

world fauna in the Coleopterorum Catalogus 

series. Peck and Anderson (1985) reviewed 

the taxonomy, phylogeny, and biogeography 

of the silphids of Latin America. 

A preliminary checklist of the silphids 

of Nebraska was published by Meserve in 

1936, and he recognized 15 species for the 

state. The silphids of other states have been 

listed or treated taxonomically by Fall and 

Cockerell (1907) for New Mexico, Blatchley 

(1910) for Indiana, Hatch and Rueter (1934)

16 


for Washington, Hatch (1957) for the Pacific 

Northwest, Lago and Miller (1983) for 

Mississippi, Lingafelter (1995) for Kansas, 

and Cuthrell and Rider (in press) for the 

Dakotas. Checklists of the North American 

fauna were prepared by Leng (1920), 

Blackwelder and Arnett (1974), and Peck 

and Miller (1993). 

COlleCTING SIlPhIDS 

Carrion beetles are relatively easy to 

collect because they are so readily attracted 

to carcasses or bait that can be easily 

manipulated. The most obvious way of 

col-lecting these beetles is to find them at 

naturally-occurring carrion. In the earlier 

stages of the decay process, Nicrophorus 

species can be found beneath carcasses 

either feeding or preparing to bury the 

remains. If the carcass is slowly rolled 

over, Nicrophorus species can be collected 

with fingers or forceps as they run for 

cover. Species of silphines can be collected 

in much the same way from carcasses that 

are slightly older in the decay cycle. 

Carcasses can also be deliberately 

placed in selected areas in order to attract or 

survey for silphids. Whole animal carcasses 

can often be obtained from pig, chicken, or 

turkey farms where there is always juvenile 

mortality. Frozen laboratory rats are also a 

more expensive option. These remains can 

be periodically checked for silphids in the 

same way as naturally-occurring carrion. 

With smaller carcasses that are likely to 

be buried by Nicrophorus species, an 18inch 

length of dental floss can be tied to a 

rear leg. The dental floss remaining above 

the ground serves as a marker that can be 

followed down to the buried remains after 

several days. In this way, both the adults 

and larvae can be collected. 

One of the most popular methods of collecting 

is the use of baited pitfall traps. In 

this method, a wide-mouth jar, can, bottle, 

or plastic bucket containing rotting meat 

as a bait is placed into the ground with the 

lip of the container level with the surface of 

the soil. Soil should be packed around the 

opening of the trap to enable an unobstructed 

approach by beetles walking to the trap. 

Silphids are attracted to the odor of the bait 

and fall into the container where they may 

be either live-trapped in a sand substrate 

or killed in a preservative solution such as 

ethylene glycol (diluted automobile antifreeze) 

or soapy water. There are probably 

as many trap designs as there are people to 

invent them, but some do seem to be more 

successful than others in trapping beetles. 

Figure 15 shows a generalized schematic for 

a baited pitfall trap. The experience of the 

people working in my lab seems to indicate 

that traps with larger surface areas tend to 

have more beetles in them when the amount 

of bait remains the same. Similarly, traps 

with more bait attract more beetles even 

when the trap container size varies. Whole 

animal remains seem to attract more carrion 

beetles than when only parts (e.g., beef 

liver, pieces of fish, or chicken gizzards) are 

used. As always, proper aging of bait (two to 

three days in warm weather) helps to ensure 

greater trapping success. Ripening of bait in 

a closed container will keep flies from ovipositing 

on the bait, thus making it unusable. 

Fig. 15. Graphic representation of a baited pitfall trap.

The jar or container should not be tightly 

sealed in order to allow for the escape of gases 

that will be generated; otherwise, a nasty 

surprise could greet anyone opening the jar 

for the first time as the rotted bait explodes 

from the container. Ripened bait can then be 

suspended in one of several ways in the trap 

(as in Fig. 15) or placed in a small receptacle 

on the sandy bottom of a live trap (such as a 

small jar inside of a five-gallon bucket). Bait 

should always be wrapped or containerized 

to prevent beetles from actually coming into 

contact with it. 

Placement of traps is also important. 

Traps placed in dense woods attract fewer 

beetles than those placed in more open 

woods, meadows, or prairies. This is probably 

because flying through dense undergrowth 

while searching for carrion is more 

difficult. Traps exposed to direct sun may 

attract fewer beetles if the bait inside the 

trap becomes dried out and mummified. A 

rain cover/sun shield should always be placed 

over pitfall traps so as to protect both the 

bait and trapped beetles from the elements. 

Traps should generally not be placed in areas 

so low that they might flood if it rains heavily. 

Traps should also not be placed near an 

ant nest because the ants may usurp the bait 

and deter beetles from approaching. 

Vertebrate scavengers will often attempt 

to get at the bait in traps, thus disrupting 

any trapping program. Raccoons 

especially are tenacious in their efforts to 

get at rotting bait. A one-foot-square piece 

of one-inch mesh screen wire will prevent 

such scavenging if it is securely staked down 

above the opening of the pitfall trap. Even 

so, I have had very persistent scavengers 

dig entire traps out of the ground in order to 

eat the bait. Shubeck (1984b) designed an 

inexpensive carrion beetle trap that might 

inhibit such scavenging although it is more 

labor intensive for the initial construction and 

more cumbersome to transport, especially if 

a large number of them are being used. 

When live-trapping (as for the endangered 

American burying beetle or to acquire 

other species alive for behavioral studies) 


great care must be taken to prevent beetle 

mortality in the trap resulting from flooding, 

overcrowding, or from heat stress. For nocturnal 

species, traps should be serviced daily and 

preferably in the morning before daytime high 

temperatures cause traps to become lethal. 

Pitfall traps should always be completely 

and securely covered or removed when they 

are no longer in use to prevent additional, 

unnecessary mortality to non-target beetles 

because additional beetles will fall into them, 

die, rot, and then attract more beetles. 

Key TO The SUBFAMIlIeS 

OF ADUlT SIlPhIDAe IN 

NeBRASKA 

1. Elytra with apices rounded, not truncate 

or shortened, usually exposing 1-2 

segments (Thanotophilus truncatus has 

truncate elytra, but it is entirely black and 

has a gradually widening antennal club). Antenna 

with 11 distinct segments, gradually 

widening into club . . . . . . . . . . . Silphinae 

1’. Elytra with apices truncate, shortened, 

usually exposing 3-4 segments.Antenna 

apparently 10-segmented (actually 11-segmented 

but second small, nearly hidden in 

apex of first), last 4 segments widened into 

distinct, compact club . . . Nicrophorinae 

Key TO The SUBFAMIlIeS OF 

ThIRD INSTAR lARVAe OF 

SIlPhIDAe IN NeBRASKA 

(after Anderson and Peck 1985) 

1. Tergites large, laterally produced, each 

usually with posterior angles attenuated 

(Fig. 16). Ventral surface with sternites large, 

sclerotized, and pigmented. Head on each side 

with 6 pigmented stemmata . . . . Silphinae 

1’. Tergites small, those on abdomen each 

with 4 small spines (Fig. 17). Ventral surface 

soft, creamy white, lacking sclerotization. 

Head on each side with 1 unpigmented 

stemma . . . . . . . . . . . . . . Nicrophorinae

18 


Fi g s. 16-17. Body form of 16, a larval silphine, Necrophila americana and 17, a larval 

nicrophorine, Nicrophorus investigator.

SUBFAMIly SIlPhINAe 

The Silphinae is comprised of ten genera 

containing about 120 species (Hatch 

1928, Lawrence and Newton 1995). They 

are worldwide in distribution with most 

of the species occurring in Eurasia and 

North America although there are two 

genera (Diamesus and Ptomaphila) with 

five species that reach Australia. In North 

America there are seven genera represented, 

and five of these genera (representing 

seven species) occur in Nebraska. 

As with the Nicrophorinae, the status 

of the category Silphinae has shifted 

back and forth from tribe to subfamily 

depending on which specialist’s views 

held sway at the time. The taxonomic 

category seems finally to have settled at 

the subfamily level following the works 

of Anderson and Peck (1985), Peck and 

Anderson (1985), and Lawrence and 

Newton (1995). 


Key TO The GeNeRA AND SPe- 

CIeS OF ADUlT SIlPhINAe IN 

NeBRASKA 

(modified from Anderson and Peck 1985) 

1. Pronotum black with yellow margins 

(Fig. 57). Elytra with reticulate sculpturing 

. . . . . . . . . . . . . . . . . . . . . . . . . Necrophila 

1’. Pronotum entirely black or black with 

reddish orange margins. Elytra with or without 

reticulate sculpturing . . . . . . . . . . . . 2 

2. Eyes large (Fig. 49). Pronotum distinctly 

orbicular, widest at middle, basal angle 

broadly rounded. Male with metafemora 

enlarged and with tooth near apex (Fig. 18) 

. . . . . . . . . . . . . . . . . . . . . . . . . . Necrodes 

2’. Eyes small. Pronotum not orbicular, 

widest toward base, basal angles subangulate. 

Males with metafemora not enlarged 

or toothed . . . . . . . . . . . . . . . . . . . . . . . . 3 

3. Head with distinct row of stout setae 

behind eyes (best seen in anterior view) 

(Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . 4 

Fi g s. 18-19. Ventral view of left posterior femur and tibia of (18) male and (19) female 

Necrodes surinamensis. Fig. 20. Anterior view of head of Oiceoptoma inaequale showing 

row of stout setae behind eye.

20 


Fi g s. 21-33. Left elytron of 21-22, Heterosilpha ramosa, male and female; 23-24, Necrodes surinamensis, male and 

female; 25-26, Necrophila americana, male and female; 27-28, Oiceoptoma inaequale, male and female; 29-30, O. 

novaboracense, male and female; 31-32, Thanatophilus lapponicus, male and female; 33, T. truncatus (either sex).

3’. Head may have short setae, but not with 

stout setae in a distinct row behind eye . . . 5 

4. Humerus with small tooth (Figs. 27-30) 

or acute angle. Elytra without reticulate 

sculpturing . . . . . . . . . . . . . . . . Oiceoptoma 

A. Pronotum entirely black . . . . . . . . 

. . . . . . . . . . . . . . . O. inaequale (Fabr.) 

A’. Pronotum black with reddish orange 

margins . . . . O. novaboracense (Forster) 

4’. Humerus rounded, lacking tooth. Elytra 

with reticulate sculpturing . . Heterosilpha 

5. Humerus with small tooth (Figs. 31-33). 

Labrum shallowly emarginate. Mesocoxae 

widely separated (by about width of mesocoxa) 

. . . . . . . . . . . . . . . . Thanatophilus 

A. Elytra smooth, lacking costae 

(Figs 33, 64) . . . . T. truncatus (Say) 

A’. Elytra tuberculate, with costae (Figs. 

31, 32, 62) . . . . T. lapponicus (herbst) 

5’. Humerus rounded. Labrum deeply 

emarginate. Mesocoxae narrowly separated 

(by about half or less width of mesocoxa) . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Aclypea 


Key TO The GeNeRA AND SPe- 

CIeS OF ThIRD INSTAR lARVAe 

OF SIlPhINAe IN NeBRASKA 


1. Urogomphi distinctly longer than 10th 

abdominal segment (by at least half their 

length) (Figs. 34-35). Sternum of 2nd abdominal 

segment with 3 large sclerites . . 2 

1’. Urogomphi equal to or slightly longer 

than 10th abdominal segment (Figs. 36-39). 

Sternum of 2nd abdominal segment with 1 

large sclerite . . . . . . . . . . . . . . . . . . . . . . . 3 

2. Basal segment of urogomphus up to 

twice as long as 10th abdominal segment 

(Fig. 35). Dorsal color dark brown to black . . 

. . . . . . . . . . . . . . . . . . . . . Thanatophilus 

2’. Basal segment of urogomphus more 

than twice as long as 10th abdominal segment 

(Fig. 34). Dorsal color reddish brown 

. . . . . . . Necrodes surinamensis (Fabr.) 

3. Second segment of antenna with 1 large 

sensory cone (Fig. 40). Prothoracic tergite 

Fi g s. 34-39. Abdominal apex of larval silphine (dorsal view): 34, Necrodes surinamensis; 35, Thanatophilus truncatus; 

36, Necrophila americana; 37, Oiceoptoma novaboracense; 38, Aclypea bituberosa; 39, Heterosilpha ramosa (Figs. 

38-39 after Anderson and Peck 1985).

22 


emarginate anteriorly at middle (Figs. 43-44). 

Dorsal color reddish brown . . . Oiceoptoma 

A. Prothoracic tergite deeply emargiate 

anteriorly (Fig. 43). Thoracic and ab- 

dominal tergites 1-8 with lateral margins 

pale, pale areas with small, dark, spots 

or oblique lines (Fig. 43) . . . . . . . . . 

. . . . . . O. novaboracense (Forster) 

A’. Prothoracic tergite shallowly emarginate 

anteriorly (Fig. 44). Meso- and 

metathoracic tergites as well as abdominal 

tergites 1-8 with pale areas limited 

to posterolateral angles (Fig. 44) . . . . 

. . . . . . . . . . . . O. inaequale (Fabr.) 

3’. Second segment of antenna with 1 or 

more plates on sensory area (Figs. 41-42). 

Prothoracic tergite not emarginate anteriorly 

(Fig. 45). Dorsal color dark brown . . . . . 4 

4. Second and third segments of antenna 

subequal in length. Urogomphi distinctly 

2-segmented (Fig. 36) . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . Necrophila americana (l.) 

4’. Third segment of antenna distinctly 

longer than second segment. Urogomphi 

apparently with 1 segment (Figs. 38-39) . . 5 

5. Second segment of antenna with 1 plate 

on sensory area (Fig. 41). Last segment of 

maxillary palpus about twice as long as 

wide . . . . Aclypea bituberosa (leConte) 

5’. Second segment of antenna with numerous 

plates (Fig. 42). Last segment of 

maxillary palpus about 3 times as long as 

wide . . . . . Heterosilpha ramosa (Say) 

Fi g s. 40-42. Second antennal segment of larval Silphinae: 

40, Oiceoptoma novaboracense; 41, Aclypea bituberosa; 

42, Necrophila americana or Heterosilpha ramosa 

(Fig. 41 after Anderson and Peck 1985). 

Fi g s. 43-45. Prothoracic and abdominal tergites of larvae 

of 43, Oiceoptoma novaboracense; 44, O. inaequale; 

45, Necrophila americana. 

Genus AclypeA 

The genus Aclypea has about 22 species 

(Hatch 1928). They are Holarctic in 

distribution. Only two species are found 

in the United States. 

Aclypea bituberosa (LeConte) (Fig. 

46) is included in this work because there 

are some old references indicating its 

presence in Nebraska. This species does 

not occur in Nebraska. I believe that the 

earlier citations of its occurrence here were 

a result of misidentification where Silpha 

opaca (later thought to be bituberosa) was 

confused with Oiceoptoma inaequale or 

Heterosilpha ramosa.

In a report by Lawrence Bruner on 

the “Insect injuries in Nebraska during 

the summer of 1892,” Bruner discusses 

second hand information from a Mr. Huxman 

that “Silpha opaca” was a pest of 

sugar beets at West Point. Huxman had 

told Bruner that he could not be mistaken 

about the identity of the insect because he 

had seen so many of them in Europe that 

he knew them on sight. I believe that 

Huxman had the very similar appearing 

Oiceoptoma inaequale or Heterosilpha 

ramosa (which does not occur in Europe) 

and not Aclypea bituberosa. In a 1916 

letter from Myron Swenk (entomologist 


Fig. 46. Aclypea bituberosa (LeConte). 

at the Nebraska Experiment Station) 

to R. Cooley, it is again mentioned that 

“Silpha” bituberosa was found in beet 

fields in Nebraska . . . but, it is important 

to note, Swenk obtained this information 

from Bruner (who, by extension, received 

it from Huxman). There are no actual 

specimens to support these claims. Cooley 

(1917), in a detailed report on the biology, 

distribution, and description of Aclypea 

(his Silpha) bituberosa, then erroneously 

concluded that there is “positive information 

of its occurrence in Nebraska.” 

Meserve (1936), in his list of Nebraska 

silphids, also mentioned a single, no-data

24 


record of Silpha opaca, but, again, there 

is no voucher specimen. Anderson and Peck 

(1985) and Peck and Kaulbars (1987), using 

the information in Cooley (1917), included 

Nebraska in the distribution of this species. 

This species occurs just west of the Rocky 

Mountains in Wyoming, Montana, Utah, 

and Idaho. 

According to Anderson and Peck 

(1984), both adults and larvae of Aclypea 

bituberosa are phytophagous and eat the 

leaves and shoots of native species of Solanaceae 

and Chenopodiaceae, introduced 

weeds, and at least 12 species of plants of 

agricultural or horticultural importance, including 

squash, pumpkin, spinach, wheat, 

beet, radish, rhubarb, potato, lettuce, cabbage, 

rapeseed, and turnip. Occasionally, 

this species has been considered a pest of 

some of these crops. 

Genus HeTeROSIlpHA 

Heterosilpha Portevin 1926: 83. 

The genus Heterosilpha consists of 

two species that are both endemic to North 

America. Heterosilpha aenescens (Casey) 

occurs near the west coast from southern 

Oregon to northern Baja California in 

Mexico (Miller and Peck 1979, Peck and 

Anderson 1985) and H. ramosa (Say) is 

found in most of western North America 

from southern Canada to northern Mexico 

(Peck and Anderson 1985). Only the latter 

species is found in Nebraska. 

Prior to Portevin’s (1926) establishment 

of the genus Heterosilpha, these species 

were included in Silpha, a catch-all 

genus at the time. Arnett (1946) suggested 

that aenescens was a synonym of ramosa, 

but this has not been adopted by subsequent 

authors. 

Species of Heterosilpha are unique 

among North American silphids because of 

the presence of three, branching costae on 

each elytron. Larvae are characterized by 

having the sensory area of the second antennal 

segment with numerous small plates 

(as in Fig. 42) and the urogomphi subequal 

to the tenth abdominal segment. 

Heterosilpha ramosa (Say) 

(Figs. 21-22, 39, 47-48) 

Silpha ramosa Say 1823: 193. 

Silpha cervaria Mannerheim 1843: 252. 

Diagnosis. Length 11.2-16.7 mm. Thorax: 

Color entirely black. Surface finely, densely 

punctate. Elytra: Color black. Surface tricostate, 

costae with short, lateral branches. 

Males with apex rounder, females with apex 

slightly elongated and attenuated (Figs. 21- 

22). Legs: Males with tarsomeres 1-4 on first 

and second pair of legs laterally expanded 

and densely pubescent beneath; females 

with tarsomeres normal, not expanded. 

Distribution. Heterosilpha ramosa is 

found west of a line from northeastern 

Minnesota to south-central New Mexico; 

it also occurs in southern Canada west of 

Lake Superior to British Columbia as well 

as in northern Baja, California (Anderson 

and Peck 1985, Peck and Kaulbars 1987). 

In Nebraska, this species is more abundant 

in the western half of the state with populations 

extending eastward as far as Grand 

Island. There are two old Lincoln records 

(pre-1920), but this species does not now 

occur in Lancaster Co. 

locality Records (Fig. 48). 78 Nebraska 

specimens examined or recorded. 

CHERRY CO. (43): Hackberry Lake, Trout 

Lake, Valentine, Valentine Wildlife Refuge; 

CUSTER CO. (2): Anselmo, 17 mi. E. 

Anselmo; DAWES CO. (1): Pepper Creek; 

GARDEN CO. (4): Crescent Lake, Oshkosh; 

HALL CO. (2): Alda, Mormon Island Refuge; 

KEITH CO. (15): Cedar Point Biological Station; 

KEYA PAHA CO. (1): Mills; LINCOLN 

CO. (8): Box Elder Canyon, Moran Canyon, 

North Platte, Wellfleet; LOGAN CO. (1): No 

data; McPHERSON CO. (1): Sandhills Ag 

Lab; MORRILL CO. (3): No data; SCOTTS 

BLUFF CO. (1): Mitchell.


Fig. 47. Heterosilpha ramosa (Say). 

Fig. 48. Nebraska distribution of Heterosilpha ramosa.

26 


Temporal Distribution. Rangewide: 

March to October (Peck and Kaulbars 1987). 

Nebraska: May (2), June (37), July (34), 

August (10), October (3). 

Remarks. Heterosilpha ramosa is easily 

recognized because it is the only silphid in 

North America that is entirely black, with 

distinctly tricostate elytra, and with the 

costae branching and weakly shining against 

a dull black background. 

The larval stage was described by 

Gissler (1880), Dorsey (1940), Brewer and Bacon 

(1975), and Anderson and Peck (1985). 

Brewer and Bacon (1975) studied the 

natural history of this species in Colorado, 

and their observations are probably representative 

for Nebraska as well. Adults 

overwinter and become active the following 

spring when temperatures become warm. 

Eggs are laid in the soil around a carcass, 

and this stage typically lasts 5 days. The 

first instar takes 4-5 days, the second instar 

5-6 days, the third instar 8-10 days, and the 

pupal stage 8-9 days. The period from egg 

to adult lasted about 30 days. There are two 

generations a year with adults of the first 

brood (in Nebraska) appearing in June and 

those of the second in late July and August. 

Genus NecRODeS 

Necrodes Leach 1815: 88. 

Asbolus Bergroth 1884: 229. 

Protonecrodes Portevin 1922: 508. 

The genus Necrodes contains five 

species (Hatch 1928) distributed in North 

America, Europe, and Asia. There is only 

one species in North America, and it is found 

throughout Nebraska. There is no modern 

taxonomic treatment of the genus, and the 

most recent world catalog is Hatch (1928). 

Madge (1980) reported that Bergroth 

(1884) believed that the name Asbolus was 

validly published by Voet (1778) and proposed 

that it replace Necrodes Leach 1815. 

However, Voet’s work was not consistently 

binomial and thus is not available for zoo- 

logical nomenclature according to the Code. 

Asbolus, therefore, dates from Bergroth 

(1884) whose action can be regarded as the 

proposal of an unnecessary nomen novum. 

While life history information is probably 

generally known for all of the species, 

only the North American N. surinamensis 

has been studied in detail (Ratcliffe 1972). 

The genus Necrodes is easily recognized 

because of its large eyes, broadly orbicular 

pronotum, strongly tricostate elytra, and 

males with enlarged posterior femora. The 

larvae are distinctive because the basal segment 

of the urogomphus is more than twice 

as long as the tenth abdominal segment. 

Necrodes surinamensis (Fabr.) 

(Figs. 18-19, 23-24, 34, 49-56) 

Silpha surinamensis Fabricius 1775: 72. 

Protonecrodes surinamensis bizonatus Portevin 

1926: 165. 

Diagnosis. Length 12.0-24.0 mm. Head: 

Color black, widest across large eyes. Antenna 

11-segmented, gradually clavate. 

Labrum broadly, shallowly emarginate. 

Thorax: Pronotum shining black, orbicular, 

widest near middle. Surface densely punctate, 

punctures small. Elytra: Color black, 

usually with subapical, transverse row of 

1-5, small, reddish orange spots variously 

combined; occasionally with subbasal, transverse 

row of 1-3 spots; rarely immaculate. 

Surface strongly tricostate, with short costa 

at base between costae 2-3. Surface densely 

punctate, punctures moderately large. Legs: 

Foretarsi of males with segments 1-4 usually 

expanded, as wide as long; in females, segments 

a little longer than wide. Males with 

hind femora usually enlarged and with acute 

tooth on posterior edge; femora not enlarged 

or toothed in females (Figs. 18-19). Posterior 

tibia usually curved in males, straight in 

females. 

Distribution. Necrodes surinamensis is 

broadly distributed in the eastern United 

States east of the Rocky Mountains and in

certain areas of the Pacific Northwest, Montana, 

and Utah; it is also found in southern 

Canada from Newfoundland to British 

Columbia (Ratcliffe 1972). This species is 

distributed throughout Nebraska and seems 

to show a preference for wooded areas. 

locality Records (Fig. 50). 2,479 Nebraska 


ADAMS CO. (10): No data; BOYD CO. (9): 

Spencer; BUFFALO CO. (2): Kearney; CASS 

CO. (7): Plattsmouth, South Bend, Union; 

CHASE CO. (1): Enders Reservoir; CHERRY 

CO. (5): Ft. Niobrara National Wildlife Refuge; 

CLAY CO. (1): No data; DAWSON CO. 

(350): No data; DOUGLAS CO. (6): Omaha; 

DUNDY CO. (1): S side Republican River E of 

Benkelman; FILLMORE CO. (5): Fairmont; 


Fig. 49. Necrodes surinamensis (Fabr.), male. 

FRANKLIN CO. (1): No data; FRONTIER 

CO. (46): Farnam, Medicine Creek Reservoir, 

Red Willow Reservoir; GAGE CO. (3): 

Beatrice; GOSPER CO. (338): Elwood Reservoir, 

Lexington, Smithfield; HALL CO. (21): 

Alda; HOLT CO. (1): O’Neill; JEFFERSON 

CO. (14): Fairbury; JOHNSON CO. (11): 

No data; KEYA PAHA CO. (16): Mills, Norden; 

KNOX CO. (18): Bazile Creek Wildlife 

Management Area; LANCASTER CO. (40): 

Hickman, Lincoln, Reller Prairie; LINCOLN 

CO. (49): Box Elder Canyon, Brady, Cottonwood 

Canyon, Moran Canyon, North Platte, 

Wellfleet; OTOE CO. (1,048): Nebraska City; 

PAWNEE CO. (11): No data; PHELPS CO. 

(28): Bertrand; PLATTE CO. (2): Columbus; 

RED WILLOW CO. (3): McCook; RICH- 

ARDSON CO. (2): Indian Cave State Park;

28 


SALINE CO. (1): Dewitt; SARPY CO. (5): 

Bellevue, Fontenelle Forest; SAUNDERS CO. 

(424): Ashland. 


January through December (Ratcliffe 1972). 

Nebraska: April (8), May (5), June (54), July 

(269), August (2,016), September (12), October 

(10), November (3). The large number 

for August is a result of a survey in Otoe 

County in 1995. 

Fig. 50. Nebraska distribution of Necrodes surinamensis. 

Remarks. Necrodes surinamensis is distinguished 

from other members of the Silphinae 

by its large eyes, completely black, orbicular 

pronotum, strongly tricostate elytra, and 

enlarged femora in the males. The variation 

in elytral spots is considerable and was 

documented by Ratcliffe (1972). 

The egg, larval, and pupal stages were 

described in detail by Ratcliffe (1972). 

The life history of this species was 

studied in detail by Ratcliffe (1972), and a 

brief synopsis is provided here. After locating 

carrion, adults of Necrodes surinamensis 

feed actively on the dipterous larvae that are 

present. They mate during the one to seven 

days they are at the carcass. The females 

oviposit relatively large, cream colored eggs 

one at a time randomly on the soil near the 

carcass. The eggs (Figs. 51-52) gradually 

darken to resemble the soil on which they 

rest. Larvae of different instars are found 

on the same carcass because eggs are laid 

over a span of several days. Larvae normally 

hatch from the eggs in 2-4 days and 

immediately seek the shelter of the carcass 

to begin feeding. Under favorable conditions, 

first instar larvae (Fig. 53) molt in 1-2 days. 

Second instar larvae molt after 2-5 days. The 

third instar stage (Fig. 54) lasts 3-5 days 

and, as in all previous stages, may be greatly 

extended due to cold or wet weather or poor 

food supply. When ready to pupate the larvae 

wander a short distance from the carcass 

to form earthen pupal cells in the ground. 

