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HARVARD UNIVERSITY

Library of the
Museum of

Comparative Zoology

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A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.

VOLUME 21

NUMBER 1

1985

Publication of Quaestiones Entomologicae was started in 1965 as part of a
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QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.

Volume 21 1985

CONTENTS

Bell and Bell-Rhysodini of the World. Part IV. Revisions of Rhyzodiastes Fairmaire and
Clinidium Kirby, with New Species in Other Genera (Coleoptera: Carabidae or Rhysodidae) 1

Book Review-Belton, P. 1983. Review of Mosquitoes of British Columbia 173

Book Notice-Griffiths, G.C.D. (Editor). 1983-84. Flies of the Nearctic Region. Part 2,

Numbers 2-3. Anthomyiidae 174

Fredeen-Some Economic Effects of Outbreaks of Black Flies ( Simulium luggeri Nicholson

and Mickel) in Saskatchewan 175

Ward-The Nearctic Species of the Genus Pseudomyrmex (Hymenoptera: Formicidae) 209
Peck and Anderson-Taxonomy, phytogeny and biogeography of the carrion beetles of Latin

America (Coleoptera: Silphidae) 247

Hilchie-The tiger beetles of Alberta (Coleoptera: Carabidae, Cicindelini) 319

Ball-Characteristics and evolution of elytral sculpture in the tribe Galeritini (Coleoptera:

Carabidae) 349

Book Review-Manson, D.C.M. 1984. Fauna of New Zealand; Number 4 369

Faunal Influences on Soil Structure - A Symposium 371.1

Quaest. Ent., 1985, 21 (4)

QUAES TIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.

Volume 21 Number 1 1985

CONTENTS

Bell and Bell-Rhysodini of the world. Part IV. Revisions of Rhyzodiastes Fairmaire
and Clinidium Kirby, with new species in other genera (Coleoptera: Carabidae or

Rhysodidae) 1

Book Reviews 173

RHYSODINI OF THE WORLD

PART IV. REVISIONS OF RHYZODIASTES FAIRMAIRE AND CLINIDIUM KIRBY,
WITH NEW SPECIES IN OTHER GENERA (COLEOPTERA: CARABIDAE OR

RHYSODIDAE)

Ross T. Bell
Department of Zoology
Marsh Life Science Building
University of Vermont
Burlington, VT, 05405-0086, U. S. A.

Joyce R. Bell

24 East Terrace

South Burlington, VT, 05401

U. S. A Quaestiones Entomologicae

21:1-172 1985

ABSTRACT

This paper is fourth of a series which will constitute a revision of Rhysodini of the world.
Rhyzodiastes Fairmaire and Clinidium Kirby are revised. New subgenera of Rhyzodiastes are:
Rhyzotetrops NEW SUBGENUS, rype-Rhyzodiastes janus, n. sp., one sp., Fiji ; Rhyzoarca
NEW SUBGENUS, rype-Rhyzodiastes montrouzieri (Chevrolat), three spp. Australia, New
Zealand, New Caledonia; Temoana NEW SUBGENUS, r>>/?e-Rhyzodiastes spissicornis
( Fairmaire ), 25 spp. Southeast Asia, Southwest Pacific; Rhyzostrix NEW SUBGENUS,
r>y?e-Rhyzodiastes maderiensis ( Chevrolat ), five spp. S. America. Five species from South
America remain in Rhyzodiastes s. str.

The following new species are described ( type localities indicated): Rhyzodiastes
(Rhyzotetrops) janus (FIJI, Viti Levu); Rhyzodiastes (Temoana) bipunctatus ( SOLOMON
ISLANDS, Guadalcanal, Mt. Austen); Rhyzodiastes (Temoana) indigens (SUMATRA, Si
Rambe); Rhyzodiastes (Temoana) convergens (NEW BRITAIN, Gisiluve); Rhyzodiastes
(Temoana) preorbitalis (THAILAND, Doi Sutep ); Rhyzodiastes (Temoana) vadiceps
(BORNEO?); Rhyzodiastes (Temoana) patruus (MALAYA, Johor, Sedili Kechil );
Rhyzodiastes (Temoana) denticauda (SARAWAK, Mt. Murud); Rhyzodiastes (Temoana)
propinquus (NICOBAR IS.); Rhyzodiastes (Temoana) bonsae (SUMATRA, Mt. Tenggamoes);
Rhyzodiastes (Temoana) alveus (VIET NAM, Hoa Binh); Rhyzodiastes (Temoana) fossatus
(VIET NAM, hills sw. of Kui Chau); Rhyzodiastes (Rhyzostrix) davidsoni (BRAZIL,
Amazonas, Taruma Falls); Rhyzodiastes (Rhyzostrix) nitidus (BRAZIL, Santarem);
Rhyzodiastes (Rhyzostrix) menieri (FRENCH GUIANA, Haut-Carsevenne ); Rhyzodiastes (s.
str.) pentacyclus (BRAZIL, Alto da Serra); Rhyzodiastes (s. str.) suturalis (BRAZIL, Espiritu
Santo, Sooretama); Clinidium (Mexiclinidium) championi (GUATEMALA, Quiche Mtns.);
Clinidium (Mexiclinidium) newtoni (MEXICO, Chiapas, Pueblo Nuevo); Clinidium
(Mexiclinidium) halffteri (MEXICO, Vera Cruz, Amates); Clinidium (Mexiclinidium) balli
(MEXICO, Hidalgo, 25.6 km n. of Zimapan); Clinidium (Mexiclinidium) triplehorni
(MEXICO, 1 1.3 km n.e. of Jacala; Clinidium (Mexiclinidium) iviei (MEXICO, Oaxaca, 3.2
km s. of Cerra Pelon); Clinidium (s. str.) impressum (FRENCH GUIANA, Saint Laurent du

Bell and Bell

Maroni); Clinidium (s. str.) hammondi ( COLOMBIA , Bogota); Clinidium (s. str.) howdenorum
( TRINIDAD , Morne Bleu); Clinidium (s. str.) jolyi ( VENEZUELA , Merida, La Azulita);
Clinidium (s. str.) alleni ( PANAMA , Cerro Jefe); Clinidium (s. str.) whiteheadi ( PANAMA ,
Cerro Campana); Clinidium (s. str.) trionyx ( DOMINICAN REP., Cazabita); Clinidium (s.
str.) dormans (PANAMA, Chiriqui, Finca Lerida, near Boquete); Clinidium (s. str.)
penicillatum (COLOMBIA, Dept. Valle, Represa Calima); Clinidium (s. str.) segne
(VENEZUELA, Aragua, Rancho Grande); Clinidium (s. str.) kochalkai (COLOMBIA, Casa
Antonio, Loma, Cebolleta, Sierra Nevada de Santa Marta); Clinidium (s. str.) microfossatum
(MARTINIQUE); Clinidium (s. str.) smithsonianum (DOMINICA); Clinidium (s. str.)
bechyneorum (VENEZUELA, Carabobo, Hac. Montero, Montalban); Clinidium (s. str.)
excavatum (VENEZUELA, Carabobo, Montalban Oeste); Clinidium (s. str.) pala
(VENEZUELA, Miranda, Guatopo Nat. Pk., 50 km se Caracas); Clinidium (s. str.) humile
(NEW GRANADA (Colombia or Panama)); Clinidium (s. str.) curvatum (COLOMBIA,
Santander del Norte, Oroque); Clinidium (s. str.) crater (PANAMA, Cerro Jefe, Azul Ridge);
Clinidium (s. str.) spatulatum (PANAMA, Colon, Sta. Rita ridge); Clinidium (s. str.)
moldenkei (COSTA RICA, Rincon de Osa); Clinidium (s. str.) argus (PHILIPPINES (?)
Horns of Negros); Dhysores biimpressus (TANZANIA, Usumbura, Neu Bethel); Kaveinga (s.
str.) poggii (D’ENTRECASTEAUX ISLANDS, Goodenough Is.); Grouvellina hexadon
(COMORO IS., Mayotte, Mamouzou); Yamatosa kryzhanovskyi (VIET NAM, mts. n.e. of
Thai Nguen); Yamatosa Kabakovi (VIET NAM, mountains of S ha-Pa Province);
Omoglymmius (Pyxiglymmius) opacus (SUMATRA, Padang); Omoglymmius (s. str.) gressitti
(NEW GUINEA, Wau, Mt. Missim.); Omoglymmius (s. str.) craticulus (NEW GUINEA,
Moroka); Omoglymmius (s. str.) largus (NEW GUINEA, Fly R.); Omoglymmius (s. str.) tolai
(NEW BRITAIN, Rabaul); Omoglymmius (Laminoglymmius) perplexus (SUMATRA);
Omoglymmius (Navitia) peckorum (FIJI, Viti Levu, Nandarivatu).

Clinidium beccarii Grouvelle is removed from Rhyzodiastes and returned to Clinidium (s.
str.). Rhysodes punctatolineatus Grouvelle is assigned to Arrowina.