Pupal cells made in laboratory rearing 

chambers were constructed 5 cm below the 

surface. The pupal cell is formed by sharp, 

convulsive thrashing of the abdomen that 

gradually forms an oval, hollow chamber 

with firmly packed walls (Fig. 55). After 

construction of the pupal cell, a period of quiescence 

follows lasting from 5-8 days. This 

is the pharate pupal stage, so called because 

the new pupa is developing inside the cuticle 

of the last instar. Ecdysis eventually occurs


Fi g s. 51-52. Eggs of Necrodes surinamensis: 51, eggs 2 hours old; 52, eggs one day old. Egg on right has 

turned a cryptic brown. Figs. 53-54. First and third instar, respectively, of N. surinamensis. Fig. 55. Ventral 

aspect of pharate pupa of N. surinamensis within its pupal chamber. Fig. 56. Final molt to pupal stage of 

N. surinamensis.

30 


and exposes the cream-white pupa (Fig. 56). 

The length of the pupal period varies from 12- 

17 days. After emerging from the pupa, the 

adult becomes sclerotized in about 24 hours, 

after which it digs its way out of the subterranean 

cell. Adults overwinter beneath the 

soil or in areas that afford protection. 

Adults feed primarily on fly larvae during 

the active decay stage of a carcass when 

maggots are usually abundant, but they will 

also consume carrion as well. Typically, an 

adult N. surinamensis seizes a maggot with 

its mandibles, restrains it with its forelegs 

placed on either side of the maggot, and 

raises its head to lift the struggling maggot 

off of the substrate and so prevent it from 

pulling away. After easily breaking the integument 

of the maggot with its mandibles, 

the beetle chews rapidly and extracts the soft 

body contents. Young (1985) observed that 

adults may, on occasion, consume noctuid 

moth larvae and dead insects. 

While the adults are predators on maggots 

and may feed on carrion to a limited 

extent, the larvae normally feed on carrion 

upon which they are free living and, in addition, 

feed on fly larvae to a limited extent. 

During active decay of a carcass, larvae 

feed on decomposing flesh and semi-liquid 

putrefaction. When maggot feeding, a larvae 

seizes a maggot in the mid-region of the body 

with the mandibles while the forelegs assist 

in immobilizing the prey. After puncturing 

the cuticle, the soft contents are consumed. 

Necrodes surinamensis is unique among 

silphids because it can eject anal fluid as 

a spray (rather than an ooze) (Reed 1958, 

Ratcliffe 1972, Eisner and Meinwald 1982). 

The abdominal tip, which projects beyond 

the posterior margins of the elytra, serves 

as a revolvable turret by which ejections are 

actually aimed (personal observation, Eisner 

and Meinwald 1982). Eisner and Meinwald 

noted that N. surinamensis is anomalous in 

that it expels its aimed, secretory discharges 

from the anus (admixed with enteric matter). 

Other beetles that spray (i.e., Carabidae) 

also discharge from the tip of the abdomen, 

but the glands responsible for the spray are 

integumental and open beside the anus. According 

to Schildknecht and Weis (1962), the 

high concentration of ammonia in the spray 

is probably derived from decaying, ingested 

animal protein and may serve for defense. 

Adults are nocturnal and are strongly 

attracted to lights at night. And it is for this 

reason that I have concerns for the future 

welfare of this species. During the mid- 

1970s when I conducted extensive research 

on the biology of N. surinamensis, this was 

an abundant species in the Lincoln, NE, 

area both at carrion and at lights. During 

the early 1990s, however, I have rarely seen 

this insect near the city at lights or at carrion, 

even in places where it used to be abundant. 

Artificial lighting may be a contributing 

factor because I believe that lights decrease 

populations of some nocturnally active insects. 

Insects are attracted to lights where 

they are congregated and easily preyed upon 

by vertebrate scavengers such as toads, 

opossums, and raccoons. Even if they are 

not eaten, they are effectively drawn away 

from their natural habitats and, instead of 

breeding, die from exposure on hot pavement 

or are run over by cars and trucks. I 

believe there is a direct correlation between 

the electrification of rural America and the 

decline of some nocturnal insects. 

Genus NECROPHILA 

Nerophila Kirby and Spence 1828: 509. 

Necrobora Hope 1840: 151. 

Necrotropha Gistel 1848: 121. 

Eusilpha Semenov-Tian-Shanskij 1891: 299. 

Calosilpha Portevin 1920: 396. 

Deutosilpha Portevin 1920: 396. 

Chrysosilpha Portevin 1921: 538. 

Deuterosilpha (misspelling); in Hatch 1928: 

112. 

Necrophila is a genus consisting of about 

20 species (Hatch 1928, Peck and Miller 

1993). With the exception of N. americana, 

all species are found in Asia and India (Hatch 

1928). Necrophila americana is the only species 

found in North America. It is broadly

distributed throughout the eastern half of 

the United States and southern Canada. 

Necrophila americana is the only 

North American silphid with a mostly yellow 

pronotum. That, in combination with 

its broadly oval body shape and reticulate 

elytral sculpturing will easily distinguish 

the only member of this genus in Nebraska. 

Anderson and Peck (1985) indicated the 

larvae are characterized by their black color, 

short two-segmented urogomphi, second 

sternite with a single sclerite, and the presence 

of numerous plates on the sensory area 

of the second antennal segment. 

Necrophila americana (L.) 

(Figs. 25-26, 36, 41-42, 57-58) 

Silpha americana Linnaeus 1758: 360. 

Silpha peltata Catesby 1748: Plate 10, Fig. 7. 

Oiceoptoma terminata Kirby 1837: 103. 

Oiceoptoma affine Kirby 1837: 103. 

Oiceoptoma canadense Kirby 1837: 104. 


Fig. 57. Necrophila americana (L.), male. 


Color yellow, usually with a large, discal, 

black spot. Surface extremely finely punctate 

to rugopunctate, punctures and rugae 

longitudinal. Elytra: Color black, females 

with apical tips usually yellow or rarely all 

black. Surface tricostate (lateral costa occasionally 

indistinct), intervals confusedly 

tuberculate, tubercles irregular in shape 

and usually connected to costae. Apex 

in males rounded, slightly elongated in 

females (Figs. 25-26). 

Distribution. Necrophila americana is 

found in the eastern half of the United 

States and southern Canada (Peck and 

Anderson 1985, Peck and Kaulbars 1987). 

This species occurs throughout Nebraska. 

While there are no records for the panhandle, 

they may also occur there. 

Locality Records (Fig. 58). 3,562 Nebraska 

specimens examined or recorded.

32 


ANTELOPE CO. (2): Neligh; CASS CO. 

(396): Plattsmouth; CHASE CO. (3): Enders 

Reservoir, Imperial; CHERRY CO. (27): 

Dewey Lake, North Loup River, Pelican 

Lake, Valentine; CUSTER CO. (94): Anselmo, 

Sargent; DIXON CO. (8): Aowa Creek; 

DOUGLAS CO. (1): Omaha; FRONTIER 

CO. (5): Farnam; GAGE CO. (2): Wolf-Wildcat 

Creek; HALL CO. (4): Alda; JOHNSON 

CO. (100): No data; KEITH CO. (19): Cedar 

Point Biological Station; KEYA PAHA CO. 

(88): Mills, Norden; KNOX CO. (755): Bazile 

Creek Wildlife Management Area; LIN- 

COLN CO. (4): Box Elder Canyon, Brady, 

North Platte, Wellfleet; LOGAN CO. (1): 

No data; OTOE CO. (53): Nebraska City; 

PAWNEE CO. (105): No data; SALINE CO. 

(1): Swan Creek; SARPY CO. (6): Fontenelle 

Forest, Schramm Park; SAUNDERS CO. 

(3): Wahoo; SHERIDAN CO. (2): Gordon; 

THOMAS CO. (59): Halsey Forest; WASH- 

INGTON CO. (5): Ft. Calhoun. 

Temporal Distribution. Rangewide: March 

to September (Peck and Kaulbars 1987). 


August (1,080), September (4). The August 

numbers reflect a concerted trapping effort 

near Niobrara, NE, in 1994 and is not reflective 

of a true August peak in the population. 

Fig. 58. Nebraska distribution of Necrophila americana. 

Remarks. Necrophila americana is distinctive 

because of the relatively large, 

broadly rounded, dorso-ventrally flattened 

body, and conspicuous, yellow pronotum. 

As in Heterosilpha ramosa, males have 

rounded elytral apices and females have the 

apices slightly prolonged (Figs. 25-26). 

The larvae were described in detail by 

Dorsey (1940) and by Anderson and Peck 

(1985). 

According to Anderson (1982c), adults 

overwinter and begin reproducing in late 

May through mid-July. Larvae were numerous 

during this time, but they appeared 

later than the larvae of other species. Anderson 

observed teneral adults in late July 

through August indicating the emergence 

of the first brood of larvae. These adults 

evidently overwintered (despite their relatively 

early emergence in the year), which 

suggests that there is only one generation 

a year. Development from egg to the 

adult stage took about 10-12 weeks. This 

is a diurnal species (personal observation, 

Shubeck 1971). Clark (1895) and Steele 

(1927) observed adults feeding on maggots 

at carrion. Dorsey (1940) observed 

both adults and larvae feeding on decaying 

flesh, usually near softer parts and excretions. 

However, the larvae fed principally

on the dried remains of hide, sinew, and 

shreds of flesh that are left after the dipterous 

larvae have finished with the carcass. 

Trapping in Nebraska indicated this species 

shows a strong preference for marshy 

and forested areas, a result also observed 

by Lingafelter (1975) in Kansas. 

Fisher and Tuckerman (1986) observed 

that Psithyrus ashtoni (Cresson), a cuckoo 

bumble bee, is mimicked by N. americana. 

Both model and mimic occur in the fieldforest 

interface, have flight periods that 

coincide, share similar flight behavior, and 

have similar coloring and overall appearance 

when flying. Necrophila americana presumably 

acquires some protection from predation 

by mimicking a large, stinging bee. 

Genus OICEOPTOMA 

Oiceoptoma Leach 1815: 89. 

Oeceoptoma Agassiz 1847: 256 (unjustified 

emendation). 

Isosilpha Portevin 1920: 398. 

Seven species are placed in the genus 

Oiceoptoma (Hatch 1928, Peck and Miller 

1993). Three species occur in North America, 

and the remainder are found in Asia with 

one of those reaching Europe. Two species 

are found in Nebraska while the third North 

American species, O. rugulosum Portevin, is 

restricted to the southern United States. 

North American species of the genus 

Oiceoptoma are characterized by having a 

head with a short row of long setae on the 

posterior margin of the eye (Fig. 20), small 

eyes, a pronotum that is widest at the base 

and entirely black or with a black disc and 

dull orange margins, and an elytral shoulder 

with a small tooth (Figs. 27-30). It should 

be noted that while Thanatophilus truncatus 

(Say) also has setae behind the eye, these 

setae are shorter, directed anteriorly, and are 

in a small field behind the posterior margin 

of the eye. 

The larvae are characterized by having 

the urogomphi subequal in length to the 

tenth abdominal segment (Fig. 37) and the 


second segment of the antenna with a large 

sense cone (Fig. 40). 

Oiceoptoma inaequale (Fabricius) 

(Figs. 20, 27-28, 44, 59-60) 

Silpha inaequalis Fabricius 1781: 87. 


Color black. Surface finely and densely punctate, 

with short, black setae. Elytra: Color 

black. Surface finely and densely punctate, 

tricostate. Apex rounded in males, attenuated 

in females (Figs. 27-28). 

Distribution. Oiceoptoma inaequale occurs 

from southern Ontario and Quebec to 

Florida and extends west to Texas and the 

Dakotas (Anderson and Peck 1985, Peck 

and Kaulbars 1987). This species is found 

in Nebraska in approximately the eastern 

two-thirds of the state; there are no records 

for the panhandle or the southwest corner. 

Locality Records (Fig. 60). 427 Nebraska 


ANTELOPE CO. (3): Neligh; CASS CO. 

(4): Plattsmouth, South Bend; CHASE CO. 

(3): Enders Reservoir; CUMING CO. (3): 

West Point; CUSTER CO. (1): Anselmo; 

DAWES CO. (1): No data; DIXON CO. (2): 

Aowa Creek; DOUGLAS CO. (5): Omaha; 

DUNDY CO. (20): 1.5 mi. SW Max, Republican 

River E of Benkelman; FILLMORE CO. 

(1): Fairmont; FRANKLIN CO. (75): 5 mi. S 

Franklin; FRONTIER CO. (108): Farnam, 

Medicine Creek Reservoir, Red Willow 

Reservoir; GOSPER CO. (1): Lexington; 

HALL CO. (3): Alda; HARLAN CO. (8): Republican 

River S of Orleans; JEFFERSON 

CO. (67): Fairbury; LANCASTER CO. (63): 

Lincoln, Reller Prairie, Sprague; LINCOLN 

CO. (16): North Platte, 2 mi. S Sutherland, 

Wellfleet; NUCKOLLS CO. (14): No data; 

SARPY CO. (28): Bellevue, Schramm Park; 

THOMAS CO. (1): Halsey Forest. 

Temporal Distribution. Rangewide: January 

to October (Peck and Kaulbars 1987).

34 


Nebraska: April (21), May (57), June (206), 

July (139), August (3), September (1). 

Remarks. The characters in the key and in 

the diagnosis will serve to separate this species 

from others. It is similar in appearance 

to O. noveboracense (Forster) but lacks the 

dull orange pronotal margin of that species. 

It also vaguely resembles T. lapponicus 

(Herbst) but lacks the distinctive elytral 

tubercles present in that species. 

Dorsey (1940) described the larval 

stage in detail, and Anderson and Peck 

(1985) provided a diagnosis of the larva. 

The shallow emargination of the anterior 

margin of the prothoracic tergite (Fig. 44) 

will separate the larva of this species from 

that of O. noveboracense, which has the 

anterior margin of the prothoracic tergite 

deeply emarginate (Fig. 43). 

Anderson and Peck (1985) suggested 

there is only one generation per year based 

Fig. 59. Oiceoptoma inaequale (Fabr.), female. 

on data from Howden (1950), Reed (1958), 

and Anderson (1982c). In Nebraska, oviposition 

probably occurs from late May to 

June. Goe (1919) observed oviposition in the 

soil, and the egg-laying period in one female 

lasted 36 days with an average of two eggs 

laid per day (range 1-7 eggs/day, total of 62 

eggs). One pair of eggs hatched in six days. 

The length of the first stadium was ten days, 

the second stadium was four days, and the 

third about eight days. Adults emerged 17- 

20 days after the third instars entered the 

soil to pupate. Cole (1942) observed the eggs 

to hatch in about seven days and a larval 

duration of 20 days; pupation took 2-3 weeks. 

Overwintering was in the adult stage. 

Oiceoptoma inaequale is a diurnal species 

(personal observation, Shubeck 1971). 

Lampert (1977) observed that, during flight, 

the elytra are raised to the vertical and held 

together over the back like the wings of a 

resting butterfly. The ventral side of the

elytra are exposed and are a bright, metallic 

blue. He noted that, as the beetle flew, 

the vertically oriented elytra created the 

impression of a much larger insect because 

the ventral surfaces of the elytra flashed 

metallic blue as they reflected light. Upon 

landing and lowering the elytra, the large 

metallic beetle suddenly vanished leaving 

only a small, black one. This has also been 

observed in T. lapponicus (J. Bedick, pers. 

comm., September 1995). The adaptive 

value of this strategy is an abrupt “disappearance” 

from the search image of a 

potential predator. 

This species is most commonly found 

throughout its range in areas of deciduous 

forest (Anderson and Peck 1985). Most of 

the records from eastern Nebraska are also 

from forested areas although specimens 

from west central Nebraska come from 

mostly prairie habitats. Results of a study 

by Shubeck (1984a) showed a slight preference 

for open areas while that of Lingafelter 

(1995) demonstrated a slight preference for 

forests. Although a broad zone of geographic 

overlap occurs between O. inaequale and O. 

noveboracense, one of the two species always 

seems to be locally rare or absent when both 

occur in the same area (Anderson and Peck 

1985). Steele (1927) observed adults feeding 

on maggots at carrion. 


Fig. 60. Nebraska distribution of Oiceoptoma inaequale and O. novaboracense. 

Oiceoptoma noveboracense (Forster) 

(Figs. 16, 29-30, 37, 40, 43, 60-61) 

Silpha noveboracensis Forster 1771: 17. 

Silpha marginalis Fabricius 1776: 215. 

Oiceoptoma marginata Kirby 1837: 100. 


Disc black with broad margins pale 

orange or tan. Surface finely and densely 

punctate, with several longitudinal, dense 

rows of short, tawny setae. Elytra: Color 

light brown (most common) to black. Surface 

finely and densely punctate, tricostate. 

Apex rounded in males, attenuate in females 

(Figs. 29-30). 

Distribution. Oiceoptoma noveboracense is 

a widely distributed species in the eastern 

part of the United States (although it does 

not occur in Florida, Louisiana, and Texas). 

Its range extends west to Oklahoma, Kansas, 

Nebraska, and the Dakotas. There are 

only sporadic records for Wyoming and Montana. 

According toAnderson and Peck (1985) 

and Peck and Kaulbars (1987), it is recorded 

from the southern part of Canada and extends 

westward to Alberta. In Nebraska, 

this species is found statewide except for the 

southern portion of the panhandle as well as 

the southwestern corner.

36 




BUFFALO CO. (3): Kearney; CASS CO. 

(7): Plattsmouth; CHASE CO. (9): Enders 

Reservoir; CHERRY CO. (6): Ft. Niobrara 

National Wildlife Refuge; CUSTER CO. 

(50): Anselmo, Sargent; DAWES CO. (28): 

Ash Creek, Chadron; DIXON CO. (18): Aowa 

Creek; DUNDY CO. (1): 1.5 mi. SW Max; 

FRANKLIN CO. (32): 5 mi. S Franklin; 

FRONTIER CO. (40): Farnam, Medicine 

Creek Reservoir, Red Willow Reservoir; 

GOSPER CO. (3): Lexington; HALL CO. (8): 

Grand Island; HARLAN CO. (6): Republican 

River S of Orleans; JEFFERSON CO. (15): 

No data; KEITH CO. (1): Cedar Point Biological 

Station; KEYA PAHA CO. (38): Mills, 

5 mi. SW Norden; KNOX CO. (13): Bazile 

Creek; LANCASTER CO. (11): Lincoln; 

LINCOLN CO. (158): Box Elder Canyon, 

Cottonwood Canyon, Moran Canyon, North 

Platte, Sutherland, Wellfleet; MERRICK 

CO. (3): No data; NEMAHA CO. (1): Auburn; 

NUCKOLLS CO. (1): No data; OTOE CO. (1): 

Fig. 61. Oiceoptoma novaboracense (Forster), female. 

Nebraska City; SARPY CO. (15): Fontenelle 


(4): Cedar Bluffs, Wahoo; SIOUX CO. (29): 

Ft. Robinson, Gilbert-Baker State Wayside 

Area, Monroe Canyon, Warbonnet Canyon; 


Temporal Distribution. Rangewide: February 

to October (Anderson and Peck 1985; 

Peck and Kaulbars 1987). Nebraska: March 

(1), April (13), May (32), June (66), July (211), 

August (172), September (1). 

Remarks. Oiceoptoma noveboracense is a 

smaller silphid that may be readily identified 

because of its usually light brown color and 

distinctive dull orange color of the pronotal 

margins. Only O. inaequale (Fabr.) is similar, 

but it lacks the marginal coloration of 

the pronotum. 

The larvae were described in detail by 

Dorsey (1940) and by Anderson and Peck 

(1985). The deep emargination of the anterior 

margin of the prothoracic tergite (Fig. 43)

will distinguish the larva of this species 

from O. inaequale, which has the anterior 

margin of the prothoracic tergite shallowly 

emarginate (Fig. 44). 

Adults of this species are diurnal 

(personal observation, Shubeck 1971), and 

the adults are reproductively active in the 

spring (Pirone 1974, Shubeck 1976, Anderson 

1982c). Anderson and Peck (1985) noted 

that in more northerly localities, this species 

is the dominant early season silphine of forested 

areas. Shubeck (1975b) studied flight 

activity as influenced by temperature for O. 

noveboracense. He found that the normal 

temperature range of flight activity was 23° C 

to 30° C with a pronounced peak of activity at 

25° C. Beetles did not fly if the temperature 

was above 30° C or below 23° C. 

In the eastern part of its range, O. 

noveboracense is most commonly found in forested 

habitats (Shubeck 1984a, Lingafelter 

1995), but in west central Nebraska it seems 

to be just as abundant in prairie habitats. 

In Nebraska, there is probably one generation 

a year. According to Anderson and 

Peck (1985), mating and egg laying usually 

occur from about mid-April to late May. A 

female lays 8-10 eggs in the warm soil surrounding 

a carcass. Anderson and Peck 

noted that the egg stage lasted 5-6 days, the 

first instar 4-5 days, the second instar 7-8 

days, and the third instar 10 days. Third 

instar larvae then pupated in the soil. The 

duration of the pupal stage was 2-3 weeks. 

New adults began to appear in July, and 

these individuals then overwintered. 

Clark (1895) noted some feeding by 

adults on fly larvae at carrion. 

Genus THANATOPHILUS 

Thanatophilus Leach 1815: 89. 

Pseudopelta Bergroth 1884: 229. 

Philas Portevin 1903: 331. 

Silphosoma Portevin 1903: 333. 

Chalcosilpha Portevin 1926: 31. 


The genus Thanatophilus contains 20 

species and was last revised by Schawaller 

(1981). Interestingly, Schawaller did not 

include the North American T. truncatus 

in his revision. I do not know if this was 

an oversight or whether he purposefully 

excluded it from the genus . . . in which case 

I would have expected him to mention this 

in his revision. Five species are found in 

North America (Anderson and Peck 1985, 

Peck and Miller 1993) while the remaining 

species occur in Europe and Asia with two 

species in sub-Saharan Africa (Schawaller 

1981). In Nebraska, there are two species. 

Species of Thanatophilus are characterized 

by having widely separated mesocoxae, 

small eyes that lack a short row of long setae 

on the posterior margin of the eye, and elytra 

that lack costae or else have tubercles in the 

intervals. It should be noted that while T. 

truncatus has setae behind the eye, these 

setae are not in a distinct row (but in a small 

field), are directed forward, and are short. 

Only two of the five North American 

species of larvae have been described (including 

only T. lapponicus from the Nebraska 

species). The larvae of these species are 

distinguished by the urogomphi being longer 

than the tenth abdominal segment (Fig. 

35) and the presence of three large sclerites 

on the second sternite (Anderson and Peck 

1985). 

Thanatophilus lapponicus (Herbst) 

(Figs. 31-32, 62-63) 

Silpha lapponica Herbst 1793: 209. 

Silpha caudata Say 1823: 192. 

Silpha tuberculata Germar 1824: 81. 

Silpha granigera Chevrolat 1834: 1. 

Silpha californica Mannerheim 1843: 253. 

Silpha sachalinica Kieseritzky 1909: 126. 

Thanatophilus irregularis Portevin 1914: 221. 

Thanatophilus lapponicus mulleri Portevin 

1932: 58. 

Diagnosis. Length 9.4-14.0 mm. Head and 

Thorax: Color black. Surface with dense, 

tawny setae. Elytra: Color black. Surface

38 


tricostate, densely setose, intervals each with 

a row of small tubercles. Apices in males 

rounded, females with apex attenuate (Figs. 

31-32). 

Distribution. Thanatophilus lapponicus 

ranges broadly throughout Canada and 

Alaska, across the northern United States 

from coast to coast, and south to southern 

California, Arizona, and New Mexico (Anderson 

and Peck 1985, Peck and Kaulbars 

1987). It is also found in northern Europe 

and Asia (Hatch 1928, Schawaller 1981). 

In Nebraska, this species probably occurs 

statewide although it is more abundant in 

the north and west. 

Fig. 62. Thanatophilus lapponicus (Herbst), male. 



ADAMS CO. (2): No data; ARTHUR CO. (2): 

Arapaho Prairie; BUFFALO CO. (5): Kearney, 

Ravenna; CHERRY CO. (25): Dewey 

Lake, Valentine, Ft. Niobrara National Wildlife 

Refuge; CHEYENNE CO. (2): Dalton; 

CUMING CO. (1): West Point; CUSTER CO. 

(100): Anselmo, Milburn, Sargent; DAWES 

CO. (15): Chadron, Pine Ridge area; DAW- 

SON CO. (3): Johnson Lake, Lexington; 

DIXON CO. (2): Concord; DUNDY CO. (6): 

Haigler, 1.5 mi. SW Max, Republican River E 

of Benkelman; FRANKLIN CO. (4): No data; 

FRONTIER CO. (71): Farnam, Medicine 

Creek Reservoir, Red Willow Reservoir;

GARDEN CO. (2): Crescent Lake; GOSPER 

CO. (36): Elwood Reservoir, Lexington, 

Smithfield; HALL CO. (12): Alda; KEITH 

CO. (7): Cedar Point Biological Station; 

KEYA PAHA CO. (8): Mills, Norden; KNOX 

CO. (8): Bazile Creek; LANCASTER CO. 

(5): Lincoln; LINCOLN CO. (235): Box Elder 

Canyon, Cottonwood Canyon, Moran 

Canyon, North Platte, Sutherland, Wellfleet; 

MORRILL CO. (2): Bayard; PHELPS 

CO. (6): Bertrand; PLATTE CO. (3): No 

data; SCOTTS BLUFF CO. (27): Mitchell; 

SIOUX CO. (17): Crawford, Ft. Robinson, 

Gilbert-Baker State Wayside Area, Glen, 

Monroe Canyon, Warbonnet Canyon; 



March to October (Peck and Kaulbars 

1987). Nebraska: April (11), May (24), June 

(149), July (232), August (206), September 

(8), October (5). 

Remarks. Thanatophilus lapponicus is 

readily identified because of the presence 

of a row of small tubercles on each of the 

elytral intervals (Figs. 31-32, 62); it is the 

only silphid in North America with this 

distinctive form of elytral sculpturing. 


Fig. 63. Nebraska distribution of Thanatophilus lapponicus and T. truncatus. 

The larval stage was described by Dorsey 

(1940), and a brief synopsis was given by 

Anderson and Peck (1985). The larva of this 

species is characterized by a dark brown to 

black color on the dorsal surface, urogomphi 

that are about two times the length of the 

10th abdominal segment, and antennae with 

a large sense cone on the second segment (as 

in Fig. 40). 

Overwintering adults become reproductively 

active in late April and May in 

Nebraska. There are two generations a year 

in Nebraska; Anderson (1982c) reported two 

generations a year also in Ontario, Canada. 

Adults of the first generation appear in June 

and early July when they mate and lay eggs. 

Adults of the second generation appear in 

late July through September, and these are 

the overwintering adults. Anderson and 

Peck (1985) reported that individual females 

lay about ten eggs in the soil surrounding a 

carcass. The egg stage lasts 5-6 days, the 

first instar about 7 days, the second instar 

8-10 days, and the third instar 10-12 days. 

Anderson and Peck did not observe pupae. 

Anderson and Peck noted that T. 

lapponicus is a cold-adapted species that 

occurs at higher elevations in the western 

mountains of North America. It is often the

40 


only silphid present in some of these areas. 

Thanatophilus lapponicus shows a strong 

preference for open areas (Anderson 1982c). 

Emetz (1975) reported that this species 

is sometimes injurious to furs, meats, and 

dried fish. Clark (1895) observed extensive 

predation on fly larvae by adult beetles. 

Thanatophilus truncatus (Say) 

(Figs. 33, 63-64) 

Silpha truncata Say 1823: 193. 

Diagnosis. Length 10.5-15.9 mm. Head 

and Thorax: Color black. Surface with small, 

dense punctures, punctures with minute, 

black, adpressed setae. Elytra: Color black. 

Surface with small, dense punctures, costae 

Fig. 64. Thanatophilus truncatus (Say). 

or tubercles absent. Apices of elytra truncate, 

not attenuated, in both sexes (Fig. 33). 

Distribution. Thanatophilus truncatus is 

found from Nebraska southwest to Kansas, 

Colorado, Texas, New Mexico, and Arizona 

(Peck and Kaulbars 1987). It occurs in 

much of Mexico also, ranging as far south 

as southcentral Mexico (Peck and Anderson 

1985). In Nebraska, this species has been 

found as far east as Lincoln and as far north 

as Custer county. 