RESUME

Cet article est la quatrieme d’une serie qui constiteront une revue taxonomique des Rhyzodini du monde. On reviset
les genres Rhyzodiastes Fairmaire et Clinidium Kirby. Les sous-genres nouveaux de Rhyzodiastes sont: Rhyzotetrops
NOUVEAU SOUS-GENRE, type-Rhyzodiastes janus, n. sp., un sp., Fiji; Rhyzoarca NOUVEAU SOUS-GENRE,
rype-Rhyzodiastes montrouzieri (Chevrolat), trois sp. Australia, Nouvelle Zelande, Nouvelle Caledonie; Temoana
NOUVELLE SOUS-GENRE, rype-Rhyzodiastes spissicornis ( Fairmaire ), 25 spp. Asie de sud-est, Pacifique de sud-ouest;
Rhyzostrix NOUVEAU SOUS-GENRE, fype-Rhyzodiastes maderiensis (Chevrolat), cinque spp., America du Sud.
Cinque species de I’Amerique du Sud restaient en Rhyzodiastes s. str.

On deer it les esp&ces nouvelles que void (en indiquant pour chacune la locality du speimen type): Rhyzodiastes
(Rhyzotetrops) janus (FIJI, Viti Levu); Rhyzodiastes (Temoana) bipunctatus (ILES DE SOLOMON, Guadalcanal, Mt.
Austen); Rhyzodiastes (Temoana) indigens (SUMATRA, Si Rambe); Rhyzodiastes (Temoana) convergens (NOUVELLE
BRETAGNE, Gisiluve); Rhyzodiastes (Temoana) preorbitalis (THAILAND, Doi Sutep); Rhyzodiastes (Temoana)
vadiceps (BORNEO?); Rhyzodiastes (Temoana) patruus (MALAYA), Johor, Sedili Kechil ); Rhyzodiastes (Temoana)
denticauda (SARAWAK, Mt. Murud); Rhyzodiastes (Temoana) propinquus (ILES DE NICOBAR); Rhyzodiastes
(Temoana) bonsae (SUMATRA, Mt. Tenggamoes); Rhyzodiastes (Temoana) alveus (VIET NAM, Hoa Binh);
Rhyzodiastes (Temoana) fossatus (VIET NAM, sur les collines au sud-ouest de Kui Chau); Rhyzodiastes (Rhyzostrix)
davidsoni (BRESIL, Amazonas, Sault de Taruma); Rhyzodiastes (Rhyzostrix) nitidus (BRESIL, Santarem);
Rhyzodiastes (Rhyzostrix) menieri (GUYANE FRANQAISE, Haut-Carsevenne); Rhyzodiastes (s. str.) pentacyclus
(BRESIL, Alto da Serra); Rhyzodiastes (s. str.) suturalis (BRESIL, Espiritu Santo, Sooretama); Clinidium
(Mexiclinidium) championi (GAUTEMALA, Quiche Mtns.); Clinidium (Mexiclinidium) newtoni (MEXIQUE, Chiapas,
Pueblo Nuevo); Clinidium (Mexiclinidium) halffteri (MEXIQUE, Vera Cruz, Amates); Clinidium (Mexiclinidium) balli
(MEXIQUE, Hidalgo, 25.6 km n. de Zimapan); Clinidium (Mexiclinidium) triplehorni (MEXIQUE, 11.3 km n.e. de
Jacala; Clinidium (Mexiclinidium) iviei (MEXIQUE, Oaxaca, 3.2 km s. de Cerra Pelon); Clinidium (s. str.) impressum

Quaest. Ent., 1985, 21 (1)