BUFFALO CO. (2): Kearney; CUSTER CO. 

(5): Sargent; DUNDY CO. (5): Haigler, 1.5 

mi. SW Max; FRANKLIN CO. (1): No data; 

FRONTIER CO. (101): Farnam, Medicine

Creek Reservoir, Red Willow Reservoir; 

GOSPER CO. (22): Lexington, Smithfield; 

HALL CO. (1): Alda; KEITH CO. (4): Cedar 

Point Biological Station; LANCASTER CO. 

(16): Lincoln; LINCOLN CO. (183): Box 

Elder Canyon, Cottonwood Canyon, Moran 


PHELPS CO. (5): Bertrand; RED WILLOW 

CO. (1): McCook. 

Temporal Distribution. Rangewide: May 

to October (Peck and Kaulbars 1987). Nebraska: 

March (1), April (1), May (7), June 

(5), July (279), August (45), September (7), 

October (1). 

Remarks. Thanatophilus truncatus may be 

readily distinguished by its dull black color, 

lack of costae or tubercles on the elytra, and 

truncate elytra. The truncate elytra might 

lead one to conclude that this species is a 

nicrophorine, but the gradually widening antennae 

will place it in the Silphinae. Among 

the Nebraska fauna, this beetle is unique 

in its appearance and is easily identified. 

A cautionary note: T. truncatus possesses 

setae behind the eye that might lead one to 

key it to Oiceoptoma species. In Oiceoptoma, 

however, there is normally a distinct row of 

long setae exactly on the posterior margin of 

the eye whereas in T. truncatus these setae 

are shorter, directed anteriorly, and are in 

a small but distinct field just behind the 

posterior margin of the eye. 

The immature stages of this species 

remain undescribed. 

Virtually nothing is known of the biology 

of this species. Peck and Kaulbars (1987) 

indicated it lives in such diverse habitats as 

grasslands, arid scrub desert, oak-pinyonjuniper 

woodlands, pine forests, and montane 

meadows. Lingafelter’s (1995) study 

in Kansas showed this species had a strong 

preference for open meadows. In Nebraska, 

it has been collected in short grass prairie, 

sandhills, juniper canyonlands, deciduous 

gallery forests, and heavily disturbed tall 

grass prairie habitats. I have collected a 

specimen from dog feces in Arizona. 


SUBFAMILY NICROPHORINAE 

The Nicrophorinae contains the genera 

Nicrophorus (with about 85 species), Ptomascopus 

(three extant and one fossil species), 

and Paleosilpha (one fossil species) (Hatch 

1927, 1928, Peck and Anderson 1985). The 

species of Nicrophorus are found throughout 

the Americas (including the first record from 

the Caribbean region, a new species from the 

Dominican Republic being described by Davidson 

and Rawlins; J. Rawlins, pers. comm., 

October 1995), Europe, and Asia. Most of the 

species are north temperate in distribution. 

They are not found in subsaharan Africa, 

Australia, or India. 

Ptomascopus has three extant species in 

Asia and one fossil species (P. aveyronensis 

Flach) from the Oligocene of France (Hatch 

1927). The monotypic Paleosilpha fraasii 

Flach is also known only from the Oligocene 

of France. 

During most of the last century, the 

Nicrophorinae has shifted back and forth 

from tribe to subfamily status depending 

on the views of the particular specialist at 

the time. Both Hatch (1928) and Peck and 

Miller (1993), in their checklists of the world 

and North American faunas respectively, 

give tribal status to the taxon. Conversely, 

Anderson and Peck (1985), in their treatment 

of the Canadian and Alaskan fauna, Peck 

and Kaulbars (1987), in their distribution 

and bionomics of U.S. carrion beetles, Peck 

and Anderson (1985), in their treatment of 

the carrion beetles of Latin America, and 

Lawrence and Newton (1995), in their new 

classification of beetle families, use the subfamily 

level for the taxon. Subfamily status 

appears to be the current consensus. 

Genus NICROPHORUS 

Nicrophorus Fabricius 1775: 71. 

Necrophorus Thunberg 1789: 7. 

Necrophagus Leach 1815: 88. 

Crytoscelis Hope 1840: 149. 

Acanthopsilus Portevin 1914: 223. 

Necrocharis Portevin 1923: 141.

42 


Necroxenus Semenov-Tian-Shanskij 1926: 46. 

Eunecrophorus Semenov-Tian-Shanskij 1933: 

152. 

Necrocleptes Semenov-Tian-Shanskij 1933: 

153. 

Necrophorindus Semenov-Tian-Shanskij 1933: 

153. 

Necrophoriscus Semenov-Tian-Shanskij 1933: 

152. 

Nesonecrophorus Semenov-Tian-Shanskij 

1933: 153. 

Necropter Semenov-Tian-Shanskij 1933: 154. 

Nesonecropter Semenov-Tian-Shanskij 1933: 

154. 

Stictonecropter Semenov-Tian-Shanskij 1933: 

154. 

Neonicrophorus Hatch 1946: 99. 

The genus Nicrophorus presently contains 

about 85 species distributed in Europe, 

Asia, North and South America (Hatch 1928, 

Peck and Anderson 1985). Most of the species 

occur in Europe and Asia. There are 

15 species in the United States, and all but 

four of these occur in Nebraska (N. defodiens 

Mannerheim, N. nigrita Mannerheim, N. 

sayi Laporte, and N. vespilloides Herbst). 

Nicrophorus defodiens Mannerheim is found 

throughout much of South Dakota, and further 

sampling may indicate its presence in 

northern Nebraska. 

While there is an abundance of literature 

on the taxonomy of the genus and the 

life histories of its species, there has been 

no modern, comprehensive treatment. The 

North American taxa were last reviewed by 

Anderson and Peck (1985). The most recent 

synoptic world catalog is that of Hatch (1928) 

while the latest North American catalog was 

provided by Peck and Miller (1993). Peck and 

Anderson (1985) conducted a preliminary 

phylogenetic analysis of the species groups 

of Nicrophorus as well as a general overview 

of biogeography. 

In the past, the spelling of this genus 

name varied from Nicrophorus to Necrophorus 

and back again. Fabricius (1775) established 

the name Nicrophorus, and it was 

subsequently used in this form by himself 

in other publications, by Olivier (1790), and 

by other contemporaries. Thunberg (1789), 

however, used the spelling Necrophorus, 

and from that point confusion has reigned. 

Hatch (1932) first explored the spelling dilemma, 

but he did not conclude which name 

should be accepted. Herman (1964) provided 

a detailed history of these two names. He 

concluded that Necrophorus was an incorrect 

transliteration (hence, an unjustified 

emendation), and that the original spelling 

should be maintained. 

The adults of nearly all of the species 

of Nicrophorus show parental care in rearing 

their young, and this has resulted in the life 

histories of several species becoming well 

studied, mostly by ecologists. For other species, 

however, very little is actually known 

of their life history or immature stages. 

Fi g s. 65-66. Foretarsus of (65) male and (66) female 

Nicrophorus marginatus showing dimorphism in 

tarsomeres. Also note bisetose empodium. Fig. 67. 

Abdominal tergite of larval Nicrophorus investigator 

showing 4 posterior spines.

Adults in the genus Nicrophorus are 

easily recognized because of the presence 

of truncate elytra that are usually marked 

with conspicuous orange or reddish bands 

or spots. The males of most species possess 

expanded segments of the foretarsus (Fig. 65) 

whereas in females the tarsi are only slightly 

expanded (Fig. 66). Bliss (1949) observed 

that some species of Nicrophorus show secondary 

sexual characters although there are 

no characters common to the entire genus. 

Larvae of Nicrophorus species are distinctive 

because of the presence of quadrispinose 

abdominal tergites (Fig. 67) and 

reduced sclerotization. Anderson (1982b) 

described the larvae of ten Nearctic species 

of Nicrophorus. 

NICROPHORINE BIOLOGY 

Flying upwind against a gentle breeze, 

silvery moonlight reflecting dully from hardened 

wing covers held high over her body, 

orange-tipped antennae fully extended and 

quivering in the warm night airs, a female 

burying beetle searches for the odor of recent 

death. She is seeking the relatively fresh 

remains of a recently dead animal so that 

she and a prospective mate can quickly bury 

it and use it as a food source for themselves 

and their young. 

It is a truism indeed that what passes 

for food to some is absolutely repulsive to 

others. This is due, in part, to culture, familiarity, 

and available food resources because, 

after all, various foods are composed of the 

same basic set of proteins, carbohydrates, 

fats, and sugars. Fortunately, burying 

beetles have no culture, and they dig in (literally) 

with a necessary haste that reflects a 

competitive principle of “better to eat quickly 

than to let the flies have it.” Calliphorid flies 

are often the first to oviposit at carrion and, 

if the eggs are not detected and destroyed 

by Nicrophorus adults, the carcass may be 

consumed by developing fly larvae, causing 

the beetles to abandon the resource. 

The majority of silphids are scavengers 

on dead animals, dung, and decaying 


plant materials, and some prey on snails. 

Although not all silphids bury carrion, the 

orange and black banded species in the genus 

Nicrophorus inter small, dead vertebrates in 

the ground, hence their common names of 

carrion or sexton beetles. There the beetles 

lay eggs and process the remains in order 

to provide a food source for their developing 

larvae. 

Se a r c h i n g Be h a v i o r 

Burying beetles are found primarily in 

temperate regions of the world. They are 

rare or absent in the tropics because they are 

simply out-competed by more efficient carrion 

feeding ants and vultures. Most burying 

beetles are nocturnal, and they search widely 

for carrion. They are remarkably adept 

at detecting the odor of animals that have 

recently died. Using the organs of smell located 

on their antennae, they can find a dead 

mouse, for example, within an hour of death 

and from as far away as two miles (Petruska 

1975)! Attesting to their extreme sensitivity 

in detecting odors is the fact that humans do 

not usually consider that there is any odor 

associated with remains within an hour of 

death. Customarily, however, beetles find 

a carcass after a day or two. Experiments 

conducted by Shubeck (1975a) demonstrated 

that vision did not play a role in searching 

behavior. Most searching behavior is guided 

by the sense of smell. 

Dethier (1947) conducted olfaction experiments 

with several Nicrophorus species, 

Oiceoptoma novaboracense, and Necrophila 

americana. He concluded that the ability to 

perceive odors from a distance resided in the 

antennae while the end organs of the palpi 

detected odors from a short distance. He observed 

that beetles that had their antennae 

surgically removed could still locate carrion 

from short distances (30 inches); it was on 

this basis that Abbott (1927a-b, 1936) had 

previously and erroneously concluded that 

the antennae of Nicrophorus orbicollis, N. 

tomentosus, and Oiceoptoma inaequale were 

of little importance in orientation to odors.

44 


Dethier concluded that the major difference 

between long range and short range odor 

perception probably involved thresholds of 

individual sensillae. These sensillae are 

sensitive to hydrogen sulfide and some cyclic 

carbon compounds (Waldow 1973) that are 

released as a carcass decays. The antennal 

lamellae (the apical three segments) possess 

several hundred sensillae whereas the 

remaining segments have few to none. 

Shubeck (1968) conducted a large mark/ 

recapture experiment using 460 individuals 

of Oiceoptoma novaboracense and 205 individuals 

of N. tomentosus and N. orbicollis. 

Only 2% of released O. novaboracense returned 

to carrion from a distance of 75 m, and 

only 28% returned when released 5 m from 

carrion. Rates of return were even lower 

for the Nicrophorus species. The return 

rate increased as the distance was reduced. 

Shubeck suggested that these species were 

not very efficient at locating carrion. Perhaps 

the “trauma” of capture or some environmental 

factors may have deleteriously influenced 

the return rate inasmuch as these insects 

rely on finding ephemeral patches of carrion 

for survival and breeding. In other words, 

the results of this study may not be indicative 

of the searching ability of silphids. In pitfall 

studies conducted by Wilson et al. (1984) in 

Michigan, 94% of pitfall traps baited with 

mice were discovered within 24 hr, and 95% 

of the discoverers were Nicrophorus species. 

These beetles seem to be abundant relative 

to their resources. 

In studies conducted by Conley (1982) 

using N. carolinus, location of carcasses 

varied from 24-100 hours; when putrefying 

carrion was present, the location time 

was less than six hours. Locating efficiency 

ranged from 10-80% (mean=36%) of available 

carcasses. Of 37 carcasses colonized, 14 

(38%) were occupied by single beetles, and 

10 (27%) were occupied by two beetles. 

Pukowski (1933) observed a colonization 

pattern in six European species of 

Nicrophorus in which male beetles locate 

carrion, produce pheromones to attract a 

female, and then the pair vigorously rebuffs 

all other beetles attempting to colonize the 

carcass. Milne and Milne (1944) found that 

first colonizers of N. orbicollis and N. tomentosus 

may be of either sex, that there was no 

advertisement by males, and that burial and 

brooding was often accomplished by several 

pairs of beetles. Conley’s observations of N. 

carolinus concurred with those of the Milnes. 

Conversely, Wilson and Fudge (1984) reported 

that while several beetles may find 

and bury a carcass together, a single pair 

will eventually drive off the others and 

secure the carcass for themselves. Wilson 

and Knollenberg (1984) noted that success in 

finding carrion is influenced by many factors, 

including density of competing vertebrate 

scavengers, density of competing individuals 

of Nicrophorus, individual searching ability, 

reproductive condition, and temperature. 

A male that is successful at locating a 

carcass emits a sex pheromone that serves 

to attract a sexually receptive female (Pukowski 

1933, Eggert and Müller 1989b). 

Often climbing to a higher perch, the male 

assumes a “headstand” position with the 

head held down and the fully extended abdomen 

pointing upward (Fig. 68). This exposes 

the last abdominal segment from which 

the pheromone is released as the tip of the 

tip is moved up and down slightly in such 

a way that the intersegmental membranes 

can be seen (Eggert and Müller 1989b). 

This segment is supplied with a number of 

cuticular pores and lined with specialized 

epithelial gland cells (Eggert and Sakaluk 

1995). Studies by Eggert (1992) showed 

that Nicrophorus males emit pheromones 

both when they have found a carcass and 

when they have not. It seems likely that 

females cannot always tell whether a male 

is emitting pheromones on or off a carcass 

until there is physical contact between the 

two (Eggert and Müller 1989a). What, 

then, is the benefit accruing to a female 

responding to a pheromone emitter without 

a carcass? Eggert suggested that the 

benefit obtained by the female is obtaining 

an adequate sperm supply for when she 

finds a carcass on which no mate is present.

A few field studies have shown that some 

Nicrophorus females can often raise their 

brood without a mate (Scott and Traniello 

1990b, Eggert 1992). Under these circumstances, 

females depended on sperm transferred 

from a male during previous matings 

for fertilization of their eggs. Eggert (1992) 

demonstrated with N. vespilloides that sperm 

stored in a female’s spermatheca started to 

become infertile three weeks after insemination, 

even when the female had not produced 

any eggs in the meantime. The reproductive 

period of the female is longer than that of the 

male and may last several months. 

Fig. 68. Male Nicrophorus sp. emitting pheromone in 

typical “headstand” position. Illustration from cover 

of Behavioral Ecology 3(3), 1992. Used by permission 

of Oxford University Press. 

Bu r i a l a n d Pr e P a r a t i o n o f t h e ca r c a s s 

After arriving at a carcass, a male/ 

female pair will first examine the body (Fig. 

69) and assess its size by trying to move it. 

Adults make short trips to nearby terrain 

to find a suitable spot for burial (Scott and 

Traniello 1989). Less interest seems to be 

shown if the carcass is too large for burial although 

it can be an important food resource 


for the adults (Wilson and Knollenberg 

1984). This may be because large carcasses 

are more difficult to transport or because 

a pair of beetles cannot bury it before the 

arrival of competitors, whether they be conspecifics, 

other Nicrophorus species, or flies 

(Trumbo 1990b, Eggert and Müller 1992). 

Smaller species, such as N. tomentosus and 

N. defodiens, bury carcasses just below the 

leaf litter while larger species take carcasses 

to greater depths beneath the soil (Pukowski 

1933, Wilson and Knollenberg 1987). 

According to Muths (1991), Nicrophorus 

species were tested in the laboratory 

to determine if they discriminated 

between different substrates when burying 

a carcass. His results suggested beetles do 

discriminate, preferring substrates with 

higher “bulk” (i.e., grass clippings) over 

those without. Factors other than those 

influencing the speed of concealment probably 

influence burial site selection. A stable, 

non-collapsing burial chamber to hold the 

carcass and provide a nursery for the larvae 

is important for successful reproduction 

(Pukowski 1933, Milne and Milne 1976). 

Muths indicated that discrimination may be 

explained mechanistically in terms of substrate 

qualities such as ease of excavation 

and suitability in the substrate for stable 

burial chamber construction. Alternatively, 

discrimination can be explained functionally 

in terms of response to competition, 

where immediate burial insures exclusive 

resource use. It is not clear which explanation 

is most appropriate. There may be a 

tradeoff; immediate burial in a less than 

optimum substrate may be the best choice 

in situations where competition is intense, 

but delayed burial may be a better strategy 

where competition is less intense and 

optimum substrate is available only some 

distance from the carcass. 

If, after an exploration of the surrounding 

soil, the ground is found to be too hard for 

burial, the pair of beetles (working together) 

may move mouse-size remains three to four 

feet per hour for as much as three hours 

until a substrate soft enough for burial

46 


is found. It remains unknown how a pair of 

beetles can “agree” on a burial site or how 

they are able to keep the carcass moving 

uniformly in one direction. The soil at the 

burial site is loosened by “plowing” through it 

in much the same fashion as does a bulldozer 

(Fig. 70). Using its head, a beetle presses 

its head in and forces soil upward until it 

crumbles. Roots are forced aside or chewed 

through. Gradually, soil from beneath the 

carcass is displaced to each side, and the 

carcass settles into the ground and is buried 

by several inches of soil (Figs. 71-72). 

Milne and Milne (1944), working with N. 

tomentosus, observed burial chambers 6.5 

cm long and 2.7 cm wide and deep. Burial 

is usually completed in five to eight hours 

although some beetles will continue for days 

if obstructions slow their work. Immediate, 

nocturnal burial is important to these beetles 

because this prevents flies from laying eggs 

on the remains. Although fly larvae can be 

eaten or killed by the adult beetles, the presence 

of numerous fly larvae would make the 

remains unsuitable for the beetles and their 

young. In smaller species of Nicrophorus, 

brood failures may be more common on large 

rather than smaller carcasses because of 

uncontrollable fly infestations (Trumbo and 

Fiore 1994). 

Scott and Traniello (1987), working 

with N. tomentosus, suggested that animals 

that require a patchy food source would be 

expected to make use of cues provided by 

that resource to trigger ovarian maturation. 

Newly eclosed females have previtellogenic 

ovaries that gradually develop to the resting 

stage in about three weeks (Wilson and Knollenberg 

1984). These authors demonstrated 

that females at this stage were reproductively 

capable, and their ovaries matured 

rapidly (48 hr) when they were given a dead 

mouse to bury. Scott et al. (1987) demonstrated 

that the particular cue that brings 

about the physiological changes for ovarian 

development is the behavior of the female 

when she assesses, prepares, and attempts to 

bury a carcass. The location and burial of a 

suitable carcass is followed by vitellogenesis 

and rapid maturation. They noted that the 

mere presence of carrion was not sufficient 

in and of itself to trigger vitellogenesis nor 

were nutritional or feeding cues, presence of 

a male, or mating. 

After burial, the beetles use their large 

jaws to strip away fur or feathers and work 

the remaining mass into a compact ball 

(Fig. 73). They will then “inoculate” the 

remains with oral and anal secretions that 

preserve the carrion and modify the course 

of decomposition (Scott et al. 1987). The 

female usually constructs a short chamber 

above the carrion in which she lays 10-30 

eggs (Fig. 74). Wilson and Fudge (1984) 

conducted lab experiments with N. orbicollis. 

They found that the number of eggs 

laid by females varied directly with the 

body size of the female and not with the 

size of the carcass. The smallest number 

of eggs laid was 22. The largest number of 

larvae that survived to pupation was 26 in 

a single brood. Nearly all females laid more 

eggs than would ultimately survive. When 

adults were experimentally removed prior 

to hatching of the eggs, many broods failed 

completely. Those that survived resulted 

in few larvae, and these were much smaller 

than when the adult was present. Hatching 

success was high, but a thick mold grew 

over the carcasses in most cases when the 

parents were absent. Müller et al. (1990), on 

the other hand, concluded that the number 

of eggs was positively correlated with carcass 

weight in the European Nicrophorus 

vespilloides, and that there was no correlation 

between female body size and clutch size 

as reported by Wilson and Fudge (1984). 

Trumbo (1992) experimented with N. 

orbicollis, N. tomentosus, and N. defodiens 

and found that they can raise a maximum of 

35-50 young. In contrast, N. pustulatus (a 

brood parasite) is unique among Nicrophorus 

species because it can raise nearly 200 young 

on larger carcasses. Trumbo observed further 

that N. orbicollis and N. sayi were extremely 

dependent on parental regurgitations, and 

that young failed to survive to the second 

instar if parents were removed. Young of N.


Fi g s. 69-74. Burial of a mouse by a pair of Nicrophorus beetles. As the beetles remove soil from beneath the 

carcass, it slips downward and is ultimately covered by about 3 cm of soil. After burial, a chamber is made. The 

skin of the mouse is removed and the remains are fashioned into a ball. A shallow depression is made on top of 

the ball to receive liquified food that the adults regurgitate there. From The Social Behavior of Burying Beetles 

by L. and M. Milne. Copyright © 1976 by Scientific American, Inc. All rights reserved.

48 


defodiens, N. pustulatus, and N. tomentosus 

survived without parental regurgitations. 

Müller and Eggert (1989) determined 

that males of Nicrophorus species are able 

to achieve a high level of paternity (mean = 

92%). The mechanism they employ is a repeat 

mating tactic, i.e., the female is mated 

frequently shortly before and during oviposition. 

Repeated matings are essential for 

a high reliability of paternity since single 

copulations resulted in the fertilization of 

only a small number of the female’s eggs. 

Pa r e n t a l ca r e 

After returning to the carcass, the female 

prepares a conical depression on the top of it 

(Fig. 74). Both parents regurgitate droplets 

of partly digested food into the depression. 

The fluid accumulates as food for the larvae 

that will hatch in a few days (cover figure). 

First instar larvae, as well as older larvae 

that have just molted, also approach any 

adult and press their mouth parts against 

the jaws or palps of the adult. This action 

stimulates regurgitation directly to the larva 

(Milne and Milne 1976). Although some species 

require parental regurgitation (at least 

initially), a few other species (including N. 

tomentosus) can develop normally without 

parental feeding (Trumbo 1992, Scott 1994b). 

Otherwise, growing larvae feed directly from 

the depression (Fig. 75) or pull fragments 

from the surface of the carrion ball. 

The larvae receive parental care during 

the entire time they are feeding and growing. 

This is an extremely rare and highly developed 

behavior in insects, a condition normally 

found only in the social bees, wasps, ants, 

and termites. Both adults regur-gitate food 

to begging larvae (Scott 1989), a behavior 

also seen in birds and their nestlings. The 

larvae grow very rapidly and are soon able 

to feed themselves. Parental regur-gitation 

to larvae is reduced by the third day, and 

after the fourth day there is very little. It is 

probable that parents continue to facilitate 

feeding throughout larval development by 

Fig. 75. Nicrophorus larvae feeding within the prepared remains of a mouse. Photo by D. S. Wilson.

preparing the feeding cavity of the carcass 

(Scott and Traniello 1990a). The oral secretions 

are very proteolytic (Wilson and Knollenburg 

1987). In a study by Fetherston 

et al. (1994), both single males and single 

females regurgitated to larvae and maintained 

carrion more frequently than paired 

males and females. Their data suggested 

male and female burying beetles increase 

their brood-care behavior to compensate for 

loss of a mate. This study may represent the 

first demonstration of compensation for mate 

loss in an invertebrate. 

Bartlett and Ashworth (1988) and Scott 

and Traniello (1990b) demonstrated that larval 

weight correlated negatively with larval 

numbers. Neither the duration of parental 

regurgitation per larva nor the duration of 

activity within the carrion had a significant 

effect on mean larval weight (Fetherston 

et al. 1990). Consequently, the number of 

larvae, rather than the duration of parental 

feeding per larva, was the critical factor that 

determined mean larval weight. The mechanisms 

that parents use to determine brood 

size are poorly understood (Trumbo 1990c). 

Sometimes, if the size of the brood is too large 

to be successfully reared on a small carcass, 

both adults will regulate brood size by selective 

infanticide of smaller larvae, usually during 

the first 24 hours after eclosion, so that 

the remaining young will have a sufficient 

food supply (Bartlett 1987, Trumbo 1990c). 

Trumbo and Fernandez (1995) examined the 

ability of male N. orbicollis to regulate brood 

size, and they also manipulated the mass and 

volume of carcasses to determine whether 

correlates of these factors were important as 

cues in the assessment of resource potential. 

They found that single males of N. orbicollis 

raise broods of similar number and mass as 

those raised by single females and pairs. The 

results of their mass and volume manipulations 

with carcasses indicated that burying 

beetles use volume-related cues (not mass) 

to assess resource potential. This study 

indicated that, in the absence of the female, 

males perform the additional task of brood 

size regulation. Direct and early culling of 


larvae in excess of the carrying capacity of 

the carcass is the most unusual feature of 

infanticide by Nicrophorus species and is a 

consequence of their particular biology (Bartlett 

1987). Such intentional parental cannibalism 

is exceptional among invertebrates 

(Trumbo 1990c). 

The effect of male presence, if he remains 

until larval development is complete, 

may be to decrease the number or weight 

of the larvae reared (Scott 1989, Scott and 

Gladstein 1993). Conversely, the male’s 

presence greatly reduces the probability that 

the carcass will be taken over by a conspecific 

intruder and the brood killed (Scott 1990, 

Trumbo 1990b, 1991, Scott and Gladstein 

1993). Scott and Gladstein (1993) noted 

that the cause of increased larval mortality 

(especially on small carcasses) when the 

male is present appears to be due to both decreased 

resources available to larvae and to 

infanticide. On small carcasses, the amount 

consumed by an adult male was equivalent 

to the amount of resources required for one 

larva. Fetherston et al. (1990), for example, 

observed females of N. orbicollis attempting 

to drive off males from carcasses of 25-30 g. 

Since the presence of the male may result 

in a decrease in total brood weight, and the 

female is subject to a lower cost than the 

male if the male deserts and is replaced by 

an intruder (Scott 1989), the female may 

prefer that the male leave the brood chamber 

earlier than the male would prefer to leave 

(Fetherston et al. 1990). 

The adults continually tend the carcass, 

removing fungi and covering the carrion ball 

with a presumed antibacterial secretion. To 

my knowledge, research has not yet been 

conducted on this phenomenal ability to 

inoculate and alter the course of decomposition. 

On the other hand, Solter et al. (1989) 

examined the midgut, hindgut and associated 

hemolymph of Nicrophorus tomentosus, 

Necrophila americana, and Oiceoptoma 

novaboracense. They found 19 species of 

bacteria. In their study, six species of bacteria 

were also associated with the skin, an 

additional four with the respiratory system,

50 


and ten species with the gastrointestinal 

tract, all of humans. 

In a similar study, Berdela et al. (1994) 

recovered 607 isolates consisting of 42 different 

strains of bacteria from Nicrophorus 

tomentosus, N. orbicollis, Oiceoptoma 

novaboracense, O. inaequale, Necrophila 

americana, and Necrodes surinamensis. Of 

these, 52.1% were gram negative bacteria, 

21.1% were coagulose-negative staphylococci, 

8.1% were obligately anaerobic bacteria, 

7.6% were streptococci, 5.4% were Bacillus 

spp., 4.4% were Aerococcus spp., and less 

than 1% were coryneform bacteria. Many 

of these species are known opportunistic 

pathogens. 

After about a week, the larvae have 

consumed all but the bones of the carcass 

and, at this time, one or both adults break 

out of the chamber and fly away. The young 

pupate in the nearby soil about two weeks 

after hatching and emerge as adults about 

a month later. Overwintering then occurs 

in the adult stage. 

ag o n i s t i c Be h a v i o r 

Suitable carcasses are scarce relative 

to the number of potential breeders (Wilson 

and Fudge 1984, Trumbo 1992), and beetles 

will accept a broad size range of carcasses 

(Trumbo and Eggert 1994). On a small carcass, 

fights reduce the resident population 

to a dominant male-female pair (Pukowski 

1933, Wilson and Fudge 1984). Intrusions 

and takeovers appear to be a regular feature 

of the breeding system of Nicrophorus species 

(Trumbo 1990a). 

Trumbo (1990a) observed that interactions 

between intruders and residents were 

agonistic, and infanticide occurred regularly 

as a consequence of a takeover. Intruders 

of both sexes generally kill larvae of the 

resident parents when they find them on 

the carcass. Once infanticide began, larvae 

were not killed all at once but opportunistically 

as the intruder inspected the carcass. 

Intruders pierced the integument of larvae 

with their mandibles and handled them for 

3-72 seconds. Those held the longest were 

almost entirely consumed while those held 

for only a few seconds were dropped, immobilized 

but alive, and later consumed by the 

intruder or the resident. Milne and Milne 

(1944) saw that N. orbicollis became quite 

excited when expelling a competitor and 

stridulated audibly by rubbing the upper 

surface of the abdomen against the bottom 

surface of the elytra. The outcome of fights 

is largely determined by relative body size 

of the combatants (Pukowski 1933, Bartlett 

and Ashworth 1988, Otronen 1988). Scott’s 

(1994a) study showed that male presence in 

N. defodiens is generally ineffective against 

larger, congeneric intruders. Competition 

with other beetles can be severe. Scott and 

Traniello (1990b) observed no significant differences 

between males and females in four 

species of Nicrophorus in the proportion of individuals 

suffering injury (loss of body parts) 

from competitive fights for carcasses. 

Müller et al. (1990) observed (in N. 

vespilloides) that females losing fights did 

not immediately abandon the carcass. Instead, 

they often stayed to lay their own 

eggs. In their lab experiments, some of the 

loser’s larvae were cared for by the winner 

and survived to adulthood in 60% of the 

cases observed. These experiments showed 

a positive correlation between the parents 

duration of stay near the carcass and her 

chances of parasitizing the other female’s 

brood. Müller and Eggert (1990) further 

observed that winning females did not discriminate 

against unrelated larvae if they 

arrived on the carcass within a window of 

time that corresponded with the oviposition 

period of the dominant female. Nicrophorus 

vespilloides females responded parentally 

only to larvae they encountered during a 

distinct and relatively short parental phase. 

Before or after that phase, larvae encountered 

were killed and eaten, irrespective of 

carrion availability. These authors believe 

that the time dependency of infanticidal behavior 

and parental care is an example of a 

simple “rule of thumb” (Dawkins 1979) that 

increases the probability that parental care

is allocated to the parents’ own offspring. 

Recognition of individuals by the parents is 

not involved and may not even occur. 

Mi t e re l a t i o n s h i P s 

Nicrophorus species have a mutualistic 

relationship with phoretic mites of the genus 

Poecilochirus (Mesostigmata: Parasit-idae). 

Springett (1968) gave the following overview. 

Deutonymphs are carried by Nicrophorus 

adults to carrion where the mites leave the 

beetle and feed on fly eggs and small larvae. 

If the carcass is buried by the beetle, the 

feeding behavior of the mite (on fly eggs) 

helps to ensure successful breeding by the 

beetle. When the carcass is buried, both 

beetles and mites reproduce underground. 

When the Nicrophorus larvae pupate, the 

adult female beetle abandons the brood 

chamber, and large numbers of deutonymphs 

are carried by her to another carcass. The 

continuity of the association is maintained 

when the mite deutonymphs join the beetle 

larvae when they pupate or encounter other 

adult Nicrophorus at a carcass. The mites 

may not be completely dependent on Nicrophorus 

species for successful reproduction, 

and Nicrophorus species would be much 

less successful in competing with the larvae 

of Calliphora fly species without the mites. 

Brown and Wilson (1992) observed that P. 

carabi usually do not reproduce on large 

carcasses but wait until their Nicrophorus 

host has found a small carcass suitable for 

its own reproduction. 

In more detailed observations of Poecilochirus 

carabi and Nicrophorus vespilloides 

in Europe, Schwarz and Müller (1992) observed 

the first deutonymphs of the new mite 

generation aggregated on the male beetle. 

The mites did not use sex-specific traits 

to discriminate between male and female 

beetles in the brood chamber but traits that 

are related to the behavior of the beetles. 

When the male beetle departed, it carried 

away nearly all the deutonymphs then present 

in the brood chamber. Deutonymphs that 

developed later congregated on the female 


beetle, which left the chamber several days 

after the male. Only those deuto-nymphs 

that missed the female’s departure dispersed 

on the beetle larvae; this meant they had to 

wait in the pupal chambers until the beetles 

completed their development. On average, 

86% of the deutonymphs left the brood chamber 

on the parent beetles. 

Springett’s (1968) laboratory experiments 

demonstrated that the presence of 

mites helps to enable successful reproduction 

of beetles whereas in cultures lacking mites 

(but with calliphorid eggs), the Nicrophorus 

adults were unsuccessful in rearing young. 

Calliphorid eggs hatched only when mites 

were absent because the mites were such 

avid predators of the fly eggs. Beetles could 

then utilize the carcass without the competition 

from fly larvae. Adult Nicrophorus do 

eat fly larvae but were never successful in 

killing all of them. Mites would also attack 

fly larvae but only those less than 5 mm. 

The presence of mites does not guarantee 

the absence of flies. In studies conducted by 

Scott and Traniello (1990b), 9.1% of carcasses 

buried (N=33) were lost to flies in June, 

44.7% (N=38) were lost in July, and 66.7% 

(N=15) were lost to flies in August. 

Deutonymphs stayed with the Nicrophorus 

larvae and remained active within 

the pupal cell throughout the pupal period 

(Springett 1968). During this time it was 

impossible for mites to free themselves from 

the pupal cell, and the beetles emerged as 

adults bearing the deutonymph mites. The 

deutonymphs are chemically attracted to 

burying beetles and prefer them to non- 

Nicrophorus species; they could not be induced 

to use other carrion beetles as hosts 

(Springett 1968, Korn 1983). 

Beninger (1993) and Blackman and 

Evans (1994), working with Nicrophorus 

vespilloides in Europe, observed the mites Poecilochirus 

carabi G. and R. Canestrini and P. 

davydovae Hyatt attacking the eggs of their 

host burying beetles with a consequent reduction 

in brood size. Their studies indicate that 

mite predation on burying beetle eggs occurs, 

and that further studies are needed.

52 

ne M a t o d e re l a t i o n s h i P s 


Richter (1993) reported that Rhabditis 

stammeri (Völk), a carrion-dwelling 

nematode, is specifically associated with 

Nicrophorus vespilloides. Juveniles of the 

nematode use the adult beetle for transport 

to carrion where both adults and larvae of 

beetles become infected. Inside the beetle 

larvae, the juvenile nematodes are transported 

to the pupal chamber via the larval 

gut. After pupation, the juvenile nematodes 

are found in the pupal chamber at protected 

places such as the exuvium or beneath the 

wings of the pupa. After emergence from 

the pupa, the nematodes migrate to the gut 

and genitalia of the adult beetles. Rhabditis 

stammeri is also transmitted from one adult 

beetle to another during copulation. Infection 

by nematodes probably occurs in many 

other species of Nicrophorus as well, and is 

an area needing additional study. 

st r i d u l a t i o n 

All species of Nicrophorus have a stridulatory 

structure in both males and females 

(Fig. 2). Stridulation is used during burial of 

the carcass, copulation, and the interaction 

between the female and her brood (Niemitz 

1972, Niemetz and Krampei 1972). Huerta 

et al. (1992) conducted lab experiments 

with N. mexicanus and found that a lack of 

stridulation in the female resulted in poor 

or no “bonding” between the female and her 

offspring, which negatively affected larval 

survival. Inhibition of stridulation in the 

male affected and sometimes precluded 

copulation. Lack of stridulation in both nest 

partners may affect the coordination of nest 

preparation (Halffter 1982, Huerta et al. 

1992, Halffter et al. 1983). 

Pr e d a t i o n 

Most ground-feeding insectivorous 

birds are probably familiar with burying 

beetles, and these beetles have been recorded 

as among the food items of several species, 

especially crows that routinely visit carrion. 

Jones (1932) conducted a number of experiments 

using N. americanus, N. orbicollis, 

and N. pustulatus to determine if a variety 

of insectivorous birds would be deterred by 

these aposematically colored beetles. Although 

he tested only seven specimens, none 

were eaten by the birds, while 42 out of 46 

other beetles (representing seven species) of 

comparable size were eaten by seven species 

of birds during 93 feeding events. Jones 

concluded that birds do avoid these brightly 

colored beetles, at least when other food is 

available. 

so c i a l i t y 

The activities of adult Nicrophorus 

species in rearing their young is the highest 

level of sociality attained in the Coleoptera 

(Wilson 1971, Wilson and Fudge 1984). On a 

large carcass, the mating system is variable. 

Larger carcasses can support greater numbers 

of larvae and support broods of greater 

total mass than smaller carcasses (Trumbo 

1992). Consexual adults often tolerate each 

other and often feed each other’s young in 

a quasisocial fashion (Scott and Traniello 

1990a, Eggert and Müller 1992, Trumbo 

1992, Scott and Williams 1993, Trumbo and 

Wilson 1993). According to Trumbo and Wilson 

(1993), females of smaller Nicrophorus 

(N. defodiens, N. tomentosus) species were 

much more likely to feed young cooperatively 

than females of, for example, N. orbicollis. 

They hypothesized that since adults cannot 

discriminate between related and unrelated 

young, they feed any larvae on the carcass 

to ensure adequate care for their own young. 

Larger carcasses were more difficult to exploit 

because: (a) they took longer to conceal 

beneath the leaf litter; (b) they were less 

likely to be rounded into brood balls; (c) they 

were more likely to be utilized by dipterans; 

and (d) they were occupied by greater numbers 

of congeners (Trumbo 1992). 

Scott (1994b) suggested that competition 

with flies promotes communal breeding 

in N. tomentosus. She demonstrated that

oods reared on small carcasses by foursomes 

were 32% larger than those reared 

by a pair, and that on larger carcasses they 

were 49% larger when the beetles were in 

competition with flies. Her study showed 

that subordinate males provided longer care 

on flyblown carcasses, which indicated there 

is a benefit gained when both males assist. 

In these cases, the subordinate remained a 

few days longer to help eliminate maggots 

and fly eggs. Scott indicated that the actual 

frequency of communal breeding in the field 

depends on the density of the beetle population 

as well as other factors because a carcass 

must be discovered by several consexual 

adults within 24 hours, or else the resulting 

late-arriving larvae will be killed by the 

resident adults. 

A female can produce two broods in 

a short time in response to partial or total 

brood failure (Müller and Eggert 1987), an 

intruder male that destroys her initial brood 

(Trumbo 1987, Scott and Traniello 1990a), or 

completing a reproductive cycle and locating 

a second carcass (Scott and Traniello 1990b). 

In similar experiments, Müller (1987), Scott 

and Traniello (1990a), and Trumbo (1990c) 

found that the number and mean mass of 

larvae declined between the first and second 

reproductive events. 

If a male fails to discover the carcass, 

a female can breed on her own using stored 

sperm (Bartlett 1988, Scott 1989, Trumbo 

1990b). Females acquire sperm by copulating 

with males that emit pheromones in the 

absence of a carcass or by copulating with 

males on large carcasses where feeding only 

occurs (Müller and Eggert 1987, Eggert and 

Müller 1989a, Trumbo and Fiore 1991). Both 

males and females can breed more than once 

in a season (Bartlett and Ashworth 1988, 

Scott and Traniello 1990a). 

Trumbo (1991) found in his studies 

of N. orbicollis that males investing time 

with their brood forfeit time spent searching 

for new breeding opportunities. Such 

costs imply that some benefits are derived 

from paternal care. On large carcasses, 

larvae raised by pairs were further along in 


development than broods raised by single 

females. Trumbo saw no differences on small 

carcasses. On large carcasses, the brood is 

vulnerable for a shorter period of time when 

the male is present. Trumbo concluded that 

at least one parent is needed throughout 

development to defend against predators, 

control microbial activity, and open new 

portions of the carcass to larval feeding. The 

major benefit, then, of male assistance lies 

in guarding against intraspecific competition 

after the carcass is buried. The presence 

of both parents dramatically reduced the 

probability that conspecifics will usurp the 

resource, replace either the male or female, 

kill the newly hatched brood, and produce a 

replacement clutch (Scott 1990). 

Eggert and Sakaluk (1995) observed 

that the reproductive interests of the sexes 

often do not coincide, and that this fundamental 

conflict probably underlies a variety 

of sex-specific behavioral adaptations. 

Sexual conflict arises when a pair of beetles 

secures a carcass that can support more 

offspring than a single female can produce. 

They noted that in such a situation, any 

male attracting a second female sires more 

surviving offspring than he would by remaining 

monogamous whereas the female’s 

reproductive success decreases if a rival 

female is attracted to the carcass. Their observations 

and those of Trumbo and Eggert 

(1994) demonstrated that monogamously 

paired males on large carcasses do attempt 

to attract additional females by means of 

pheromone emission whereas males on small 

carcasses do not. In a series of remarkable 

experiments, Trumbo and Eggert (1994) and 

Eggert and Sakaluk (1995) demonstrated 

that females physically interfered with male 

polygnous signaling using various behavioral 

tactics. These interference tactics included 

mounting, pushing him from a perch from 

which he emits pheromones, undercutting 

the male, or pinching his abdomen with her 

mandibles. 

Halffter (1991), among others, concluded 

that subsocial behavior in general is 

accompanied by a reduction in the number of

54 


offspring and that this is compensated for by 

a significant reduction in juvenile mortality. 

He observed that subsociality in Nicrophorus 

species was favored by several behavior patterns: 

(1) A reduction in the effort needed for 

both sexes to find one another. The characteristic 

discontinuity of carcass distribution 

in space and time favors the sexes meeting 

and cooperating at the food source; (2) The 

relative ease by which 

the sexes meet may simplify precopulatory 

mating behavior, thus reducing valuable 

energy expenditures; (3) The food needs to 

be manipulated by the parents before being 

ingested by the larvae. This includes 

transport, burial, processing, guarding, 

and tending the food. This is accomplished 

more efficiently by pairs of beetles; (4) The 

evolution of cooperative nesting and care of 

progeny generally favors pair bonding that is 

accompanied by chemical, acoustic, or tactile 

behaviors. 

Of principal importance to these beetles 

and their young is the burial of the food 

resource because this effectively removes 

it from the arena of intense competition by 

maggots, other carrion-feeding insects, and 

even mammal scavengers. Carrion is an 

ephemeral, unpredictably encountered food 

source, and its “bonanza” nature is so valuable 

to the prospective parents that they 

bury it to keep it from being stolen. Burying 

beetles are unique among the silphids 

because they are the only ones that break 

the cycle of competition at the food source 

while, at the same time, providing their 

larvae with a considerably safer subterranean 

environment in which to develop that 

is relatively free from predators. Burying 

beetles exhibit one of the most advanced 

forms of parental care described among 

Coleoptera (Zeh and Smith 1985). 

According to Scott and Traniello 

(1990a) the principal determinants of 

reproductive success for burying beetles 

are clear. There is a positive correlation 

between brood mass and carcass mass (Kozol 

et al. 1988, N. americanus; Wilson and 

Fudge 1984 and Scott and Traniello 1990b, 

N. orbicollis; and Robertson 1992, N. orbicollis 

and N. pustulatus). Beetles probably 

do not bury larger carcasses because they 

are too heavy to move or because a pair of 

beetles cannot bury it before the arrival of 

competitors, whether they be conspecifics, 

other Nicrophorus species, or flies (Eggert 

and Müller 1992). The size of the carcass 

is the most important factor influencing the 

total weight of the brood. Offspring size is 

increased by care from the primary parents 

throughout larval development by facilitation 

of feeding even though the presence 

of each parent may reduce the amount of 

food available to offspring (Scott 1989). The 

presence of the second parent reduces the 

probability that the carcass will be taken 

over by a competitor and the brood killed. 

Males almost always participate in defense 

and feeding of the brood (Bartlett 1988). 

KEY TO THE SPECIES OF ADULT 

NICROPHORUS IN NEBRASKA 


1. Pronotum lacking anterior, transverse 

impression; lateral margins extremely narrowly 

explanate (Fig. 76) . . carolinus (L.) 

1’. Pronotum with anterior, transverse impression; 

lateral margins broadly explanate 

(Figs. 77-83) . . . . . . . . . . . . . . . . . . . . . . . . 2 

2. Frons and pronotal disc red. Tarsal 

empodium quadrisetose . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . americanus Olivier 

2’. Frons and pronotal disc black. Tarsal 

empodium bisetose (Figs. 65-66) . . . . . . . . 3 

3. Pronotum with dense yellow pubescence 

. . . . . . . . . . . . . . . . . . . tomentosus Weber 

3’. Pronotum glabrous or with sparse setae 

on anterior or lateral margins . . . . . . . . . 4 

4. Posterior angle of metepimeron with 

dense, yellow pubescence (Figs. 84, 87, 89) . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 

4’. Posterior angle of metepimeron glabrous 

(Figs. 85, 86, 90, 91), or with dark 

setae (Fig. 88), or with only a few yellow setae 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

5. Anterior face of procoxa with short setae 

on basal half, setae about half length of those 

on humerus . . . . . . marginatus Fabr. 

5’. Anterior face of procoxa with long setae 

on basal half, setae as long or longer than 

those on humerus . . . . . . . . . . . . . . . . . . . 6 

6. Penultimate antennal segment with 

outer edge only moderately emarginate (Fig. 

94); basal segment of antennal club black or 

orange; if club orange, elytron with anterior 

black band crossing onto epipleuron (Fig. 

84); if club black, orange elytral maculations 

reduced or absent and epipleuron bicolored . 

. . . . . . . . . . . . . . . . . . guttula Motschulsky 

6’. Penultimate antennal segment with 

outer edge deeply and abruptly emarginate 

(Fig. 95); basal segment of antennal club 

black. Elytron with anterior black band 

reaching epipleural ridge but not crossing 

onto epipleuron (Fig. 89) . . obscurus Kirby 

7. Elytral epipleuron unicolorous, black or 

orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 

7’. Elytral epipleuron bicolored, black and 

orange (Fig. 88) . . mexicanus Matthews 

8. Elytral epipleuron black . . . . . . . . . 9 

8’. Elytral epipleuron orange . . . . . . . . . 10 

9. Dorsal surface of elytra with long, fine, 


erect setae (Fig. 90). Epipleural ridge short, 

extending anteriorly only to level of apex of 

scutellum . . . . . . . . . . . . . . orbicollis Say 

9’. Dorsal surface of elytra without long, 

erect setae (Fig. 91). Epipleural ridge long, 

extending anteriorly to near level of base of 

scutellum . . . . . . . . pustulatus Herschel 

10. Humerus covered by field of small, stout 

setae that extend to base of epipleural ridge 

(Figs. 85, 92). Metasternum with crescentshaped 

area immediately behind each mesocoxa 

that is glabrous or with sparse setae 

only. Posterior edge of metatrochanter of 

males with acute tooth projecting perpendicular 

(dorsally) from trochanter (Fig. 96). 

Females with small eyes, post-ocular bulge a 

little shorter than length of eye (Fig. 98) . . . . 

. . . . . . . . . . . . . hybridus Hatch & Angell 

10’. Humerus with small, stout setae but 

these ending well before base of epipleural 

ridge (Figs. 86, 93). Metasternum with long, 

dense setae immediately behind each mesocoxa. 

Posterior edge of metatrochanter of 

males with acute tooth recurving dorsally 

(Fig. 97). Females with large eyes, post-ocular 

bulge less than half length of eye (Fig. 99) 

. . . . . . . . . . . . . investigator Zetterstedt 

Fi g s. 76-83. Pronotum of 76, Nicrophorus carolinus; 77, N. guttula; 78, N. hybridus; 79, N. investigator; 

80, N. marginatus; 81, N. mexicanus; 82, N. orbicollis; 83, N. pustulatus.

56 


Fig s. 84-87. Lateral view of left side of elytra showing form and color of epipleuron and elytra, 

metepimeron, and humeral setae of 84, Nicrophorus guttula; 85, N. hybridus; 86, N. investigator; 

87, N. marginatus.


Fig s. 88-91. Lateral view of left side of elytra showing form and color of epipleuron and elytra, 

metepimeron, and humeral setae of 88, Nicrophorus mexicanus; 89, N. obscurus; 90, N. orbicollis; 

91, N. pustulatus.

58 


Fi g s. 92-93. Left elytron of 92, Nicrophorus hybridus 

and 93, N. investigator showing length of row of setae 

on humerus. 

Fig s . 94-95. Club of antenna of 94, Nicrophorus 

guttula and 95, N. obscurus. Note emargination of 

penultimate segment. 

Fi g s. 96-97. Caudal view of metatrochanter of males 

of 96, Nicrophorus hybridus and 97, N. investigator. 

Note form of tooth. 

Fi g s. 98-99. Dorsal view of head of 98, Nicrophorus 

hybridus female and 99, N. investigator female. Note 

size of post-ocular bulge. Males of both species are 

similar to Fig. 98.

KEY TO THE SPECIES OF THIRD 

INSTAR LARVAE 

OF NICROPHORUS IN NEBRASKA 

(modified from Anderson 1982) 

1. Abdominal segment 10 with venter 

unsclerotized at base. Abdominal segment 

9 with sclerotization of sternite fragmented 

(Fig. 102) or else truncated at lateral margins 

(Figs. 100, 103) . . . . . . . . . . . . . . . . . . . . . 2 

1’. Abdominal segment 10 with venter 

sclerotized at its base. Abdominal segment 

9 with sclerotization of the sternite entire, 

straight along posterior margin, arcuate and 

emarginate at middle along anterior margin 

(Fig. 101) . . . . . . . . . . . . . . . . . . . . . . . . . . 4 

2. Abdominal segment 9 with sclerotization 

of sternite fragmented (Fig. 102) . . . . . . 

. . . . . . . . . . . . . . . . . . . . marginatus Fabr. 

2’. Abdominal segment 9 with sclerotization 

of sternite entire and truncated along 

lateral margins (Figs. 100, 103) . . . . . . . 3 

3. Abdominal segment 3 with lateral and 

mid-dorsal spines equal in length (Fig. 103). 

Ligula without lateral lobes . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . obscurus Kirby 

3’. Abdominal segment 3 with mid-dorsal 

spines about 3 times length of lateral spines 

(Fig. 100). Ligula with small lateral lobes . . 

. . . . . . . . . . . . . . . . . guttula Motschulsky 

4. Abdominal segment 10 with y-shaped 

sclerite on venter in apical half, sclerite extending 

to bases of innermost pair of apical 

setae . . . . . . . . investigator Zetterstedt 

4’. Abdominal segment 10 not sclerotized 

on venter in apical half . . . . . . . . . . . . . . . 5 

5. Abdominal segment 1 with lateral and 

mid-dorsal spines equal in length. Dorsal 

spines at middle of segments 2-8 at least 

twice as long as lateral spines. Lateral spines 

of segments 1-8 short, not exceeding diameter 

of spiracle . . . . . tomentosus Weber 

5’. Abdominal segments 1-3 with lateral 

spines slightly longer than or equal to middorsal 

spines. Mid-dorsal spines of segments 

4-8 at least twice as long as lateral 

spines. Lateral spines of segments 1-8 long, 

approximately twice diameter of spiracle on 

segments 6-8 . . . . . . . . . . . . . . . . . . . . . . . .6 


6. Abdominal segment 9 with large lateral 

spines. Distance between the base of 

urogomphus and base of lateral spine subequal 

to length of lateral spine . . . . . . . . . 

. . . . . . . . . . . . hybridus Hatch & Angell 

6’. Abdominal segment 9 with minute 

lateral spines, appearing as small cones. 

Distance between base of urogomphus and 

base of lateral spine at least 4 times length 

of lateral spine . . . . . . . . orbicollis Say 

Fi g s. 100-103. Ventral view of ninth abdominal segment 

of larva of 100, Nicrophorus guttula; 101, N. 

investigator; 102, N. marginatus; 103, N. obscurus 

(after Anderson and Peck 1985).

60 


Nicrophorus americanus Olivier 

(Figs. 104-107) 

Nicrophorus americanus Olivier 1790: 6. 

Nicrophorus virginicus Frölich 1792: 123. 

Necrophorus grandis Fabricius 1801: 247. 


Frons orange. Club of antenna orange. 

Thorax: Pronotum orbicular. Lateral and 

basal margins broad, black. Disc orange; 

anterior transverse impression distinct. 

Metasternum with dense, yellow pubescence. 

Metepim-eron with sparse, light brown setae. 

Elytra: Each elytron with two transverse, orange 

maculae, maculae not reaching suture. 

Epipleuron completely orange. Legs: Poste- 

Fig. 104. Nicrophorus americanus Olivier. 

rior tibia slightly curved. Tarsal empodium 

quadrisetose. 

Distribution. Nicrophorus americanus was 

formerly distributed throughout 35 states 

and three Canadian provinces in temperate 

eastern North America from Nova Scotia 

to western Nebraska and from the upper 

peninsula of Michigan to Texas (U.S. Fish 

and Wildlife Service 1991). During this 

century, it has disappeared from over 90% 

of its historic range (Lomolino et al. 1995) 

(Fig. 105). It is now known from only five 

states: on Block Island off the southern coast 

of Rhode Island (Kozol 1989, 1991), eastern 

Oklahoma/western Arkansas (U.S. Fish 

and Wildlife Service 1991), the Sand Hills

in north-central Nebraska (Ratcliffe and 

Jameson 1992, Ratcliffe on-going studies), 

and, as of late 1995, southern South Dakota 

(Backlund and Marrone 1995). 

Anderson (1982a) suggested that N. 

americanus was an obligate denizen of primary 

forest, and that the coincident decline 

of this species and the destruction of this 

habitat were linked. Anderson suggested 

that dependence on larger vertebrate carcasses 

for breeding may have restricted N. 

americanus to ecosystems with deeper soils, 

hence mature forests with deep, humic soils. 

Lomolino et al. (1995) tested this hypothesis 

and found it to be false; N. americanus exhibited 

the widest niche breadth of the four most 

common Nicrophorus species present in their 

studies with the Arkansas and Oklahoma 

silphids. They found that N. americanus 

was broadly distributed across habitats, 

and that there was no preference for forest 

or shrub cover. The extant populations on 

treeless Block Island in Rhode Island and in 

the grassland areas of Ft. Chaffee Military 


Fig. 105. Present and historical distribution of Nicrophorus americanus. 

Reservation in Arkansas also do not support 

Anderson’s idea nor do the records from 

relatively treeless west-central Nebraska 

and South Dakota. Creighton et al. (1993) 

reported beetles occur both in oak-hickory 

forests and in grasslands in Oklahoma; 

higher numbers were observed in grasslands 

than in bottomland forests (where, presumably, 

foraging flight is severely hampered by 

denser undergrowth). Considering the broad 

geographic range formerly occupied by the 

American burying beetle, it is unlikely that 

vegetation or soil type were historically limiting 

(U.S. Fish and Wildlife Service 1991). 

Today, the American burying beetle seems 

to be largely restricted to areas most undisturbed 

by human influence. 

In Nebraska, the Sandhills is just such 

an area, and it is there that the beetles have 

been recently rediscovered. Gothenburg, 

Brady, North Platte, the Valentine National 

Wildlife Refuge, and Jamison are all locales 

in which beetles have been found during 

1994 and 1995. Mark Peyton, Senior District

62 


Biologist for the Central Nebraska Public 

Power and Irrigation District, made the 

first discovery of a substantial population 

near Gothenburg in 1994 when he collected 

40 specimens (Peyton 1994). During the 

summer of 1995, Peyton and Jon Bedick 

(my research assistant who is conducting 

studies on the Gothenburg population of 

the American burying beetle) captured more 

than 300 specimens. Contrary to the earlier 

belief that the insects were associated with 

eastern deciduous woodlands, it now seems 

that carrion availability (appropriate size as 

well as numbers) is more important than the 

type of vegetation or soil structure. Habitats 

in Nebraska where these beetles have been 

recently found consist of grassland prairie, 

forest edge, and scrubland (Ratcliffe 1995). 



ANTELOPE CO. (1): Neligh; CHERRY CO. 

(9): Valentine National Wildlife Refuge, 8 

mi. N Valentine National Wildlife Refuge; 

CUSTER CO. (1): Milburn; DAWSON CO. 

(173): Darr Strip Wildlife Management Area, 

Gallagher Canyon, 6 mi. S Gothenburg , 

Midway Lake; FRONTIER CO. (2): 3 mi. S 

Farnam; GOSPER CO. (3): Elwood; KEYA 

PAHA CO. (4): Jamison, Mills; LANCASTER 

Fig. 106. Nebraska distribution of Nicrophorus americanus. 

CO. (3): Lincoln; LINCOLN CO. (146): Box 

Elder Canyon, Brady, Cottonwood Canyon, 

9 mi. S Cozad, Jeffries Canyon, Moran Canyon, 

Snell Canyon, Wellfleet, North Platte; 



February to September (Peck and Kaulbars 

1987). Nebraska: April (1), June (93), July 

(107), August (96), September (2), October 

(2). One specimen from the Valentine National 

Wildlife Refuge was taken on 29 October 

1995. It was crawling in the grass, and 

the temperature was 42°F! Several inches of 

snow had fallen about two weeks earlier. 

Remarks. The American burying beetle, 

Nicrophorus americanus, is the largest (up to 

35 mm) carrion beetle in North America. It is 

easily distinguished from other orange banded 

species of Nicrophorus by its large size and 

by the orange pronotal disc. Males are easily 

distinguished by the large, orange rectangle 

on the clypeus whereas females have a small, 

orange triangle on the clypeus. 

Although all the immature stages are 

known, none appear to have been formally 

described in the literature. My student, Jon 

Bedick, and I will be rectifying this by describing 

the larval stage in the future.

U.S. Fish and Wildlife Service (1991) 

gives the most complete information regarding 

the ecology of this species. Adults 

are active at night (Schweitzer and Master 

1987, Peck and Kaulbars 1987), searching for 

carrion on which to feed and lay eggs; they 

are occasionally attracted to lights. These 

beetles seek out large carcasses (up to 300 g) 

of mostly birds and mammals, although carrion 

sources between 50 and 200 grams are 

apparently adequate for rearing their young 

(Fig. 107) (Schweitzer and Master 1987). 

Nicrophorus americanus, like many 

other Nicrophorus species, provides parental 

care for its young. Kozol, Scott, and Traniello 

(1988) demonstrated that carrion was 

prepared by the parents for the larvae in a 

fashion similar to that described for other 

Nicrophorus species. Nicrophorus americanus 

may cooperate in burying carrion, but individuals 

of both sexes are capable of burying 

carrion alone. The carrion is buried, shaved of 

fur or feathers, rolled into a ball, and treated 

with anal and oral secretions that favorably 


Fig. 107. Adult Nicrophorus americanus on a kangaroo rat in west central Nebraska. Photo by M. L. Jameson. 

alter the decay process. The female lays 

eggs in the soil near the carcass, and larvae 

hatch within a few days and move toward the 

carcass. The larvae are first fed regurgitated 

food by both the male and female parents. 

The larvae grow rapidly and are soon able to 

feed themselves. Approximately two weeks 

after burial of the carrion, the larvae complete 

their development and pupate in the 

soil nearby. Adults emerged from the pupal 

stage from 48 to 65 days later. 

They observed further that, in laboratory 

studies, reproductive success (measured 

by total brood weight and by the number of 

tenerals eclosed) is significantly correlated 

with carcass size as has also been shown 

in laboratory broods of N. orbicollis (Wilson 

and Fudge 1984). The negative correlation 

between the number of adults reared per 

brood and their average weight suggested 

that N. americanus parents make a tradeoff 

between a larger number of small offspring 

or a smaller number of large offspring. The 

results of this tradeoff may depend on carcass

64 


size, prior reproductive history of the parents, 

and possibly a “prediction” of future reproductive 

opportunities for the offspring. 

In the 1980s, entomologists documented 

the decreasing abundance of N. americanus 

across North America (Davis 1980, Anderson 

1982a, Kozol et al. 1988). Since that 

time, beetles of this species have not been 

collected or sighted in the United States 

except for the known and widely separated 

populations previously mentioned. Because 

of the precipitous decline in the distribution 

of this species, it was included in the IUCN 

Invertebrate Red Book as an endangered 

species (Wells et al. 1983). It was proposed as 

an endangered species in the United States 

Federal Register in 1988 (Recce 1988) and 

was placed on the Endangered Species List 

on 14 August 1989. 

Scott et al. (1987) noted that Nicrophorus 

species diversity is highest at northern 

latitudes, and it is likely that congeneric 

competition would be greatest where species 

diversity is highest. Kozol et al. (1988) 

observed that body size appears to be the 

most important determinant of success in 

competition for securing carrion; the largest 

individuals invariably displace smaller burying 

beetles. Because N. americanus are the 

largest carrion beetles in North America and 

even the smallest N. americanus overlap in 

size only slightly with the largest N. orbicollis 

and N. marginatus, it seems unlikely that 

N. americanus have been outcompeted by 

other Nicrophorus species. However, factors 

other than size that might affect the outcome 

of competition remain to be examined. 

Kozol et al. (1994) studied genetic variation 

within and between two populations of 

N. americanus, one from Rhode Island (Block 

Island) and the other from Oklahoma and 

Arkansas. They used the polymerase chain 

reaction RAPD-PCR with single short primers 

to randomly amplify polymorphic DNA. 

Comparable low levels of genetic variation 

were observed in both the island and distantly 

removed mainland populations. These 

populations were not differentiated, and this 

suggested that no genetic isolation had yet 

occurred as a result of the relatively recent 

geographic isolation. 

The prevailing theory explaining the 

disappearance of the American burying 

beetle involves habitat fragmentation. 

Fragmentation of large expanses of natural 

habitat changed the species composition and 

lowered the reproductive success of prey species 

required by the American burying beetle 

for optimum reproduction. Fragmentation 

also resulted in an increase in edge habitat 

that supported and increased the occurrence 

and density of vertebrate predators 

and scavengers such as crows, raccoons, 

foxes, opossums and skunks, all of which 

compete with burying beetles for available 

carrion. Fragmented habitats not only support 

fewer or lower densities of indigenous 

species that historically may have supported 

burying beetle populations, but there is also 

now a great deal more competition for those 

limited resources among the “new” predator/ 

scavenger community. 

Determining a single cause for the decline 

of the American burying beetle would 

simplify and facilitate its recovery. Unfortunately, 

the decline is probably the result of an 

interplay of several complex factors that may 

include (1) artificial lighting that decreases 

populations of nocturnally active insects, 

(2) changing sources of carrion because of 

habitat alteration, (3) isolation of preferred 

habitat because of land use changes, (4) 

increased edge effect harboring more vertebrate 

competitors for carrion and (5) the 

possibility of reduced reproduction because of 

some genetic characteristic of the species. 

The U.S. Fish and Wildlife Service, in 

cooperation with the scientific community, 

has formulated a recovery plan that is now 

being implemented. Surveys at several 

places in the United States are being conducted 

to find remnant populations so they 

can be protected. 

Beetles are being reintroduced in 

Massachusetts from a laboratory colony at 

Boston University, and other introductions 

are planned. Life history studies are being 

conducted in order to determine possible

factors responsible for the decline of the species. 

Similarly, DNA studies are ongoing to 

ascertain what, if any, genetic differences 

exist among the known populations. Knowledge 

of those differences could be important 

for future breeding programs. 

There is now an ongoing, unprecedented 

loss of species diversity throughout the world 

as well as a decline in the absolute numbers 

of organisms from the smallest microorganism 

to the largest mammal. The current 

loss of biota has several causes. One is the 

destruction, conversion, or degradation of 

entire ecosystems with the consequent loss 

of entire assemblages of species. Another 

is the accelerating loss of individual species 

within communities or ecosystems as a result 

of habitat disturbance, pollution, and overexploitation. 

Third and more subtle is the loss 

of genetic variability. Selective pressures 

such as habitat alteration, the presence of 

chemical toxins, or regional climate changes 

may eliminate some genetically distinct 

parts of the population yet not cause extinction 

of the entire species. 

Why worry about one insect that most 

of us have never seen? The World Wildlife 

Fund perhaps said it best: “All that lives 

beneath Earth’s fragile canopy is, in some 

elemental fashion, related. Is born, moves, 

feeds, reproduces, dies. Tiger and turtle 

dove; each tiny flower and homely frog; the 

running child, father to the man and, in ways 

as yet unknown, brother to the salamander. 

If mankind continues to allow whole species 

to perish, when does their peril also become 

ours?” 

Nicrophorus carolinus (Linnaeus) 

(Figs. 76, 108-109) 

Silpha carolina Linnaeus 1771: 530. 

Necrophorus mediatus Fabricius 1801: 334. 

Necrophorus mystacallis Angell 1912: 307. 

Necrophorus carolinus scapulatus Portevin 

1923: 142. 

Necrophorus carolinus dolosus Portevin 1923: 

307. 



Club of antenna completely orange. Thorax: 

Pronotum cordate, with lateral margins 

very narrow; basal margin moderately 

wide; anterior, transverse impression lacking 

(Fig. 76). Metasternum with dense, 

yellow pubescence. Metepimeron glabrous. 

Elytra: Epipleuron extremely narrow. Each 

elytron with two transverse, orange maculae; 

maculae variably reduced, anterior 

macula often broken into two spots. Legs: 

Posterior tibia slightly curved. 

Distribution. Nicrophorus carolinus occurs 

widely from the central United States 

south to Texas and Arizona, east along the 

Gulf Coastal Plain to Florida, and north 

along the Atlantic Coastal Plain to Virginia 

(Anderson and Peck 1985, Peck and Kaulbars 

1987). In Nebraska, this species is 

known from the western half of the state. 

One specimen was collected in Lincoln in 

1930, but no additional specimens have 

been taken since that time. 



ARTHUR CO. (31): Arapaho Prairie; CHASE 

CO. (1): Imperial; CHERRY CO. (40): Bloomington, 

Dewey Lake, Ft. Niobrara National 

Wildlife Refuge; CUSTER CO. (16): Anselmo, 

Milburn; DAWES CO. (1): Marsland; DAW- 

SON CO. (1): Gothenburg; DUNDY CO. 

(1): Haigler; FRONTIER CO. (1): Farnam; 

GARDEN CO. (1): Oshkosh; KEITH CO. (5): 

Cedar Point Biological Station; KEYA PAHA 

CO. (3): Mills, Norden; LANCASTER CO. 

(1): Lincoln; LINCOLN CO. (172): Box Elder 

Canyon, Brady, Cottonwood Canyon, Moran 

Canyon, Sutherland, Wellfleet; McPHER- 

SON CO. (1): Sandhills Ag Lab; NANCE CO. 

(1): Genoa; SCOTTS BLUFF CO. (10): Mitchell; 

SHERIDAN CO. (2): Gordon; THOMAS 

CO. (134): Halsey Forest, Thedford. 


March to October (Peck and Kaulbars 1987). 


August (224), September (8), October (1).

66 


Remarks. Nicrophorus carolinus is easily 

distinguished from all other North American 

species by the largely unsculptured 

pronotum (lacking a transverse, anterior 

impression and with very narrow explanate 

margins laterally) (Fig. 76). The elytral epipleuron 

is also narrow in comparison to other 

species of Nicrophorus. Anderson and Peck 

(1985) and Peck and Kaulbars (1987) noted 

that the elytral maculations are discontinuous 

and reduced in populations occurring in 

the north-central states, and nearly all the 

Nebraska material exhibits this trait. 

Fig. 108. Nicrophorus carolinus (L.). 

The immature stages, while undoubtedly 

known, have not been described in the 

literature. 

Anderson and Peck (1985), Peck and 

Kaulbars (1987), and Lingafelter (1995) 

remarked that N. carolinus occurs most 

frequently in areas with loose or sandy soils. 

The Nebraska specimens seem to corroborate 

this. Scott et al. (1987) found N. carolinus 

equally in fields and forests. Arnett (1946) 

described briefly how a dead snake was 

buried by these beetles, and Conley (1982) 

concluded that this species (at least in New

Mexico) had relatively low efficiency of 

carrion-locating behavior. I have collected 

this species commonly in western Nebraska 

using pitfall traps baited with fish. Otherwise, 

little is known of its biology. 

Nicrophorus guttula Motschulsky 

(Figs. 77, 84, 94, 100, 109-110) 

Necrophorus guttula Motschulsky 1845: 53. 

Necrophorus hecate Bland 1865: 382. 

Nicrophorus guttula punctostriatus Pierce 

1949: 66. 

Nicrophorus hecate immaculosus Hatch 1957: 

15. 


Club of antenna completely orange or with 

basal segment black or piceous; penultimate 

segment with outer edge emarginate (Fig. 

94). Thorax: Pronotum cordate, with lateral 

margins narrow; basal margin wide; anterior, 

transverse impression deep, distinct (Fig. 

77). Metasternum and metepimeron with 

dense, yellow pubescence. Elytra: Pattern 

variable; each elytron with two transverse, 

orange maculae, maculae frequently coalesced 

near center, anterior macula usually 

reaching suture, posterior macula not quite 

extending to suture; or, maculae reduced 


Fig. 109. Nebraska distribution of Nicrophorus carolinus, N. guttula, N. hybridus, N. investigator. 

to two separate bands, or spots, or entirely 

absent. Epipleuron usually orange with 

anterior black band of elytra crossing onto it 

(Fig. 84); in specimens with predominantly 

black elytra, only extreme base of epipleuron 

orange, remainder black. Legs: Posterior 

tibia straight. Anterior face of procoxa with 

long setae on basal half. 

Distribution. Nicrophorus guttula is widely 

distributed in the western half of the United 

States, southern British Columbia, Alberta, 

and Saskatchewan in Canada, and northern 

Baja California in Mexico (Anderson and 

Peck 1985, Peck and Kaulbars 1987). In Nebraska, 

this species is recorded from the west 

primarily, but there is a Lincoln record. 



BOX BUTTE CO. (1): No data; CHERRY CO. 

(7): Ft. Niobrara National Wildlife Refuge; 

DAWES CO. (6): Chadron; LANCASTER 

CO. (1): Lincoln; SIOUX CO. (56): Gilbert 

Baker Wildlife Area, Glen, Monroe Canyon, 

Warbonnet Canyon; THOMAS CO. (46): 

Halsey Forest Reserve. 

Temporal Distribution. Rangewide: May 

to September (Anderson and Peck 1985).

68 



August (15). 

Remarks. Nicrophorus guttula is similar 

to N. obscurus and is distinguished by the 

form of the third antennal segment (emarginate 

as opposed to the deeply emarginate 

segment of N. obscurus) (Figs. 94-95), the 

variable color of the first antennal segment 

(always black in N. obscurus), and the 

variably colored elytral epipleuron (always 

Fig. 110. Nicrophorus guttula Motschulsky. 

completely orange in N. obscurus). All 

but one of the Nebraska specimens have 

broad, orange elytral maculae. The single 

Lincoln specimen has completely black 

elytra, characteristic of southwestern 

coastal areas of the United States where 

melanistic forms occur. Nicrophorus guttula 

also resembles N. marginatus but is 

distinguished from it by the presence, in 

N. guttula, of long setae (as long or longer 

than those on humerus) on the anterior

face of the procoxa in the basal half; N. 

marginatus has short setae (shorter than 

those on humerus). 


Anderson (1982b), and a brief diagnosis 

was given by Anderson and Peck (1985). 


(1985), this species exhibits a broad range 

of ecological tolerances and inhabits dry 

forests, prairies, and deserts. Adults are 

diurnal, and they have been collected at 

human and coyote dung as well as carrion. 

In our studies done in Nebraska, numerous 

specimens have been taken using pitfall 

traps baited with the flesh of mice, rats, 

and rhinoceros. Yes, rhinoceros. 


Fig. 111. Nicrophorus hybridus Hatch and Angell. 

Nicrophorus hybridus Hatch and 

Angell 

(Figs. 78, 85, 92, 96, 98, 109, 111) 

Necrophorus hybridus Hatch and Angell 1925: 

216. 


Club of antenna with basal segment black, 

remaining three segments orange. Both 

sexes with small eyes; post-ocular bulge in 

female a little shorter than length of eye (Fig. 

98). Thorax: Pronotum with lateral and basal 

margins wide; anterior transverse impression 

deep, distinct (Fig. 78). Metasternum 

with dense, yellow pubescence except for a

70 


crescent-shaped, sparsely setose or nearly 

glabrous area immediately behind margin 

of each mesocoxa. Metepimeron glabrous. 

Elytra: Each elytron with 2 broad, complete, 

transverse, orange bands. Humerus 

covered by field of short, stout setae that 

extend posteriorly to base of epipleural ridge 

(Figs. 85, 92). Epipleuron orange (Fig. 85). 

Legs: Anterior face of procoxa with sparse, 

minute setae on basal half. Posterior tibia 

straight. Posterior edge of metatrochanter 

with acute tooth projecting perpendicular 

from trochanter (Fig. 96). 

Distribution. Nicrophorus hybridus ranges 

from the southern part of western Canada 

southward through the north-central United 

States to northern Arizona and New Mexico 


1987). In Nebraska, this species is known 

from the northern portion of the state only 

but may also occur in the panhandle. 


specimens examined. 

CUMING CO. (2): West Point; SIOUX CO. 

(1): Warbonnet Canyon. 

Temporal Distribution. Rangewide: June 

to September (Peck and Kaulbars 1987). 

Nebraska: No data. 

Remarks. Nicrophorus hybridus is best identified 

by the combination of key characters, 

especially when trying to separate it from N. 

investigator. Nicrophorus hybridus is distinctive 

because of its broad elytral maculations, 

crescent-shaped region immediately behind 

each mesocoxa that is glabrous or sparsely 

setose, and stout humeral setae that extend 

to the base of the epipleural ridge. Both the 

males and females have small eyes. The posterior 

edge of the metatrochanter in the males 

has an acute tooth that projects perpendicular 

to the plane of the trochanter (Fig. 96). Contrast 

this with Fig. 97 for N. investigator. 

The larval stage was described by Anderson 

(1982), and a diagnosis was given by 

Anderson and Peck (1985). 

Little is known of the biology of this 

species. Peck and Kaulbars (1987) characterized 

their habitat as prairie, sage steppe, and 

montane meadow. Adults are probably diurnal 

and are reproductively active during the 

summer (Anderson and Peck 1985). Overwintering 

occurs in the prepupal stage. 

Nicrophorus investigator Zetterstedt 

(Figs. 17, 79, 86, 93, 97, 99, 101, 107, 110) 

Necrophorus investigator Zetterstedt 1824: 154. 

Necrophorus maritimus Guérin-Méneville 

1835: Pl. 17, Fig. 8. 

Necrophorus melsheimeri Kirby 1837: 97. 

Necrophorus particeps Fischer von Waldheim 

1844: 139. 

Necrophorus aleuticus Gistel 1848: 190. 

Necrophorus pollinctor Mannerheim 1853: 169. 

Necrophorus infodiens Mannerheim 1853: 170. 

Necrophorus confossor LeConte 1854: 20. 



remaining segments orange. Male with small 

eyes, post-ocular bulge subequal to length 

of eye. Female with large eyes, post-ocular 

bulge less than half length of eye (Fig. 99). 

Thorax: Pronotum subquadrate, with lateral 

and basal margins wide; anterior, transverse 

depression deep (Fig. 79). Metasternum with 

dense yellow pubescence, including area just 

posterior of each mesocoxa. Metepimeron glabrous. 

Elytra: Pattern variable; each elytron 

with two broad, transverse, orange bands; or 

anterior band reduced to one, two, or three 

spots. Epipleuron orange (Fig. 86). Legs: 

Anterior face of procoxa with minute setae on 

basal half. Posterior tibia straight. Posterior 

edge of metatrochanter of males with acute 

tooth recurving dorsally (Fig. 97). 

Distribution. Nicrophorus investigator is 

widely distributed throughout Canada and 

Alaska and along the Rocky Mountains to 

New Mexico and Arizona. It is occasionally 

found in the northeastern United States. It 

is also broadly distributed in Europe and 

Asia (Anderson and Peck 1985, Peck and

Kaulbars 1987). In Nebraska, this species 

is known only from North Platte, and this 

locality represents a NEW STATE RECORD. 

These specimens were prepared by me from 

samples collected in a stand-alone light trap 

in 1966. Unfortunately, no specimens have 

been collected since 1966 even though recent 

trapping programs have been conducted both 

to the immediate east and west of North 

Platte. 


specimens examined. 

LINCOLN CO. (2): North Platte. 


Fig. 112. Nicrophorus investigator Zetterstedt. 


May to October (Peck and Kaulbars 1987). 

Nebraska: July (2). 

Remarks. Nicrophorus investigator is most 

reliably identified by the combination of key 

characters, but especially by the length of 

the field of small setae on the humerus when 

differentiating it from N. hybridus. Although 

variation in elytral pattern is extensive (with 

darker forms occurring in the northwestern 

coastal populations), all the specimens from 

Nebraska, Wyoming, and Colorado have 

broad bands of orange on the elytra.

72 



Anderson (1982) and a brief synopsis was 

given by Anderson and Peck (1985). 

Most of the work on the biology of this 

species has been conducted by Katakura and 

Fukuda (1975) in Japan, Pukowski (1933) 

and Mroczkowski (1949) in Europe, and 

Smith and Heese (1995) in the United States. 

According to these authors, adults appear 

in June or July and begin reproductive activities. 

Their later emergence in the year is 

reflective of their more northerly distribution 

or at greater elevations where summer-like 

conditions are brief. Overwintering occurs 

in the prepupal stage. 

Smith and Heese (1995), working at 

2,900 m in the Colorado Rockies, found a 

distinct preference by these beetles for particular 

carcasses based on size. They rejected 

carcasses less than 16 g or larger than 55 g. 

The beetles apparently “weigh” carcasses to 

determine size by crawling under them and 

experimentally lifting them while on their 

backs beneath the carcass. Results from this 

study indicated that choice of a carcass of a 

certain minimum size had important fitness 

consequences because beetles that buried 

small carcasses did not raise any larvae, 

and average brood mass was larger on larger 

carcasses. 

Smith and Heese (1995) also found that 

carcasses located in shady, hence cooler, 

habitats were not utilized as frequently as 

those in sunnier habitats. Larvae reared in 

shaded conditions developed more slowly. 

While this could cause negative fitness for 

the larvae (if they don’t develop sufficiently 

before winter), these authors propose that 

slowed development may have negative 

effects on the total reproductive output 

of brood-tending parents by reducing the 

number of broods they can rear in a season. 

Hines and Smith (1995), as part of the same 

study, found that the average elytral length 

was significantly longer at higher elevations 

in the Rocky Mountains. 

Lane and Rothschild (1965) reported 

that N. investigator is a very good visual 

and sound mimic of the bumblebees Bombus 

lucorum (L.). Beetles presumably acquire 

some degree of protection from predators by 

mimicking a large, stinging bee. 

Nicrophorus marginatus Fabricius 

(Figs. 65-66, 80, 87, 102, 113-114) 

Necrophorus marginatus Fabricius 1801: 334. 

Necrophorus requiescator Gistel 1848: 190. 

Necrophorus montezumae Matthews 1888: 92. 

Necrophorus marginatus cordiger Portevin 

1924: 84. 

Nicrophorus guttula labreae Pierce 1949: 63. 

Nicrophorus mckittricki Pierce 1949: 66. 

Nicrophorus obtusiscutellum Pierce 1949: 67. 

Nicrophorus investigator latifrons Pierce 1949: 

67. 


Club of antenna completely orange. Thorax: 

Pronotum subtrapezoidal, widest anteriorly, 

with lateral margins narrow; basal margin 

wide; anterior, transverse impression present, 

occasionally only weakly indicated 

(Fig. 80). Metasternum with dense, yellow 

pubescence. Metepimeron with dense, yellow 

pubescence. Elytra: Each elytron with 

two transverse, orange maculae, anterior 

orange macula nearly always reaching median 

suture while posterior macula does 

not, both maculae usually joined laterally; 

maculae rarely reduced to spots. Epipleuron 

completely orange (Fig. 87). Legs: Posterior 

tibia slightly curved. Anterior face of procoxa 

with short setae on basal half. 

Distribution. Nicrophorus marginatus occurs 

throughout most of the United States 

(excluding Florida and northwestern Washington), 

across southern Canada, and into 

northern Mexico (Anderson and Peck 1985, 

Peck and Kaulbars 1987). It is the most 

widely distributed of the North American 

Nicrophorus. It is found across the entire 

state of Nebraska. 


specimens examined or recorded.

ADAMS CO. (30): No data; ARTHUR CO. 

(15): Arapaho Prairie; BOX BUTTE CO. (4): 

No data; BUFFALO CO. (6): Elm Creek, 

Kearney; BUTLER CO. (1): David City; 

CASS CO. (260): Plattsmouth; CHASE CO. 

(3): Enders Reservoir; CHERRY CO. (77): 

Dewey Lake, Sparks, Valentine, Ft. Niobrara 

National Wildlife Refuge; CHEYENNE CO. 

(1): Sidney; CLAY CO. (2): No data; CUM- 

ING CO. (1): West Point; CUSTER CO. 

(491): Anselmo, Milburn, Sargent; DAKOTA 

CO. (1): South Sioux City; DAWES CO. (9): 

Ash Creek, Chadron; DIXON CO. (5): Aowa 

Creek, Concord; DUNDY CO. (3): Haigler, 

Republican River east of Benkelman; FILL- 

MORE CO. (1): Fairmont; FRANKLIN CO. 

(3): Bloomington; FRONTIER CO. (597): Curtis, 

Farnam, Medicine Creek Reservoir, Red 

Willow Reservoir; GAGE CO. (1): Beatrice; 

GOSPER CO. (274): Elwood Reservoir, Gothenburg, 

Lexington, Smithfield; GREELEY 


Fig. 113. Nicrophorus marginatus Fabr. 

CO. (2): Greeley; HALL CO. (54): Alda, Grand 

Island; JEFFERSON CO. (33): No data; 

JOHNSON CO. (4): No data; KEITH CO. 

(21): Cedar Point Biological Station; KEYA 

PAHA CO. (7): Mills, Norden; KNOX CO. 

(9): Bazile Creek Wildlife Management Area, 

Niobrara; LANCASTER CO. (38): Lincoln, 

Sprague; LINCOLN CO. (3,332): Box Elder 

Canyon, Brady, Cottonwood Canyon, Moran 


MERRICK CO. (1): Central City; NANCE 

CO. (6): Genoa; OTOE CO. (14): Nebraska 

City; PAWNEE CO. (5): No data; PHELPS 

CO. (231): Bertrand; PIERCE CO. (1): No 

data; POLK CO. (10): No data; RED WIL- 

LOW CO. (1): McCook; RICHARDSON CO. 

(1): Verdon; SALINE CO. (4): Swan Creek; 

SARPY CO. (1): Bellevue; SAUNDERS 

CO. (2): Wahoo; SCOTTS BLUFF CO. (43): 

Mitchell, Scottsbluff; SHERMAN CO. (1): 

Loup City; SIOUX CO. (59): Gilbert Baker

74 


Wildlife Area; THOMAS CO. (60): Halsey 

Forest Reserve; WASHINGTON CO. (2): Ft. 

Calhoun; WEBSTER CO. (1): Red Cloud; 

YORK CO. (1): No data. 


February to October (Peck and Kaulbars 

1987). Nebraska: April (5), May (11), June 

(230), July (1,944), August (2,221), September 

(296), October (270), November (3). The 

large numbers in July and August reflect, 

in part, a concerted trapping program in 

Lincoln County in 1995. 

Remarks. Nicrophorus marginatus cannot 

be separated from N. obscurus and N. 

guttula based on overall appearance. It can 

be separated from these other two species 

because it has short setae (shorter than those 

on humerus) on the anterior face of the procoxa 

in the basal half; the two other species 

have long setae (as long or longer than those 

on humerus). 


(1982b), and a brief larval diagnosis 

was given by Anderson and Peck (1985). 

Adults become active in the spring 

when the weather is consistently warm; most 

adults become active in June in Nebraska. 

Reproduction occurs in June and July, and 

the new generation of adults appears in 

July and August. Overwintering is in the 

adult stage. According to Peck and Kaulbars 

(1987), most collections have been from 

open fields, montane meadows, prairies, and 

desert woodlands. This species is abundant 

in the grasslands of western Nebraska. Lomolino 

et al. (1995) and Lingafelter (1995) 

demonstrated that this species shows a 

strong preference for meadows or open, 

grassy areas. 

This species is probably diurnal even 

though some specimens have been taken 

at lights. I and my students have taken N. 

marginatus in large numbers using baited 

pitfall traps. Clark (1895) observed extensive 

feeding by adults on fly larvae at carrion. 

Nicrophorus mexicanus Matthews 

(Figs. 81, 88, 114-115) 

Necrophorus mexicanus Matthews 1888: 91. 



remaining segments orange. Thorax: 

Pronotum subquadrate, both anterior 

and posterior angles rounded. Lateral 

Fig. 114. Nebraska distribution of Nicrophorus marginatus, N. mexicanus, N. obscurus.

and basal margins wide; anterior transverse 

impression deep, distinct (Fig. 81). 

Metasternum with dense, dark brown 

pubescence. Metepim-eron with small, 

central patch of dark brown setae. Elytra: 

Each elytron with two transverse, orange 

maculae, anterior orange macula joined 

at suture, posterior macula long but not 

quite reaching suture. Epipleuron orange 

with anterior black band crossing onto it 

(Fig. 88). Legs: Posterior tibia straight. 

Anterior face of procoxa with minute setae 

on basal half. 

Distribution. Nicrophorus mexicanus was 

formerly known from the southern Rocky 


Fig. 115. Nicrophorus mexicanus Matthews. 

Mountain states southward through Mexico 

to El Salvador (Peck and Kaulbars). The 

specimen listed below from central Nebraska 

represents a NEW STATE RECORD 

and represents a range extension from the 

closest known current collection localities 

in central Colorado. 


specimen examined. 

CUSTER CO. (1): 17 mi. E. Anselmo. 

Temporal Distribution. Rangewide 

(including Mexico): All months. United 

States: May to October (Peck and Kaulbars 

1987). Nebraska: July (1).

76 


Remarks. Nicrophorus mexicanus adults 

are most easily distinguished by the dark 

setae on the metepimeron, subquadrate 

pronotum, and bicolored epipleuron. 

The larval stage has apparently not 

been described. 

Peck and Anderson (1985) recorded 

this species from habitats ranging from 

semi-arid and open thorn scrub to moist, 

closed-canopy cloud forests. The Nebraska 

specimen was collected in a baited pitfall 

trap from an area of scrub vegetation in the 

floodplain of the Loup River. 

Fig. 116. Nicrophorus obscurus Kirby. 

Nicrophorus obscurus Kirby 

(Figs. 89, 95, 103, 114, 116) 

Nicrophorus obscurus Kirby 1837: 97. 

Necrophorus melsheimeri LeConte (not Kirby) 

1853: 275. 



apical three segments orange; penultimate 

segment with outer edge deeply emarginate 

(Fig. 89). Thorax: Pronotum subtrapezoidal, 

widest anteriorly, with the lateral margins

narrow; basal margin wide; anterior, transverse 

impression present (as in Fig. 80). 

Metasternum and metepimeron with dense, 

yellow pubescence. Elytra: Each elytron 

with two transverse, orange maculae, anterior 

orange macula nearly always reaching 

suture, posterior macula not quite extending 

to suture; surface with sparse, short, 

slender, decumbent, black setae. Epipleuron 

completely orange (Fig. 89) Legs: Posterior 

tibia slightly curved. Anterior face of procoxa 

with long setae on basal half. 

Distribution. Nicrophorus obscurus occurs 

from the southern Canadian prairie provinces 

into the north-central United States as 

far south as Nebraska and northern Colorado 


1987). Nebraska records indicate this 

species is found statewide. 



CHERRY CO. (19): Ft. Niobrara National 

Wildlife Refuge; CUMING CO. (1): West 

Point; CUSTER CO. (5): Sargent; DAWES 

CO. (7): Pine Ridge area; FRONTIER CO. 

(42): Farnam, Medicine Creek Reservoir, Red 

Willow Reservoir; GOSPER CO. (10): Elwood 

Reservoir, Lexington, Smithfield; KEITH 

CO. (1): Cedar Point Biological Station; 

KEYA PAHA CO. (1): Carns; LANCASTER 

CO. (1): Lincoln; LINCOLN CO. (290): Box 


Canyon, Sutherland, Wellfleet; PHELPS 

CO. (1): Bertrand; SIOUX CO. (1): Harrison; 

THOMAS CO. (3): Halsey Forest Reserve. 


March to September (Peck and Kaulbars 

1987). Nebraska: June (3), July (180), August 

(193), September (4), October (1). 

Remarks. The deeply emarginate outer 

edge of the third antennal segment (Fig. 85) 

is particularly characteristic of this species. 

Nicrophorus guttula is similar in appearance, 

but has a simply emarginate third antennal 

segment (Fig. 84). Nicrophorus obscurus 


also closely resembles N. marginatus but is 

distinguished from it by the presence, in N. 

obscurus, of long setae on the anterior face of 

the procoxa in the basal half; N. marginatus 

has short setae. 


(1982b), and a brief diagnosis was 


Anderson and Peck (1982) observed 

that this is a diurnal prairie species, and 

that adults have been collected at carrion 

and human feces. I have taken this species 

in moderate numbers in western Nebraska 

using baited pitfall traps. 

Nicrophorus orbicollis Say 

(Figs. 82, 90, 117-118) 

Necrophorus orbicollis Say 1825: 177. 

Necrophorus halli Kirby 1837: 98. 

Necrophorus quadrisignatus Laporte 1840: 1. 



remaining segments orange. Thorax: 

Pronotum suborbicular, with lateral and 

basal margins broad; anterior, transverse 

impression deep, distinct (Fig. 82). Surface 

usually with setae along margins and 

in transverse impression laterally, these 

sometimes worn away. Metasternum 

with dense, light brown pubescence. Metepimeron 

with sparse, dark brown setae. 

Elytra: Elytra with long setae over entire 

surface, setae occasionally abraded away 

and present only along lateral margins (best 

seen in oblique lighting). Each elytron with 

an anterior, transverse, orange macula and a 

posterior, orange spot; markings not reaching 

suture. Epipleuron black; epipleural ridge 

extending anteriorly only to level of apex of 

scutellum (Fig. 90). Legs: Posterior tibia 

straight. Anterior face of procoxa with short 

setae on basal half. 

Distribution. Nicrophorus orbicollis is 

widely distributed in the eastern half of North 

America to southeastern Saskatchewan to 

eastern Texas (Anderson and Peck 1985,

78 


Peck and Kaulbars 1987). This species 

occurs throughout Nebraska. 



ADAMS CO. (2): No data; ANTELOPE CO. 

(1): Clearwater; BUFFALO CO. (28): Gibbon, 

Kearney; CASS CO. (232): Plattsmouth, 

South Bend; CHASE CO. (44): Enders 

Reservoir; CHERRY CO. (98): Ft. Niobrara 

Wildlife Refuge; COLFAX CO. (1): No data; 

CUMING CO. (1): West Point; CUSTER CO. 

Fig. 117. Nicrophorus orbicollis Say. 

(301): Anselmo, Ansley, Milburn, Sargent; 

DAWES CO. (57): Ash Creek, Chadron; 

DAWSON CO. (26): Cozad, Gothenburg, 

Lexington; DIXON CO. (316): Aowa Creek; 


River E of Benkelman; FRANKLIN 

CO. (83): Franklin; FRONTIER CO. (727): 

Farnam, Medicine Creek Reservoir, Red Willow 

Reservoir; GAGE CO. (38): Wolf-Wildcat 

Creek; GOSPER CO. (299): Elwood Reservoir, 

Gothenburg, Lexington, Smithfield; GRANT 

CO. (2): No data; HALL CO. (138): Alda,

Mormon Island; HARLAN CO. (17): Republican 

River S of Orleans; HOLT CO. (1): 

Spencer; HOOKER CO. (1): Mullen; JEF- 

FERSON CO. (16): No data; JOHNSON CO. 

(75): No data; KEITH CO. (61): Cedar Point 

Biological Station; KEYA PAHA CO. (7): 

Mills, Norden; KNOX CO. (45): Bazile Creek 

Wildlife Management Area; LANCASTER 

CO. (64): Lincoln, Reller Prairie, Sprague; 

LINCOLN CO. (1,142): Brady, Box Elder 

Canyon, Cottonwood Canyon, Moran Canyon, 

North Platte, 2 mi. S Sutherland, Wellfleet; 

McPHERSON CO. (1): No data; NUCK- 

OLLS CO. (1): No data; OTOE CO. (476): 

Nebraska City; PAWNEE CO. (706): No 

data; PHELPS CO. (27): Bertrand; PLATTE 

CO. (2): Columbus; RICHARDSON CO. (2): 

Indian Cave State Park; SALINE CO. (431): 

Swan Creek; SARPY CO. (612): Fontenelle 


(680): Mead, Wahoo; SIOUX CO. (35): Gilbert 

Baker Wildlife area, Monroe Canyon; 

THOMAS CO. (165): Halsey Forest Reserve; 

WASHINGTON CO. (629): Ft. Calhoun. 


February to October (Peck and Kaulbars 

1987). Nebraska: May (58), June (432), July 

(2,349), August (5,188), September (597), 

October (214). 

Remarks. Nicrophorus orbicollis is the 

most abundant and commonly encountered 

species of Nicrophorus in the state. The long, 

elytral setae are diagnostic for this species, 

but they are occasionally abraided away and 

so caution should be used when identifying 

specimens. It most closely resembles N. 

mexicanus, but N. mexicanus has a bicolored 

epipleuron whereas N. orbicollis has a black 

epipleuron. 


Anderson (1982), and a brief larval diagnosis 

was provided by Anderson and Peck 

(1985). 

Shubeck (1984a, 1993) and Lomolino 

et al. (1995), in studies in New Jersey, Oklahoma 

and Arkansas, demonstrated that this 

species seems to show a preference for for- 


ested areas. Contrary to these studies, this 

species is found abundantly in the grasslands 

of western Nebraska as well as the forested 

areas of eastern Nebraska. Lingafelter (1995) 

observed a similar pattern in Kansas. Nicrophorus 

orbicollis is the most commonly collected 

species in eastern North America (Peck 

and Kaulbars 1987) and seems to be the most 

abundant species of Nicrophorus in Nebraska. 

Trumbo (1990d) reported that they are the 

dominant burying beetle on small carcasses 

in the woodlands of the eastern United States. 

Shubeck (1971) reported they are nocturnal 

and are often taken at light traps; this has 

been my experience also. 

Adults overwinter and become active in 

the spring at which time reproduction takes 

place. Nicrophorus orbicollis is successful on 

a higher proportion of carcasses of all sizes 

and gains substantial benefits by excluding 

rivals; it rarely tolerates consexuals in the 

nest (Trumbo 1995). Generally, the male 

parent remains in the nest 5-12 days and 

the female parent for 10-15 days (Scott and 

Traniello 1990b, Trumbo 1991). In another 

study (under natural conditions in the field), 

Scott (1990) found that N. orbicollis males 

remained with the brood a mean of 9.5 days 

and females 17.2 days; however, 22% of the 

broods were reared by single females. In 

her experiments examining male assistance 

in guarding against conspecifics, 87% of 

the males and 93% of the females were still 

present after eight days; females normally 

remain with the brood longer than eight days 

post-burial. This long period of parental care 

represents a considerable investment for 

an animal with a reproductive span of two 

months or less, especially in view of the fact 

that both parents are foregoing additional 

reproductive success while they remain 

with their current brood. Larvae completed 

development about seven days after hatching 

and dispersed to pupate in the soil, emerging 

as adults about 30 days later (Scott and 

Traniello 1990b). Adults of the new generation 

are found in late July and early August 

in Nebraska. Wilson et al. (1984) suggested 

that N. orbicollis appear to enter reproductive

80 


diapause in late summer before temperatures 

become too cold to find carrion. This is 

an adaptation to season length since only individuals 

that have enough time to reach the 

adult stage can successfully overwinter. 

Peck and Kaulbars (1987) noted that 

they have been taken on human and carnivore 

dung and on rotting fruits as well as 

on carrion. I and my students have taken 

them in large numbers in Nebraska in pitfall 

traps baited with rotting fish, beef liver, and 

chicken hearts and gizzards. Adults have 

been observed to feed on maggots (Clark 

1895, Steele 1927). 

Pronotum subquadrate, with lateral and 

basal margins wide; anterior, transverse 

impression distinct (Fig. 83). Metasternum 

with sparse, light brown pubescence. 

Metepim-eron glabrous. Elytra: Each 

elytron with a small to medium, orange 

spot on lateral edge at about middle and 

two small to medium-sized spots near 

apex; elytra lacking long, distinctive setae. 

Epipleuron entirely black; epipleural ridge 

extending anteriorly to near level of base of 

scutellum (Fig. 91). Legs: Posterior tibia 

straight. Anterior face of procoxa glabrous 

on basal half. 

Fig. 118. Nebraska distribution of Nicrophorus orbicollis, N. pustulatus, N. tomentosus. 

Nicrophorus pustulatus Herschel 

(Figs. 83, 91, 118-119) 

Necrophorus pustulatus Herschel 1807: 271. 

Necrophorus bicolon Newman 1838: 385. 

Necrophorus tardus Mannerheim 1853: 170. 

Necrophorus marginatus fasciatus Portevin 

1924: 86. 

Necrophorus marginatus unicolor Portevin 

1924: 86. 



remaining three segments orange. Thorax: 

Distribution. Nicrophorus pustulatus is 

found from southern Canada east of the 

Rocky Mountains and in the eastern half of 

the United States from North Dakota to eastern 

Texas (Anderson and Peck 1985, Peck and 

Kaulbars 1987). This species is known from 

the eastern two-thirds of Nebraska; collecting 

records are lacking for the panhandle. 


specimens examined or recorded. The 

large number for Saunders County is the 

result of an intensive pitfall survey during 

the summer of 1995.

ADAMS CO. (2): No data; CLAY CO. (1): No 

data; CUSTER CO. (5): Anselmo, Milburn; 

DAWSON CO. (4): No data; DOUGLAS CO. 

(1): Omaha; DUNDY CO. (1): Republican 

River E of Benkelman; FRANKLIN CO. (3): 

Franklin; FRONTIER CO. (11): Farnam, 

Medicine Creek Reservoir, Red Willow 

Reservoir; GOSPER CO. (3): 4 mi. S. Gothenburg, 

Lexington; HALL CO. (13): Alda, 

Mormon Island; HOLT CO. (1): Spencer dam; 

JEFFERSON CO. (20): Fairbury; JOHNSON 

CO. (10): No data; KEYA PAHA CO. (19): 

Mills, Norden; KNOX CO. (1): Bazile Creek 

Wildlife Management Area; LANCASTER 

CO. (5): Lincoln, Sprague; LINCOLN CO. 

(47): Box Elder Canyon, Brady, Cottonwood 

Canyon, Moran Canyon, North Platte, 


Fig. 119. Nicrophorus pustulatus Herschel. 

Sutherland, Wellfleet; OTOE CO. (1): No 

data; PAWNEE CO. (7): No data; PHELPS 

CO. (7): Bertrand; RICHARDSON CO. (11): 

Indian Cave State Park; SALINE CO. (8): 

Swan Creek; SARPY CO. (6): Bellevue; 

SAUNDERS CO. (295): Wahoo; SEWARD 

CO. (1): Garland; THOMAS CO. (4): Halsey 

Forest Reserve; WAYNE CO. (1): Wayne. 

Temporal Distribution. Rangewide: March 

to October (Peck and Kaulbars 1987). Nebraska: 

April (1), May (3), June (42), July (39), 

August (402), September (3), October (3). 

Remarks. Nicrophorus pustulatus is easily 

recognized by its primarily black color and 

the presence of small elytral spots rather than

82 


transverse bands. Some specimens of N. 

orbicollis may also have small spots, but 

then they also possess elytral setae whereas 

N. pustulatus do not. 

The larval stage remains undescribed. 

Wilson and Knollenberg (1984) and 

Anderson and Peck (1985) observed that 

N. pustulatus is one of the rarer species of 

Nicrophorus and that it may, in fact, have a 

different natural history than other species 

as exemplified by its rarity in pitfall traps, 

absence from mice carcasses, and common 

occurrence at lights. Trumbo (1992) observed 

that N. pustulatus is a formidable 

brood parasite and produces the largest 

clutches (nearly 200 young) of any Nicrophorus 

species. They are routinely able to 

parasitize the broods of N. orbicollis but 

the reverse was never observed (Trumbo 

1994). Adults are nocturnal. Adult activity, 

including reproduction, occurs in the 

spring, and teneral adults usually appear 

in mid to late summer. These adults probably 

overwinter (Peck and Kaulbars 1987). 

Anderson (1982) and Shubeck (1983) suggested 

this species has a strong preference 

for forested habitats while Lingafelter’s 

(1995) study in Kansas showed a preference 

for the ecotone between forests and fields. 

Nicrophorus tomentosus Weber 

(Figs. 118, 120) 

Necrophorus tomentosus Weber 1801: 47. 

Necrophorus velutinus Fabricius 1801: 334. 

Necrophorus velutinus angustifasciatus 

Portevin 1925: 170. 

Necrophorus velutinus aurigaster Portevin 

1925: 170. 


Club of antenna black, basal segment shining, 

remaining segments dull. Thorax: 

Pronotum subquadrate with lateral margins 

broad; basal margin wide; surface covered 

with dense, long, yellow setae. Metasternum 

with long, yellow setae, with a glabrous spot 

present posterior to each of the mesocoxae. 

Metepimeron with only a few yellow setae 

or glabrous. Elytra: Each elytron with two, 

transverse orange maculae, maculae usually 

reaching suture, occasionally connected laterally, 

occasionally coalesced on disc. Legs: 

Posterior tibia straight. 

Distribution. Nicrophorus tomentosus is an 

abundant and widely distributed species. It 

occurs in nearly all of the United States (not 

the southern halves of Texas or Florida) and 

southern Canada east of the Rocky Mountains 

(Anderson and Peck 1985, Peck and 

Kaulbars 1987). In Nebraska, it is found 

throughout the state. 



ADAMS CO. (3): Hastings; CASS CO. (166): 

Plattsmouth; CHASE CO. (76): Enders 

Reservoir; CHERRY CO. (20): Valentine, 

Ft. Niobrara Wildlife Refuge; CHEYENNE 

CO. (4): Dalton, Gurly; CUMING CO. (3): 

West Point; CUSTER CO. (250): Anselmo, 

Milburn, Sargent; DAWES CO. (11): Ash 

Creek, Chadron; DAWSON CO. (2): 5 mi. S. 

Gothenburg; DIXON CO. (192): Aowa Creek; 


River E of Benkelman; FILLMORE 

CO. (5): Fairmont; FRANKLIN CO. (19): 

Franklin; FRONTIER CO. (1,287): Farnam, 

Medicine Creek Reservoir, Red Willow Reservoir; 

GOSPER CO. (192): Elwood Reservoir, 

Lexington, Smithfield; HALL CO. (37): 

Alda; HARLAN CO. (6): Republican River S 

of Orleans; JEFFERSON CO. (36): No data; 

JOHNSON CO. (5): No data; KEITH CO. 

(10): Cedar Point Biological Station, Sand 

Creek at Hwy. 2; KEYA PAHA CO. (50): 

Carns, Mills, Norden; KNOX CO. (72): Bazile 

Creek Wildlife Mgmt. Area, Center; LAN- 

CASTER CO. (46): Lincoln, Reller Prairie, 

Sprague; LINCOLN CO. (2,719): Brady, Box 



McPHERSON CO. (1): No data; OTOE CO. 

(26): No data; PAWNEE CO. (6): No data; 

PHELPS CO. (5): Bertrand; PLATTE CO. 

(1): Columbus; POLK CO. (10): No data; 

RICHARDSON CO. (1): Indian Cave State

Park; SALINE CO. (4): Swan Creek; SARPY 

CO. (28): Bellevue, Fontenelle Forest, Schramm 

Park; SAUNDERS CO. (597): Wahoo; 

SIOUX CO. (53): Gilbert Baker Wildlife area, 

Glen, Monroe Canyon; THOMAS CO. (64): 

Halsey Forest; WASHINGTON CO. (13): Ft. 

Calhoun; YORK CO. (10): No data. 


May to October (Peck and Kaulbars 1987). 

Nebraska: June (186), July (3,157), August 

(2,413), September (182), October (98). The 

high numbers for July and August reflect, 

in part, extensive trapping programs that 

were carried out in Frontier and Lincoln 

counties during the summer of 1995. 


Fig. 120. Nicrophorus tomentosus Weber. 

Remarks. Nicrophorus tomentosus is readily 

separated from all other silphids in North 

America by the presence of long, dense, yellow 

setae covering the pronotum. 


(1982b), and a brief diagnosis was 



(1985), this species is unlike other species 

of Nearctic Nicrophorus in that adults do 

not bury the carcass. Instead, they make 

only a shallow pit into which the carcass 

sinks. It is then covered with litter. After 

a period of feeding, mature larvae move into 

the surrounding soil where they spend the 

winter as a third instar, prepupal larva.

84 


Pupation occurs the following spring, and 

adults emerge in June in Nebraska. Scott 

and Traniello (1990b) indicated they do not 

become reproductively active until August. 

Wilson et al. (1984) suggested that N. tomentosus 

does not reproduce immediately 

because of competition from larger species of 

Nicrophorus. Smaller, “subordinate” species, 

such as N. tomentosus, can persist by restricting 

themselves to activity periods when 

more dominant species are not abundant or 

by using resources for which interference is 

unprofitable for the more dominant (larger) 

species (Trumbo 1990b). 

Adults are diurnal (personal observation, 

Shubeck 1971, Wilson et al. 1984). They 

greatly resemble bumble bees when flying, 

and Milne and Milne (1944) and Fisher and 

Tuckerman (1986) suggested they are Batesian 

mimics of bumble bees. Bumble bees are 

strictly diurnal, and one would expect that 

they are mimicked only by diurnal carrion 

beetles; this appears to be the case. Clark 

(1895) and Steele (1927) observed adults 

feeding on maggots at carrion. 

Anderson (1982c), Lingafelter (1995), 

and Lomolino et al. (1995) observed that N. 

tomentosus is a habitat generalist with no 

preference for forests, shrubby areas, or open 

grasslands. Our experience in Nebraska 

corroborates these observations. 

ACKNOWLEDGMENTS 

As with all synoptic faunal works of 

this nature, a great many people contributed 

valuable assistance to enable its completion. 

I thank Mark Marcuson (former Scientific 

Illustrator at the University of Nebraska 

State Museum) for most of the habitus drawings 

of the beetles, Polly Denham (Scientific 

Illustrator for the Museum) for some of the 

habitus drawings, David Reiser (Waverly, 

NE) for the highly detailed color cover, and 

my colleague and research assistant, Mary 

Liz Jameson, for the many fine line drawings 

and maps. Charles Messenger (Collection 

Manager, University of Nebraska State 

Museum) and Mary Liz assisted in collating 

data as well as with collecting specimens. 

Jon Bedick (currently my graduate student 

working on the American burying beetle) and 

Mark Peyton (Senior District Biologist with 

the Central Nebraska Public Power and Irrigation 

District) added substantially to the 

database with their collecting activities in 

west central Nebraska. 

Over the past several years, I have 

conducted endangered species surveys for 

the American burying beetle for several 

agencies and firms. I thank the following 

for providing additional opportunities to 

sample silphids: U.S. Fish and Wildlife 

Service, U.S. Forest Service, U.S. Army 

Corps of Engineers, Natural Resources Conservation 

Service (USDA), Nebraska Dept. 

of Roads, Omaha Public Power District, 

Nebraska Public Power District, Black and 

Veatch Engineering, Peterson Environmental 

Con- sulting, Werner Construction Co., 

and Fontenelle Forest Association. Mary 

Liz Jameson, Charles Messenger, Daniel 

Schmidt (Schuyler, NE), Natalie Sunderman 

(Lincoln, NE), and Steven Lingafelter (University 

of Kansas) conducted or assisted with 

some of the surveys. I extend special thanks 

to Wally Jobman (Wildlife Biologist, U.S. 

Fish and Wildlife Service) and Mike Fritz 

(Nebraska Game and Parks Commission) 

for their continuing support of my activities 

with endangered species and for coordinating 

these activities with the respective federal 

and state guidelines governing them. 

I thank the Center for Great Plains 

Studies (University of Nebraska) for granting 

me a Summer Faculty Fellowship to conduct 

research on Nebraska’s carrion beetles. 

Charles Springer (Hastings College), Randall 

Lawson (Chadron State College), Hal Nagel 

(University of Nebraska-Kearney), and John 

Janovy (Cedar Point Biological Station) made 

available specimens for study that were in 

collections under their care. I am grateful 

to Gail Littrell (University of Nebraska State 

Museum) for typing and preparing the manuscript. 

I thank Mary Liz Jameson, Mark 

Peyton, and Steven Lingafelter for critically 

reviewing the manuscript.

LITERATURE CITED 

ABBOTT, C.E. 1927a. Experimental data 

on the olfactory sense of Coleoptera, 

with special reference to the Necrophori. 

Annals of the Entomological Society of 

America 20: 207-216. 

ABBOTT, C.E. 1927b. Further observations 

on the olfactory powers of 

the Necro-phori. Annals of the Entomological 

Society of America 20: 550-553. 

ABBOTT, C.E. 1936. On the olfactory powers 

of a necrophilous beetle. Bulletin of 

the Brooklyn Entomological Society 31: 

73-75. 

AGASSIZ, L. 1847. Nomenclatorus zoologici 

index universalis, continens nomina systematica 

classium, ordinum, familiarum 

et generum animalium omnium, tam 

viventium quay fossilium, secundum 

ordinem alphabeticum unicum disposita, 

adjectis homonymiis plantarum, nec 

non variis adnotationibus et emendationibus. 

Jent and Gassmann, Soloduri 

[Solothurn, Switzerland]. 393 pp. 

ANDERSON, R.S. 1982a. On the decreasing 

abundance of Nicrophorus americanus 

Olivier (Coleoptera: Silphidae) in eastern 

North America. Coleopterists Bulletin 

36: 362-365. 

ANDERSON, R.S. 1982b. Burying beetle 

larvae: Nearctic Nicrophorus and Oriental 

Ptomascopus morio (Silphidae). 

Systematic Entomology 7: 249-264. 

ANDERSON, R.S. 1982c. Resource partitioning 

in the carrion beetle (Coleoptera: 

Silphidae) fauna of southern Ontario: 

ecological and evolutionary considerations. 

Canadian Journal of Zoology 60: 

1314-1325. 

ANDERSON, R.S. and S.B. PECK. 1984. 

Bionomics of Nearctic species of Aclypea 

Reitter: phytophagous “carrion” beetles 

(Coleoptera: Silphidae). Pan-Pacific Entomologist 

60: 248-255. 

ANDERSON, R.S. and S.B. PECK. 1985. The 

carrion beetles of Canada and Alaska. 

The Insects and Arachnids of Canada, 

Part 13: 1-121. 


ANGELL, J.W. 1912. Two new North American 

Necrophorus. Entomological News 

23: 307. 

ANONYMOUS. 1971. The systematic biology 

collections of the United States: an 

essential resource. Part I. The great collections: 

their nature, importance, condition, 

and future. Report to the National 

Science Foundation by the Conference 

of Directors of Systematics Collections, 

the New York Botanical Garden. 33 pp. 

ARNETT, R.H., JR. 1944. A revision of the 

Nearctic Silphini and Nicrophorini based 

on female genitalia. Journal of the New 

York Entomological Society 52: 1-25. 

ARNETT, R.H., JR. 1946. Coleoptera notes 

I: Silphidae. Canadian Entomologist 78: 

131-134. 

ARNETT, R.H., JR. 1968. The Beetles of the 

United States. The American Entomological 

Institute, Ann Arbor, MI. 1,112 pp. 

BACKLUND, D. and G. MARRONE. 1995. 

Surveys for the endangered American 

burying beetle (Nicrophorus americanus) 

in Gregory, Tripp, and Todd 

Counties, South Dakota, August 1995. 

Final Report to the U.S. Fish and Wildlife 

Service, 1995. 11 pp. and 49 maps. 

BAKER, R.G. and K.A. WALN. 1985. Quarternary 

pollen records from the Great 

Plains and central United States, pp. 

191-203. In Bryant, V.M., Jr. and R.G. 

Holloway (eds.), Pollen Records of Late 

Quarternary North American Sediments. 

American Association of Stratigraphic 

Palynologists Foundation, Dallas, TX. 

BARRY, R. 1983. Climatic environment 

of the Great Plains, past and present. 

Transactions of the Nebraska Academy 

of Sciences 11 (special issue): 45-55. 

BARTLETT, J. 1987. Filial cannibalism in 

burying beetles. Behavioral Ecology and 

Sociobiology 21: 179-183. 

BARTLETT, J. 1988. Male mating success 

and parental care in Necrophorus 

vespilloides. Behavioral Ecology and 

Sociobiology 23: 297-303. 

BARTLETT, J. and C.M. ASHWORTH. 1988. 

Brood size and fitness in Necrophorus

86 


vespilloides (Coleoptera: Silphidae). 

Behavioral Ecology and Sociobiology 

22: 429-434. 

BENINGER, C.W. 1993. Egg predation by 

Poecilochirus carabi (Mesostigmata: 

Parasitidae) and its affect on reproduction 

of Nicrophorus vespilloides (Coleoptera: 

Silphidae). Environmental 

Entomology 22: 766-769. 

BERDELA, G., B. LUSTIGMAN, and P.P. 

SHUBECK. 1994. A list of bacterial 

flora residing in the mid and hindgut 

regions of six species of carrion beetles 

(Coleoptera: Silphidae). Entomological 

News 105: 47-58. 

BERGROTH, E. 1884. Bemerkungen zur 

dritten Auflage des Catalogus Coleopterorum 

Europae auctoribus L. V. 

Heydon, E. Reitter et J. Weise. Berliner 

Entomologische Zeitschrift 28: 225-230. 

BLACKMAN, S.W. and G.O. EVANS. 1994. 

Observations on a mite (Poecilochirus 

davydovae) predatory on the eggs of burying 

beetles (Nicrophorus vespilloides) with 

a review of its taxonomic status. Journal 

of Zoology (London) 234: 217-227. 

BLACKWELDER, R.E. and R.H. ARNETT, 

JR. 1974. Checklist of the beetles of 

Canada, United States, Mexico, Central 

America and the West Indies. Volume 1, 

part 3, the scarab beetles, ant-loving beetles, 

clown beetles, and related groups 

(red version). The Biological Research 

Institute of America, Lanham, NY. Parts 

separately paginated. 

BLAND, J.H.B. 1865. Descriptions of several 

new species of North American Coleoptera. 

Proceedings of the Entomological Society 

of Philadelphia 4: 381-384. 

BLATCHLEY, W.S. 1910. An illustrated 

descriptive catalog of the Coleoptera or 

beetles (exclusive of the Rhynchophora) 

known to occur in Indiana. The Nature 

Publishing Co., Indianapolis. 1,386 pp. 

BLEED, A. and C. FLOWERDAY (eds.). 

1989. An Atlas of the Sand Hills. University 

of Nebraska-Lincoln, Conservation 

and Survey Division, Resource Atlas No. 

5: 1-238. 

BLISS, R.O. 1949. Studies on the Silphidae, 

I: secondary sexual differences in the 

genus Nicrophorus. Entomological News 

10: 197-204. 

BORNEMISSZA, G.F. 1957. An analysis of 

arthropod succession in carrion and the effect 

of its decomposition on the soil fauna. 

Australian Journal of Zoology 5: 1-12. 

BREWER, J.W. and T.R. BACON. 1975. 

Biology of the carrion beetle, Silpha 

ramosa. Annals of the Entomological 


BROWN, J.M. and D.S. WILSON. 1992. 

Local specialization of phoretic mites on 

sympatric carrion beetle hosts. Ecology 

73: 463-478. 

CATESBY, M. 1748. The Natural History 

of Carolina, Florida, and the Bahama 

Islands. Edition 3, G. Edwards ed., Vol. 

2 (1743) 100 pp. and appendix (1748), 

20 pp. 

CHEVROLAT, L.A. 1834. Coléoptères de 

Mexique, Fasc. 1. Strasbourg. 25 pp. 

CHO, Y.B. and C.E. LEE. 1986. Phylogenetic 

relationships among tribes of Silphidae 

I (Coleoptera). Nature and Life (Kyungpook 

Journal of Biological Sciences) 16: 

19-25. 

CLARK, C.U. 1895. On the food habits of certain 

dung and carrion beetles. Journal 

of the New York Entomological Society 

3: 61. 

COLE, A.C. 1942. Observations of three 

species of Silpha. American Midland 

Naturalist 28: 161-163. 

CONDRA, G.E. and E.C. REED. 1943. The 

geological section of Nebraska. Bulletin 

of the Nebraska Geological Survey 14: 

1-82. 

CONLEY, M.R. 1982. Carrion locating efficiency 

in burying beetles, Nicrophorus 

carolinus (L.) (Silphidae). Southwestern 


COOLEY, R.A. 1917. The spinach carrion 

beetle. Journal of Economic Entomology 

10: 94-102. 

CREIGHTON, J.C., C.V. VAUGHN and B.R. 

CHAPMAN. 1993. Habitat preference of 

the endangered American burying beetle

(Nicrophorus americanus) in Oklahoma. 

The Southwestern Naturalist 38: 275- 

306. 

CUTHRELL, D.L. and D.A. RIDER. (in press) 

The Silphidae of the Dakotas. Schafer-Post 

Series, North Dakota Insects Publication. 

DAVIS, L.R., JR. 1980. Notes on beetle distributions 

with a discussion of Nicrophorus 

americanus Olivier and its abundance in 

collections (Coleoptera: Scarabaeidae, 

Lampyridae, and Silphidae). Coleopterists 

Bulletin 34: 245-251. 

DAWKINS, R. 1979. Twelve misunderstandings 

of kin selection. Zeitschrift für 

Tierpsycologie 51: 184-200. 

DETHIER, V.G. 1947. The role of the antennae 

in the orientation of carrion beetles 

to odors. Journal of the New York Entomological 

Society 55: 285-293. 

DORSEY, C.K. 1940. A comparative study 

of the larvae of six species of Silpha 

(Coleoptera, Silphidae). Annals of the 

Entomological Society of America 33: 

120-139. 

EARLY, M. and M.L. GOFF. 1986. Arthropod 

succession patterns in exposed 

carrion on the island of Oahu, Hawaiian 

Islands, USA. Journal of Medical 

Entomology 23: 520-531. 

EGGERT, A.-K. 1992. Alternative male 

mate-finding tactics in burying beetles. 

Behavioral Ecology 3: 243-254. 

EGGERT, A.-K. and J.K. MÜLLER. 1989a. 

Mating success of pheromone-emitting 

Necrophorus males: do attracted females 

discriminate against resource owners? 

Behavior 110: 248-258. 

EGGERT, A.-K. and J.K. MÜLLER. 1989b. 

Pheromone-mediated attraction in burying 

beetles. Ecological Entomology 14: 

235-237. 

EGGERT, A.-K. and J.K. MÜLLER. 1992. 

Joint breeding in female burying beetles. 

Behavioral Ecology and Sociobiology 31: 

237-242. 

EGGERT, A.-K. and S.K. SAKALUK. 1995. 

Female-coerced monogamy in burying 

beetles. Behavioral Ecology and Sociobiology 

37: 147-153. 


EISNER, T. and J. MEINWALD. 1982. 

Defensive spray mechanism of a silphid 

beetle (Necrodes surinamensis). Psyche 

89: 357-367. 

ELDER, J.A. 1969. Soils of Nebraska. University 

of Nebraska-Lincoln, Conservation 

and Survey Division, Resource 

Report No. 2: 1-60. 

EMETZ, V. 1975. Ergebnisse der zoologischen 

Forschungen van Dr. Z. Kaszab 

in der Mongolei. Nr. 358. Silphidae und 

Liodidae. Folia Entomologica Hungarica 

28: 57-71. 

FABRICIUS, J.C. 1775. Systema Entomologiae. 

Lipsiae. 832 pp. 

FABRICIUS, J.C. 1776. Genera Insectorum. 

Chilonii, Bartsch. 310 pp. 

FABRICIUS, J.C. 1781. Species Insectorum, 

vol. 1. Hamburgii et Kilonii, Bohn. 552 pp. 

FABRICIUS, J.C. 1801. Systema Eleutheratorum 

Secundum Ordines, Genera, 

Species. Tomus 1. Bibliopolii Academici 

Novi, Kiliae. 506 pp. 

FALL, H.C. and T.D.A. COCKERELL. 1907. 

The Coleoptera of New Mexico. Transactions 

of the American Entomological 

Society 33: 145-272. 

FETHERSTON, I.A., M.P. SCOTT, and 

J.F.A. TRANIELLO. 1990. Parental 

care in burying beetles: the organization 

of male and female brood-care behavior. 

Ethology 85: 177-190. 

FETHERSTON, I.A., M.P. SCOTT, and 

J.F.A. TRANIELLO. 1994. Behavioural 

compensation for mate loss in the 

burying beetle Nicrophorus orbicollis. 

Animal Behaviour 47: 777-785. 

FISCHER, R.L., K.L. KNIGHT, C.D. MI- 

CHENER, W.W. MOSS, P. OMAN, J.A. 

POWELL and P.D. HURD, JR. 1975. 

Report of the Advisory Committee for 

Systematics Resources in Entomology. 

Part II. The current status of entomological 

collections in North America. 

Bulletin of the Entomological Society 

of America 21: 209-212. 

FISCHER von WALDHEIM, G. 1844. 

Spicilegium Entomographiae Rossicae. 

Bulletin de la Société Impériale des

88 


Naturalistes de Moscou 17: 3-144. 

FISHER, R.M. and R.D. TUCKERMAN. 

1986. Mimicry of bumble bees and cuckoo 

bumble bees by carrion beetles (Coleoptera: 

Silphidae). Journal of the Kansas 

Entomological Society 50: 20-25. 

FITZPATRICK, L.L. 1960. Nebraska placenames. 

University of Nebraska Press, 

Lincoln, NE. 227 pp. 

FORSTER, J.R. 1771. Novae Species Insectorum. 

Canturia I. Davies and White, 

London. 100 pp. 

FRÖLICH, J.A. 1792. Bemerkungen uber 

einige seltene Käfer aus der Insect- 

Sammlun des H.R. Rudolph in Erlangen. 

Naturforscher 26: 68-165. 

FULLER, M.E. 1934. The insect inhabitants 

of carrion: a study in animal ecology. Bulletin 

of the Australian Council of Science 

and Industrial Research 82: 1-62. 

GERMAR, E.F. 1824. Insectorum Species. 

Hendelii, Halae. 624 pp. 

GISSLER, C.F. 1880. Biological studies on 

Silpha ramosa. American Entomologist 

3: 265-267. 

GISTEL, J. 1848. Naturgeschichte des Thierreichs 

für höhere Schulen. Stuttgart. 

216 pp., 32 plates. 

GOE, M.T. 1919. Life history and habits 

of Silpha inaequalis Fab. (Coleoptera). 

Entomological News 30: 253-255. 

GREAT PLAINS FLORA ASSOCIATION. 

1977. Atlas of the Flora of the Great 

Plains. Iowa State University Press, 

Ames. 600 pp. 

GUÉRIN-MÉNEVILLE, M.F.E. 1835. Iconographie 

du Regne Anímal de G. Cuvier, 

ou Représentation d’Après Nature 

de l’une des Espèces les Plus Remarquables, 

et Souvent non Encore Figurés, 

de Chaque Genre d’Animaux. Insects. 

Vol. 7 (1829-1838). Ballière, Paris. 576 

pp., 104 plates. 

HALFFTER, G. 1982. Evolved relations 

between reproductive and subsocial 

behaviors in Coleoptera, pp. 164-170. 

In Breed, M.D., C.D. Michener, and 

H.E. Evans (eds.), The Biology of Social 

Insects. Proceedings of the Ninth Con- 

gress of the International Union for the 

Study of Social Insects. Westview Press, 

Boulder, CO. 

HALFFTER, G. 1991. Feeding, bisexual 

cooperation and subsocial behavior in 

three groups of Coleoptera, pp. 281- 

296. In Zunino, M., X. Belle’s and M. 

Blas (eds.), Advances in Coleopterology. 

European Association of Coleopterology, 

Barcelona. 

HALFFTER, G., S. ANDAGUA and C. 

HUERTA. 1983. Nidification des Nicrophorus 

(Col. Silphidae). Bulletin de la 

Société Entomologique de France 88: 

648-666. 

HATCH, M.H. 1927. Studies on the Silphinae. 

Journal of the New York Entomological 

Society 35: 331-371. 

HATCH, M.H. 1928. Fam. Silphidae II. 

Coleopterorum Catalogus, partes 95: 

63-244. W. Junk, Berlin. 

HATCH, M.H. 1932. Necrophorus or Nicrophorus. 


Society 40: 391. 

HATCH, M.H. 1946. Mr. Ross H. Arnett’s 

“Revision of the Nearctic Silphini and 

Necrophorini.” Journal of the New York 


HATCH, M.H. 1957. The Beetles of the Pacific 

Northwest. Part II: Staphyliniformia. 

University of Washington Publications 

in Biology, Vol. 16. University of Washington 

Press, Seattle, WA. 386 pp. 

HATCH, M.H. and J.W. ANGELL. 1925. A 

new North American species of Nicrophorus. 


Society 23: 216. 

HATCH, M.H. and W. RUETER, JR. 1934. 

Coleoptera of Washington: Silphidae. 

University of Washington Publications 

in Biology 1: 151-161. 

HERBST, J.F. 1793. Natursystem aller befannten 

in- und auslandischen Insecten 

als eine Fortfekung der von Buffonschen 

Natur-geschichte. Der Käfer, vol. 5. 

Pauli. Berlin. 392 pp. 

HERMAN, L.H., JR. 1964. Nomenclatural 

consideration of Nicrophorus. Coleopterists 

Bulletin 18: 5-6.

HERSCHEL, J.D. 1807. Die Europäischen 

Arten von Necrophorus mit Unterscheidung 

einer neuen Art: Necrophorus 

vestigator. Magazin für Insectenkunde 

6: 268-276. 

HINES, A.M. and R.J. SMITH. 1995. Altitudinal 

variation in estimated population 

sizes and factors in brood development in 

the burying beetle, Nicrophorus investigator 

(Silphidae) (abstract). Proceedings 

of the Nebraska Academy of Sciences 

1995: 71. 

HOPE, F.W. 1840. The Coleopterists Manual, 

Part the Third, Containing Various 

Families, Genera and Species of Beetles 

Recorded by Linnaeus and Fabricius. 

Bridgewater, London. 191 pp. 

HORN, G.H. 1880. Synopsis of the Silphidae 

of the United States with reference to the 

genera of other countries. Transactions 

of the American Entomological Society 

8: 219-322. 

HOWDEN, A.T. 1950. The succession of 

beetles on carrion. Unpublished M.S. 

Thesis, University of North Carolina, 

Chapel Hill, NC. 83 pp. 

HUERTA, C., G. HALFFTER and D. 

FRESNEAU. 1992. Inhibition of stridulation 

in Nicrophorus (Coleoptera: Silphidae): 

consequences for reproduction. 

Elytron 6: 151-157. 

JOHNSON, M.D. 1974. Seasonal and microseral 

variations in the insect populations 

on carrion. American Midland Naturalist 

93: 79-90. 

JONES, F.M. 1932. Insect coloration and the 

relative acceptability of insects to birds. 

Transactions of the Entomological Society 

of London 80: 345-386, 11 plates. 

JONES, J.K., JR. 1964. Distribution and 

taxonomy of mammals of Nebraska. University 

of Kansas Publications, Museum 

of Natural History 16: 1-356. 

KATAKURA, H. and H. FUKUDA. 1975. 

Faunal makeup of ground and carrion 

beetles in Komiotoineppu, Hokkaido 

University Nakagawa Experimental 

Forest, northern Japan, with some notes 

on related problems. Research Bulletin 


of the College of Experimental Forestry, 

Hokkaido University 32: 75-92. 

KAUL, R.B. 1986. Physical and floristic characteristics 

of the Great Plains. In Barkley, 

R.L. (ed.), Flora of the Great Plains, 

p. 7-10. University of Kansas Press. 

KAUL, R.B. 1989. Plants. In Bleed, A. and 

C. Flowerday (eds.), An Atlas of the Sand 

Hills, p. 127-142. University of Nebraska, 

Conservation and Survey Division, 

Resource Atlas No. 5: 1-265. 

KIESERITZKY, V. 1909. Species nova generis 

Thanatophilus Leach (Coleoptera: 

Silphidae). Revue Russe d’Entomologie 

9: 126-127. 

KIRBY, W. 1835. Northern zoology, Pt. 4: the 

insects. 325 pp., 8 pl. In Richardson, J., 

Fauna Boreali-Americana; or the Zoology 

of the Northern Parts of British America. 

Norwich. 

KIRBY, W. and W. SPENCE. 1828. An Introduction 

to Entomology, 5th ed. Longmass, 

Rees, Orme Brown, and Green, 

London. 683 pp. 

KORN, W. 1983. Zur Vergesellschaftung der 

Gamasidenarten Poecilochirus carabi 

G.U.R. Canestrini 1882 (= P. necrophori 

Vitzthum 1930), P. austroasiaticus Vitzthum 

1930 u. P. subterraneus Müller 

1859 mit Aakäfern aus der Familie der 

Silphidae. Spixiana 6: 251-299. 

KOZOL, A.J. 1989. Studies on the American 

burying beetle, Nicrophorus americanus, 

on Block Island. Unpublished report to 

the Nature Conservancy, 294 Washington 

Street, Boston, MA. 10 pp. 

KOZOL, A.J. 1991. Annual monitoring of 

the American burying beetle on Block 

Island. Unpublished report to the Nature 

Conservancy, 294 Washington Street, 

Boston, MA. 15 pp. 

KOZOL, A.J., M.P. SCOTT, and J.F.A. 

TRANIELLO. 1988. The American 

burying beetle, Nicrophorus americanus: 

studies on the natural history of a declining 

species. Psyche 95: 167-176. 

KOZOL, A.J., J.F.A. TRANIELLO, and 

S.M. WILLIAMS. 1994. Genetic variation 

in the endangered burying beetle

90 


Nicrophorus americanus (Coleoptera: 

Silphidae). Annals of the Entomological 


LAGO, P.K. and P.R. MILLER. 1983. Records 

of Mississippi Silphidae (Coleoptera) 

with a key to the species known to occur 

in the state. Journal of the Mississippi 

Academy of Sciences 28: 83-87. 

LAMPERT JR., L.L. 1977. Notes on Silpha 

inaequalis Fabricius (Coleoptera: Silphidae). 

Coleopterists Bulletin 31: 132. 

LANE, C. and M. ROTHSCHILD. 1965. 

A case of Müllerian mimicry of sound. 

Proceedings of the Royal Entomological 

Society of London (A) 40: 156-158. 

LAPORTE, F.L., COMTE DE CASTELNAU. 

1840. Histoire Naturelle des Insectes, 

Coléoptères. Tome 2. Dumenil, Paris. 

563 pp. 

LAWRENCE, J.F. and A.F. NEWTON, JR. 

1982. Evolution and classification of 

beetles. Annual Review of Ecology and 

Systematics 13: 261-290. 

LAWRENCE, J.F. and A.F. NEWTON, JR. 

1995. Families and subfamilies of Coleoptera 

(with selected genera, notes, 

references and data on family-group 

names), pp. 779-1092. In J. Pakaluk and 

S.A. Slipinski (eds.), Biology, Phylogeny, 

and Classification of Coleoptera: Papers 

Celebrating the 80th Birthday of Roy 

Crowson, Vol. 2. Muzeum i Instytut 

Zoologi PAN, Warszawa. 

LEACH, W.E. 1815. The Zoological Miscellany; 

Being Descriptions of New, or Interesting 

Animals, vol. 2. London. 154 pp. 

LeCONTE, J.L. 1853. Synopsis of the Silphidae 

of America, north of Mexico. Proceedings 

of the Academy of Natural Sciences 


LeCONTE, J.L. 1854. Descriptions of some 

new Coleoptera from Oregon, collected by 

Dr. J.G. Cooper of the North Pacific R.R. 

expedition, under Governor J.J. Stevens. 

Proceedings of the Academy of Natural 

Sciences of Philadelphia 7: 16-20. 

LENG, C.W. 1920. Catalogue of the Coleoptera 

of America, North of America. 

John D. Sherman, Jr. Publishing, Mount 

Vernon, NY. 470 pp. 

LINGAFELTER, S.W. 1995. Diversity, 

habitat preferences, and seasonality 

of Kansas carrion beetles (Coleoptera: 

Silphidae). Journal of the Kansas Entomological 

Society 68: 214-223. 

LINNAEUS, C. 1771. Mantissa Plantarum 

Altera Generum Editionis VI et Specierum 

Editionis II. Laurentii Salvii, 

Holmiae. Pp. 144-582. 

LINNAEUS, C. 1758. Systema Naturae per 

Regna tria Naturae Secundum Classes, 

Ordines, Genera, Species, cum Characteribus, 

Different iis, Synonymis, Locis. 

Edition 10, Vol. 1. Holmiae. 823 pp. 

LOMOLINO, M.V., J.C. CREIGHTON, 

G.D. SCHNELL, and D.L. CERTAIN. 

1995. Ecology and conservation of the 

endangered American burying beetle 

(Nicrophorus americanus). Conservation 

Biology 9: 605-614. 

MADGE, R.B. 1980. A catalogue of typespecies 

in the family Silphidae (Coleoptera). 

Entomologica Scandinavica 

11: 353-362. 

MANNERHEIM, C.G. 1843. Beitrag zur 

Käfer-Fauna der Aleutischen Inselen 

und der Insel Sitkha und New-Californiens. 

Bulletin de la Société Impériale des 

Naturalistes de Moscou 16: 175-314. 

MANNERHEIM, C.G. 1853. Dritter Nachtrag 

zur Käfer-Fauna der Nord-Amerikanischer 

Laender des Russischen 

Reiches. Bulletin de la Société Impériale 

des Naturalistes de Moscou 26: 95-273. 

MATTHEWS, A. 1888. Biologia Centrali- 

Americana. Insecta, Coleoptera, Vol. 1, 

Pt. 2. Pp. 72-101. 

MESERVE, F.G. 1936. The Silphidae of 

Nebraska. Entomological News 47: 

132-134. 

MILLER, S.E. and S.B. PECK. 1979. Fossil 

carrion beetles of Pleistocene California 

asphalt deposits, with a synopsis of recent 

California Silphidae. Transactions 

of the San Diego Society of Natural 

History 19: 85-106. 

MILNE, L.J. and M.J. MILNE. 1944. Notes 

on the behavior of burying beetles

(Nicrophorus spp.). Journal of the 

New York Entomological Society 52: 

311-327. 

MILNE, L.J. and M.J. MILNE. 1976. The 

social behavior of burying beetles. Scientific 

American 235: 84-89. 

MOTSCHULSKY, V. 1845. Die Coleopterologischen 

Verhältnisse und die 

Käfer Russlands. Bulletin de la Société 

Impériale des Naturalistes de Moscou 

18: 1-127. 

MROCZKOWSKI, M. 1949. Notes on the 

successive appearance of some species 

of the genera Nicrophorus and Neonicrophorus. 

Polskie Pismo Entomologiczne 

19: 196-199. 

MÜLLER, J.K. 1987. Replacement of a lost 

clutch: a strategy for optimal resource 

utilization in Necrophorus vespilloides 

(Coleoptera: Silphidae). Ethology 76: 

74-80. 

MÜLLER, J.K. and A-K. EGGERT. 1987. 

Effects of carrion-independent pheromone 

emission by male burying beetles 

(Silphidae: Necrophorus). Ethology 76: 

297-304. 

MÜLLER, J.K. and A.-K. EGGERT. 1989. 

Paternity assurance by “helpful” males: 

adaptations to sperm competition in 

burying beetles. Behavioural Ecology 

and Sociobiology 24: 245-249. 

MÜLLER, J.K. and A.-K. EGGERT. 1990. 

Time dependent shifts between infanticidal 

and parental behavior in female 

burying beetles: a mechanism of individual 

mother-offspring recognition. 

Behavioural Ecology and Sociobiology 

27: 11-16. 

MÜLLER, J.K., A.-K. EGGERT and J. 

DRESSEL. 1990. Intraspecific brood 

parasitism in the burying beetle, Necrophorus 


Animal Behaviour 40: 491-499. 

MÜLLER, J.K., A.-K. EGGERT and E. 

FURLKRÖGER. 1990. Clutch size 

regulation in the burying beetle Necrophorus 

vespilloides Herbst (Coleoptera: 

Silphidae). Journal of Insect Behavior 

3: 265-270. 


MUTHS, E.L. 1991. Substrate discrimination 

in burying beetles, Nicrophorus orbicollis 

(Coleoptera: Silphidae). Journal 

of the Kansas Entomological Society 64: 

447-450. 

NEWMAN, E. 1838. Entomological notes. 

The Entomologist’s Monthly Magazine 

5: 372-402. 

NIEMITZ, C. 1972. Bioakustische, verhalten-physiologische 

und morpho-logische 

Untersuchungen an Necrophorus 

vespillo (Fab.). Forma et Functio 5: 

209-230. 

NIEMITZ, C. and A. KRAMPE. 1972. Untersuchungen 

zum Orientierungs-verhalten 

der Larven von Necrophorus vespillo F. 

(Silphidae, Coleoptera). Zeitschrift für 

Tierpsycologie 30: 456-463. 

OLIVIER, G.A. 1790. Entomologie, ou Histoire 

Naturelle des Insectes avec leurs 

Caractères Génériques et Spécifiques, 

leur Description, leur Synonymie, et leur 

Figure Enluminée. Coléoptères, Vol. 2. 

Genera separately paged. Paris. 

OTRONEN, M. 1988. The effect of body 

size on the outcome of fights in burying 

beetles (Nicrophorus). Annales Zoologici 

Fennici 25: 191-201. 

PAYNE, J.A. 1965. A summer carrion study 

of the baby pig Sus scrofa L. Ecology 46: 

592-602. 

PECK, S.B. and R.S. ANDERSON. 1985. 

Taxonomy, phylogeny and biogeography 

of the carrion beetles of Latin America 

(Coleoptera: Silphidae). Quaestiones 

Entomologique 21: 247-317. 

PECK, S.B. and M.M. KAULBARS. 1987. 

A synopsis of the distribution of the 

carrion beetles (Coleoptera: Silphidae) 

of the conterminous United States. Proceedings 

of the Entomological Society of 

Ontario 188: 47-81. 

PECK, S.B. and S.E. MILLER. 1993. A catalog 

of the Coleoptera of America north of 

Mexico. Family: Silphidae. USDA Agric. 

Handbook No. 529-28: 1-24. 

PETERSON, A. 1960. Larvae of Insects. Part 

II. Published by the Author. Columbus, 

OH. 416 pp.

92 


PETRUSKA, F. 1975. Vliv preladajicich 

smeru vetru na nalet nekterych druhu 

brouku z celedi Silphidae do zemnich 

pasti. Acta Univsitatis Palackianae 

Olomucensis Facultas Rerum Natalium 

Biolica 51: 155-176. 

PEYTON, M.M. 1994. A report on the collection 

of 40 specimens of the endangered 

American burying beetle (Nicrophorus 

americanus) in the Upper Platte River 

Valley. Unpublished Report to the U.S. 

Fish and Wildlife Service, 12 pp. 

PIERCE, W.D. 1949. Fossil arthropods 

of California. 17. The silphid burying 

beetles in the asphalt deposits. Bulletin 

of the Southern California Academy of 

Sciences 48: 55-70. 

PIRONE, D.J. 1974. Ecology of necrophilus 

and carpophilus Coleoptera in a southern 

New York woodland (phenology, 

aspection, trophic and habitat preferences). 

Unpublished Ph.D. Dissertation, 

Fordham University. 769 pp. 

POOL, R.J. 1929. Handbook of Nebraska 

trees. University of Nebraska, Conservation 

and Survey Division Bulletin 7: 

1-179. 

PORTEVIN, M.G. 1903. Remarques sur 

les necrophages du muséum et description 

d’especes nouvelles. Bulletin du 

Museum National d’Histoire (Paris) 9: 

329-336. 

PORTEVIN, M.G. 1914. Révision des silphides, 

liodides et clambides du Japon. 

Annales de la Société Entomologique de 

Belgique 58: 212-236. 

PORTEVIN, M.G. 1920. Revision des 

Silphini et Necrophorini de la region 

Indo-Malaise. Bulletin du Museum 

National d’Histoire Naturelle (Paris) 

26: 395-401. 

PORTEVIN, M.G. 1921. Note sur quelques 

silphides et liodides de la collection 

Grouvelle. Bulletin du Museum National 

d’Histoire Naturelle, Paris 27: 

535-538. 


silphides des collection du muséum. 

Bulletin du Muséum National d’Histoire 

Naturelle 28: 505-508. 

PORTEVIN, M.G. 1923. Révision des Necrophorini 

du globe. Bulletin du Muséum 

National d’Histoire Naturelle, Paris 29: 

64-71, 141-146, 226-233, 303-309. 




83-87, 374-377. 




165-170. 

PORTEVIN, M.G. 1926. Les Grandes 

Necrophages du Globe. Encyclopedíe 

Entomologique 6: 1-270. 


Silphidae du Deutsches Entomo-logisches 

Museum. Bulletins Mensuels, 

Société Naturalistes Luxembourgeois 

26: 58-60. 

PUKOWSKI, E. 1933. Oekologische Untersuchungun 

an Necrophorus F. Zeitschrift 

für Morphologie und Ökologie der Tiere 

27: 518-586. 

RATCLIFFE, B.C. 1972. The natural history 

of Necrodes surinamensis (Fabr.) 

(Coleoptera: Silphidae). Transactions 

of the American Entomological Society 

98: 359-410. 

RATCLIFFE, B.C. 1980. A matter of taste 

or the natural history of carrion beetles. 

UNL News 59: 1-4 (Museum Notes No. 

67: 1-4). 

RATCLIFFE, B.C. 1995. Nebraska’s threatened 

and endangered species. American 

burying beetle. Nebraska Game and 

Parks brochure. In NEBRASKAland 

Magazine 73, 6 pp. 

RATCLIFFE, B.C. and M.L. JAMESON. 

1992. New Nebraska occurrences of the 

endangered American burying beetle 

(Coleoptera: Silphidae). Coleopterists 

Bulletin 46: 421-425. 

RECCE, S. 1988. Endangered and threatened 

wildlife and plants: proposed 

endangered status for the American 

burying beetle. Federal Register 53(196): 

39, 617-39, 621.

REED, H.B. 1958. A study of dog carcass 

communities in Tennessee, with special 

reference to the insects. American Midland 


RICHTER, S. 1993. Phoretic association between 

the dauer juveniles of Rhabditis 

stammeri (Rhabditidae) and life history 

stages of the burying beetle Nicrophorus 


Nematologica 39: 346-355. 

ROBERTSON, I.C. 1992. Relative abundance 

of Nicrophorus pustulatus (Coleoptera: 

Silphidae) in a burying beetle 

community, with notes on its reproductive 

behavior. Psyche 99: 189. 

SAY, T. 1823. Descriptions of coleopterous 

insects collected in the late expedition 

to the Rocky Mountains, performed by 

order of Mr. Calhoun, Secretary of War, 

under the command of Mjr. Long. Journal 

of the Academy of Natural Sciences 


SAY, T. 1825. Descriptions of new coleopterous 

insects inhabiting the 

United States. Journal of the Academy 

of Natural Sciences of Philadelphia 5: 

160-202. 

SCHAWALLER, W. 1981. Taxonomie und 

Faunistik der Gattung Thanatophilus 

(Coleoptera: Silphidae). Stuttgarter 

Beiträge zur Naturkunde, Series A 

(Biologie) No. 351: 1-21. 

SCHILDKNECHT, H. and K.H. WEIS. 

1962. Über die chemische Abwehr 

der Aaskäfer. XIV. Mitteilung über 

Insecktenabwehrstoffe. Zeitschrift für 

Naturforschung 17b: 452-455. 

SCHWARZ, H.H. and J.K. MÜLLER. 1992. 

The dispersal behaviour of the phoretic 

mite Poecilochirus carabi (Mesostigmata, 

Parasitidae): adaptation to the 

breeding biology of its carrier Necrophorus 

vespilloides (Coleoptera, Silphidae). 

Oecologia 89: 487-493. 

SCHWEITZER, D.F. and L.L. MAS- 

TER. 1987. Nicrophorus americanus 

(American burying beetle): results of 

a global status survey. The Nature 

Conservancy, Eastern Heritage Task 


Force, Boston, MA. 13 pp. 

SCOTT, M.P. 1989. Male parental care and 

reproductive success in the burying 

beetle, Nicrophorus orbicollis. Journal 

of Insect Behavior 2: 133-137. 

SCOTT, M.P. 1990. Brood guarding and 

the evolution of male parental care in 

burying beetles. Behavioral Ecology 

and Sociobiology 26: 31-39. 

SCOTT, M.P. 1994a. The benefit of paternal 

assistance in intra- and interspecific 

competition for the burying beetle, 

Nicrophorus defodiens. Ethology, Ecology 

and Evolution 6: 537-543. 

SCOTT, M.P. 1994b. Competition with flies 

promotes communal breeding in the 

burying beetle, Nicrophorus tomentosus. 

Behavioral Ecology and Sociobiology 

34: 367-373. 

SCOTT, M.P. and D.S. GLADSTEIN. 1993. 

Calculating males? An empirical and 

theoretical examination of the duration 

of paternal care in burying beetles. 

Evolutionary Ecology 7: 362-378. 

SCOTT, M.P. and J.F.A. TRANIELLO. 

1987. Behavioural cues trigger ovarian 

development in the burying beetle, 

Nicro-phorus tomentosus. Journal of 

Insect Physiology 33: 693-696. 


1989. Guardians of the underworld. 

Natural History Magazine 6: 32-36. 


1990a. Brood guarding and the evolution 

of male parental care in burying 

beetles. Behavioral Ecology and Sociobiology 

26: 31-39. 


1990b. Behavioural and ecological correlates 

of male and female parental 

care and reproductive success in burying 

beetles (Nicrophorus spp.). Animal 

Behaviour 39: 274-283. 

SCOTT, M.P., J.F.A. TRANIELLO and I.A. 

FETHERSTON. 1987. Competition for 

prey between ants and burying beetles 

(Nicrophorus spp.): differences between 

northern and southern temperature 

sites. Psyche 94: 325-332.

94 


SCOTT, M.P. and S.M. WILLIAMS. 1993. 

Comparative reproductive success of 

communally breeding burying beetles as 

assessed by PCR with randomly amplified 

polymorphic DNA. Proceedings of 

the National Academy of Science USA 

90: 2,242-2,245. 

SEMENOV-TIAN-SHANSKIJ, A. 1891. Diagnoses 

coleopterorum novorum ex Asia 

Centrali et Orientali. Horae Societatis 

Entomologicae Rossicae 25: 262-382. 

SEMENOV-TIAN-SHANSKIJ, A. 1926. 

Analecta Coleopterologica XIX. Entomologicheskoe 

Obozrenie 20: 33-55. 

SEMENOV-TIAN-SHANSKIJ, A. 1933. De 

tribu Necrophorini (Coleoptera: Silphidae), 

classificanda et de ejua distributione 

geographica. Trudy Zoologicheski 

Instituta (Leningrad), Akademiya Nauk 

SSR 1: 149-160. 

SHUBECK, P.P. 1968. Orientation of 

carrion beetles to carrion: random or 

non-random. Journal of the New York 


SHUBECK, P.P. 1971. Diel periodicities of 

certain carrion beetles. Coleopterists 

Bulletin 25: 41-46. 

SHUBECK, P.P. 1975a. Do carrion beetles 

use sight, as an aid to olfaction, in locating 

carrion? The William L. Hutcheson 

Memorial Forest Bulletin 3: 36-39. 

SHUBECK, P.P. 1975b. Flight activity of 

certain carrion beetles: Silpha noveboracensis, 

Staphylinidae, Histeridae. 

Bulletin of the William L. Hutcheson 

Memorial Forest 3: 40-43. 

SHUBECK, P.P. 1976. Carrion beetle responses 

to poikilothermic and homiothermic 

carrion. Entomological News 89: 

265-269. 

SHUBECK, P.P. 1983. Habitat preferences 

of carrion beetles in the Great Swamp 

National Wildlife Refuge, New Jersey 

(Coleoptera: Silphidae, Dermestidae, 

Nitidulidae, Histeridae, Scarabaeidae). 


Society 91: 333-341. 

SHUBECK, P.P. 1984a. Habitat preferences 

of carrion beetles in the Great Swamp 

National Wildlife Refuge, New Jersey 

(Coleoptera: Silphidae, Dermestidae, 

Nitidulidae, Histeridae, Scarabaeidae). 


Society 91: 333-341. 

SHUBECK, P.P. 1984b. An inexpensive carrion 

beetle trap (Coleoptera: Silphidae). 

Entomological News 95: 63-64. 

SHUBECK, P.P. 1993. An ecotonal study of 

carrion beetles (Coleoptera: Silphidae) 

in the Great Swamp National Wildlife 

Refuge, New Jersey. Entomological 

News 104: 88-92. 

SMITH, H.T.V. 1965. Dune morphology and 

chronology in central and western Nebraska. 

Journal of Geology 73: 557-578. 

SMITH, R.J. and B. HEESE. 1995. Carcass 

selection in a high altitude population of 

the burying beetle, Nicrophorus investigator 

(Silphidae). The Southwestern 


SOLTER, L.F., B. LUSTIGMAN, and P. 

SHUBECK. 1989. Survey of medically 

important true bacteria found associated 

with carrion beetles (Coleoptera: Silphidae). 

Journal of Medical Entomology 26: 

354-359. 

SPRINGETT, B.P. 1968. Aspects of the relationship 

between burying beetles, Necrophorus 

spp. and the mite Poecilochirus 

necrophori Vitz. Journal of Animal Ecology 

37: 417-424. 

STEELE, B.F. 1927. Notes on the feeding 

habits of carrion beetles. Journal of the 

New York Entomological Society 35: 

77-81. 

THUNBERG, C.P. 1789. Periculum entomologicum, 

quo characteres generum 

insectorum proponit. Diss. Response, 

Tornes 10 Juni 1789, Upsaliae. Edman. 

16 pp. 

TORRE-BUENO, J.R. de la. 1937. A Glossary 

of Entomology. Brooklyn Entomological 

Society, Brooklyn. 336 pp. 

TRUMBO, S.T. 1987. The ecology of parental 

care in burying beetles (Silphidae: 

Nicrophorus). Unpublished Ph.D. Dissertation, 

University of North Carolina, 

Chapel Hill, NC.

TRUMBO, S.T. 1990a. Reproductive benefits 

of infanticide in a biparental burying 

beetle Nicrophorus orbicollis. Behavioral 

Ecology and Sociobiology 27: 267-273. 

TRUMBO, S.T. 1990b. Interference competition 

among burying beetles (Silphidae, 

Nicrophorus). Ecological Entomology 

15: 347-355. 

TRUMBO, S.T. 1990c. Regulation of brood 

size in a burying beetle, Nicrophorus 

tomentosus. Journal of Insect Behavior 

3: 491-500. 

TRUMBO, S.T. 1990d. Reproductive success, 

phenology and biogeography of burying 

beetles (Silphidae, Nicrophorus). American 

Midland Naturalist 124: 1-11. 

TRUMBO, S.T. 1991. Reproductive benefits 

and the duration of paternal care in a 

biparental burying beetle, Necrophorus 

(sic) orbicollis. Behaviour 117: 82-105. 

TRUMBO, S.T. 1992. Monogamy to communal 

breeding: exploitation of a brood 

resource base by burying beetles (Nicrophorus). 

Ecological Entomology 17: 

289-298. 

TRUMBO, S.T. 1994. Interspecific competition, 

brood parasitism, and the evolution 

of biparental cooperation in burying 

beetles. Oikos 69: 241-249. 

TRUMBO, S.T. 1995. Nesting failure in burying 

beetles and the origin of communal 

associations. Evolutionary Ecology 9: 

125-130. 

TRUMBO, S.T. and A.-K. EGGERT. 1994. 

Beyond monogamy: territory quality 

influences sexual advertisement in male 

burying beetles. Animal Behaviour 48: 

1,043-1,047. 

TRUMBO, S.T. and A.G. FERNANDEZ. 

1995. Regulation of brood size by male 

parents and cues employed to assess resource 

size by burying beetles. Ethology 

Ecology and Evolution 7: 313-322. 

TRUMBO, S.T. and A.J. FIORE. 1991. A 

genetic marker for investigating paternity 

and maternity in the burying beetle 

Nicrophorus orbicollis (Coleoptera: 

Silphidae). Journal of the New York 



TRUMBO, S.T. and A.J. FIORE. 1994. 

Interspecific competition and the evolution 

of communal breeding in burying 

beetles. American Midland Naturalist 

131: 169-174. 

TRUMBO, S.T. and D.S. WILSON. 1993. 

Brood discrimination, nest mate discrimination, 

and determinants of social 

behavior in facultatively quasisocial 

beetles (Nicrophorus spp.). Behavioral 

Ecology 4: 332-339. 

U.S. FISH AND WILDLIFE SERVICE. 

1991. American burying beetle (Nicrophorus 

americanus) recovery plan. 

Newton Corner, MA. 80 pp. 

VOET, J.E. 1778. Catalogus Systematicus 

Coleopterorum, vol. 1. Latin, French, and 

Dutch translations separately paged. 

Le Haye. 

WALDOW, U. 1973. The electrophysiology 

of a new carrion smell receptor and its 

role in the behavior of Necrophorus. 

Journal of Comparative Physiology 83: 

415-424. 

WATTS, W.A. and H.E. WRIGHT, JR. 1966. 

Late-Wisconsin pollen and seed analysis 

from the Nebraska Sandhills. Ecology 

47: 202-210. 

WEAVER, J.E. 1954. North American Prairie. 

Johnsen Publ. Co., Lincoln, NE. 348 pp. 

WEAVER, J.E. 1965. Native Vegetation of 

Nebraska. University of Nebraska Press, 

Lincoln, NE. 185 pp. 

WEAVER, J.E. and F.E. CLEMENTS. 1938. 

Plant Ecology. McGraw-Hill Book Co., 

Inc., New York, NY. 601 pp. 

WEBER, F. 1801. Observationes Entomologicae, 

Continentes Novorum quae Condidit 

Generum Characteres, et Nuper Detectarum 

Specierum Descriptiones. Bibliopolii 

Academici Novi. Kiliae. 116 pp. 

WELLS, S.M., R.M. PYLE, and N.M. COL- 

LINS. 1983. The IUCN Red Data Book. 

IUCN, Gland, Switzerland. 650 pp. 

WILSON, D.S. 1983. The effect of population 

structure on the evolution of mutualism: 

a field test involving burying beetles and 

their phoretic mites. American Naturalist 

121: 851-870.

96 


WILSON, D.S. and J. FUDGE. 1984. Burying 

beetles: intraspecific interactions 

and reproductive success in the field. 

Ecological Entomology 9: 195-203. 

WILSON, D.S. and W.G. KNOLLENBERG. 

1984. Food discrimination and ovarian 

development in burying beetles 

(Coleoptera: Silphidae: Nicrophorus). 

Annals of the Entomological Society of 

America 77: 165-170. 

WILSON, D.S. and W.G. KNOLLENBERG. 

1987. Adaptive indirect effects: the fitness 

of burying beetles with and without 

their phoretic mites. Evolutionary Ecology 

1: 139-159. 

WILSON, D.S., W.G. KNOLLENBERG 

and J. FUDGE. 1984. Species packing 

and temperature dependent competition 

among burying beetles (Silphidae, 

Nicrophorus). Ecological Entomology 9: 

205-216. 

WILSON, E.O. 1971. The Insect Societies. 

Harvard University Press, Cambridge, 

MA. 548 pp. 

WRIGHT, H.E., JR. 1970. Vegetational history 

of the Central Plains, pp. 157-176. 

In Dort, W., Jr. and J.K. Jones, Jr. (eds.), 

Pleistocene and Recent Environments of 

the Central Great Plains. University of 

Kansas Press, Lawrence, KS. 

YOUNG, O.P. 1983. The biology of the Silphidae 

(Coleoptera): a coded bibliography. 

Maryland Agricultural Experiment 

Station Miscellaneous Publication 981: 

1-47. 

YOUNG, O.P. 1985. Survival of a carrion 

beetle, Necrodes surinamensis 

(Coleoptera: Silphidae), on a diet of dead 

fall armyworm (Lepidoptera: Noctuidae) 

larvae. Journal of Entomological Science 

20: 359-366. 

ZEH, D.W. and R.L. SMITH. 1985. Paternal 

investment by terrestrial arthropods. 

American Zoologist 25: 785-805. 

ZETTERSTEDT, J.W. 1824. Nya Svenska 

Insect-Arten. Svenska Vetenskaps 

Akademien Handlingar 1824: 149-159.

GLOSSARY 

(modified from Torre-Bueno 1937) 

Antenna (pl. antennae). The paired, segmented 

sensory organs borne on the 

head. 

Anterior. Front or forward; opposite of 

posterior. 

Aposematic. Conspicuous and warning of 

danger. 

Base. On the thorax, that part nearest the 

abdomen; on the abdomen, that part 

nearest the thorax. 

Carina. A longitudinal, narrow, raised ridge. 

Club. The enlarged, distal segments of the 

antenna (Figs. 83-84). 

Clypeus. That part of the head in front of 

the frons. In dorsal view of a silphid, 

that part of the head between the frons 

and labrum (Fig. 2). 

Congeneric. Belonging to the same genus. 

Conspecific. Belonging to the same species. 

Cordate. Loosely, heart-shaped or triangular 

with the corners rounded. 

Costa. Longitudinal, elevated ridges of the 

wing covers. 

Coxa. Basal segment of the leg that articulates 

with the body (Fig. 3). 

Diurnal. Active during the day. 

Dorsal. The upper surface. 

Elytron (pl. elytra). The anterior, chitinous 

wings of beetles that serve as covers to 

the folded flight wings (Figs. 2, 21-33). 

Emarginate. Notched or with a rounded or 

obtuse section removed from a margin. 

Empodium. The single pad-like or filiform 

median structure often present in the 

insect claw. 

Epimeron. The posterior division of a 

thoracic pleuron, usually small (Fig. 3, 

85-92). 

Epipleuron. The deflexed portion of the 

lateral edge of the wing cover. 

Exuvium. The cast skin of the larva after 

metamorphosis. 

Femur. Usually the stoutest segment of 

the leg, articulated to the body by the 

trochanter and coxa and bearing the 

tibia at its distal end (Figs. 2-3). 


Frons. The upper portion of the head capsule 

behind the clypeus and before the 

vertex. 

Glabrous. Smooth, lacking setae. 

Hemolymph. The blood of arthropods. 

Hindgut. The posterior portion of the alimentary 

tract, between the midgut and 

the anus. 

Humerus. The basal, exterior angle of the 

elytra; shoulder. 

Impressed. Having shallow, depressed 

areas. 

Instar. The form between molts in the 

larva, numbered to designate the various 

periods, e.g., the first instar is the stage 

between the egg and the first molt, etc. 

Interval. The longitudinal space between 

striae or costae on the elytra. 

Labrum. The upper lip that covers the base 

of the mandibles and forms the roof of 

the mouth (Fig. 2). 

Lateral. Relating to the side. 

Longitudinal. In the direction of the long 

axis. 

Macula. A colored mark larger than a spot, 

of indeterminate shape. 

Mandibles. The stout, tooth-like pair of 

jaws in chewing insects. 

Margin. The more or less narrow part of a 

surface adjacent to the edge. 

Median. Pertaining to the middle. 

Meso. Greek prefix referring to the middle. 

Meta. Greek prefix referring to posterior 

(generally third). 

Metasternum. The ventral part of the 

metathorax. In silphids, generally the 

large plate extending from the middle to 

the posterior legs (Fig. 3). 

Metepimeron. In Nicrophorus species, the 

small lobe behind and laterad of the 

metasternum (Figs. 3, 85-92). 

Midgut. The middle portion of the alimentary 

tract. 

Necrophagous. Feeding on dead or decaying 

matter. 

Nocturnal. Active at night. 

Ocellus (pl. ocelli). The simple eye in 

adult insects. See Stemma. 

Orbicular. Round and flat.

98 


Pheromone. A chemical that is secreted to 

the outside that causes a specific reaction 

in a receiving individual of the 

same species. 

Phoretic. Referring to the relationship 

between different species where one is 

carried on the body of another. 

Piceous. Pitchy black. 

Pleuron. The lateral region of any segment 

of the insect body. 

Posterior. Rear or rearward; opposite of 

anterior. 

Posterolateral. Toward the rear and side. 

Pro. Latin prefix meaning anterior or before. 

Pronotum. The upper or dorsal surface of 

the thorax (prothorax) (Fig. 2). 

Pubescence. Short, fine setae. 

Punctate. With impressed points or punctures. 

Puncture. A small impression on the hard 

surface of the body. 

Pygidium. In dorsal view, the last segment 

usually left partially exposed by 

the elytra. 

Ruga (pl. rugae). A wrinkle. 

Rugose. Wrinkled. 

Saprophagous. Feeding on dead or decaying 

vegetable matter. 

Scape. The first or basal segment of the 

antenna. 

Sclerite. Any piece of the insect body wall 

bordered by sutures. 

Scutellum. The triangular piece between 

the bases of the elytra (Fig. 2). 

Setigerous. Bearing setae. 

Spur. A spine-like appendage, articulated 

or not, usually on the tibia (Fig. 2). 

Stadium. The interval of time between the 

molts of the larva. 

Stem. The segments of the antenna exclusive 

of the club. 

Stemma (pl. stemmata). The simple eye of 

holometabolous larvae. 

Sternite. The ventral part of a segment. 

Stria (pl. striae). A longitudinal, depressed 

line or furrow, frequently with punctures, 

usually extending from the base 

to the apex of the elytra. 

Stridulation. The sound produced by 

rubbing one surface or structure upon 

another. 

Stridulatory Ridge. In Nicrophorus 

species, the short, subparallel, transversely 

grooved ridges on top of the 

third abdominal segment that are used 

in stridulation. 

Sub. Latin prefix meaning almost or not 

quite. 

Subequal. Similar but not quite equal in 

form, size or other characters. 

Suture. The longitudinal line of juncture 

of the elytra. 

Sympatric. Species or populations whose 

distributions overlap at least in part. 

Tarsomere (pl. tarsomeres). One of the 

segments of the tarsus. 

Tarsus (pl. tarsi). The foot; the jointed 

appendage attached to the apex of the 

tibia. The distal part of the insect leg 

consisting of (in silphids) five segments 

(Figs. 2-3). 

Teneral. The condition of the adult shortly 

after emergence when it is not entirely 

hardened or fully colored. 

Tergite. The dorsal part of a segment. 

Thanatosis. The act of faking death. 

Tibia (pl. tibiae). The fourth division of 

the leg, articulated at the proximal end 

to the femur and bearing the tarsus 

on the distal end (Figs. 2-3). 

Transverse. Broader than long, or crossing 

the longitudinal axis at right angles. 

Trochanter. The second segment of the 

insect leg between the coxa and the 

femur. 

Truncate. Cut off squarely at the apex. 

Tubercle. A small, conical bump. 

Umbone. An elevated knob situated on 

the humerus or near the apical angles 

of the elytra, hence humeral umbone or 

apical umbone. 

Urogomphi (sing. urogomphus). Elongated 

processes found on the terminal segments 

of certain larvae (Figs. 34-39). 

Ventral. Pertaining to the under surface. 

Vitellogenesis. Referring to yolk production 

in the egg.


CHECKLIST OF NEBRASKA SILPHIDAE 

Life List Subfamily Silphinae 

Heterosilpha ramosa (Say) 

Necrodes surinamensis (Fabr.) 

Necrophila americana (L.) 

Oiceoptoma inaequale (Fabr.) 

Oiceoptoma novaboracense (Forster) 

Thanatophilus lapponicus (Herbst) 

Thanatophilus truncatus (Say) 

Subfamily Nicrophorinae 

Nicrophorus americanus Olivier 

Nicrophorus carolinus (L.) 

Nicrophorus guttula Motschulsky 

Nicrophorus hybridus Hatch and Angell 

Nicrophorus investigator Zetterstedt 

Nicrophorus marginatus Fabr. 

Nicrophorus mexicanus Matthews 

Nicrophorus obscurus Kirby 

Nicrophorus orbicollis Say 

Nicrophorus pustulatus Herschel 

Nicrophorus tomentosus Weber

100 


ABOUT THE AUTHOR 

Dr. Brett Ratcliffe is the Curator of the Division of Entomology and Professor at the 

University of Nebraska State Museum in Lincoln. He is a specialist in the taxonomy, biology, 

ecology, phylogeny, and biogeography of scarab beetles, especially those of the Neotropics. He 

studied silphids for his Master's degree, and he has maintained an interest in carrion beetles 

ever since. He is currently leading research programs in Nebraska to study the endangered 

American burying beetle (Nicrophorus americanus), which is now found in only five states, 

including Nebraska. 

Dr. Ratcliffe has conducted extensive field research in Japan, North America, Mexico, 

Central America, South America, and South Africa. From 1976 to 1978, he was head of the 

systematics entomology research collections for the National Institute for Amazonian Research 

(INPA) in Manaus, Brazil. During the past five years, he has collaborated closely with the 

National Institute for Biodiversity (INBio) in Costa Rica, the University of Panama, and the 

Smithsonian's Tropical Research Institute in Panama to conduct biodiversity survey programs 

of the dynastine scarab beetles of those countries. He is the author of numerous scientific 

papers and popular articles about beetles as well as a book, The Scarab Beetles of Nebraska.

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