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

(GUYANE FRANQAISE, Saint Laurent du Maroni); Clinidium (s. str.) hammondi (COLOMBIE, Bogota ); Clinidium (s.
str.) howdenorum (TRINITE, Morne Bleu); Clinidium (s. str.) jolyi ( VENEZUELA , Merida, La Azulita); Clinidium (s.
str.) alleni ( PANAMA , Cerro Jefe); Clinidium (s. str.) whiteheadi ( PANAMA , Cerro Campana); Clinidium (s. str.)
trionyx (REP. DOMINICAINE, Cazabita); Clinidium (s. str.) dormans ( PANAMA , Chiriqui, Finca Lerida, pres de
Boquete); Clinidium (s. str.) penicillatum ( COLOMBIE , Dept. Valle, Represa Colima); Clinidium (s. str.) segne
( VENEZUELA , Aragua, Rancho Grande); Clinidium (s. str.) kochalkai ( COLOMBIE , Casa Antonio, Loma Cebolleta,
Sierra Nevada de Santa Marta); Clinidium (s. str.) microfossatum ( MARTINIQUE ); Clinidium (s. str.) smithsonianum
(DOMINICA); Clinidium (s. str.) bechyneorum (VENEZUELA, Carabobo, Hac. Montero, Montalban); Clinidium (s.
str.) excavatum ( VENEZUELA , Carabobo, Montalban Oeste); Clinidium (s. str.) pala (VENEZUELA, Miranda,
Guatopo Nat. pk., 50 km se Caracas); Clinidium (s. str.) humile (NEW GRANADA (Colombie ou Panama)); Clinidium
(s. str.) curvatum (COLOMBIE, Santander del Norte, Oroque); Clinidium (s. str.) crater (PANAMA, Cerro Jefe, Azul
Ridge); Clinidium (s. str.) spatulatum (PANAMA, Colon, Sta. Rita ridge); Clinidium (s. str.) moldenkei (COSTA RICA,
Rincon de Osa); Clinidium (s. str.) argus (PHILIPPINES (?) Horns of Negros); Dhysores biimpressus (TANZANIA,
Usumbura, Neu Bethel); Kaveinga (s. str.) poggii (ILES D’ENTRECASTEAUX, Goodenough Is.); Grouvellina hexadon
(ILES COMORES, Mayotte, Mamouzou); Yamatosa kryzhanovskyi (VIET NAM, Monts du nordest de Thai Nguen);
Yamatosa kabakovi (VIET NAM, Monts de la province de Sha-Pa); Omoglymmius (Pyxiglymmius) opacus (SUMATRA,
Padang); Omoglymmius (s. str.) gressitti (NOUVELLE GUINEE, Wau, Mt. Missim.); Omoglymmius (s. str.) craticulus
(NOUVELLE GUINEE, Moroka); Omoglymmius (s. str.) largus (NOUVELLE GUINEE, Fly R.) Omoglymmius (s. str.)
tolai (NOUVELLE BRETAGNE, Rabaul); Omoglymmius (Laminoglymmius) perplexus (SUMATRA); Omoglymmius
(Navitia) peckorum (FIJI, Viti Levu, Nandarivatu).

On deplacet Clinidium beccarii Grouvelle de Rhyzodiastes et le retournet a Clinidium (s. str.). On attribuet Rhysodes
punctatolineatus Grouvelle a Arrowina.

Table of Contents

Introduction 3

Sources of material 4

. Genus Rhyzodiastes 6

New Subgenus Rhyzotetrops (Figs. 1, 5, 9, 10) 6

New Subgenus Rhyzoarca (Figs. 2-4, 6-8) 8

New Subgenus Temoana (Figs. 1 1-50, 52-54) 1 1

New Subgenus Rhyzostrix (Figs. 51, 55-65) 48

Subgenus Rhyzodiastes sensu stricto (Figs. 74) 54

Genus Clinidium Kirby 59

Subgenus Mexiclinidium (Figs. 75-86, 96-102) 60

Subgenus Protainoa (Fig. 87) 69

Subgenus Tainoa (Figs. 88-95) 70

Subgenus Arctoclinidium (Figs. 103-123) 75

Subgenus Clinidium sensu stricto (Figs. 124-213) 93

Additions, Corrections to Parts I— 1 1 1 including additional species (Figs. 214-234) 149

Acknowledgements 166

References 166

Index to Names of Taxa 169

INTRODUCTION

This paper, the fourth in a series of five, consists of revisions of the genera Rhyzodiastes
Fairmaire and Clinidium Kirby, together with descriptions of new species in several of the
genera treated in earlier parts of the series. The subgenera of Rhyzodiastes are named and
defined.

Quaest. Ent., 1985,21 (1)

Bell and Bell

SOURCES OF MATERIAL

The following abbreviations designate collections cited in this paper. The names in
parentheses are the curators of the respective institutions.

AIM

ALB

AMNH

AMS

ARK

BMNH

BMS

BPBM

BPM

BSL

BSRI

CAG

CAS

CMP

CNHM

DSIR

FLA

GEN

GVA

GLP

INPA

ISNHS

LCC

LEI

LEN

Auckland Institute and Museum, New Zealand (K.A .J. Wise)

University of Alberta, Edmonton, Canada (G. E. Ball)

American Museum of Natural History, New York (L. Herman)

Instituut voor Taxonomische Zoologie, Amsterdam, Netherlands (J.
Duffels)

U.S. Dept, of Agriculture, Harrisburg, PA (K. Valley)

University of Arkansas, Fayetteville (E. P. Rouse)

S.F. Austin State University, Nacogdoches, Texas (W. W. Gibson)

British Museum, Natural History, London (R. Pope)

Buffalo Museum of Science, NY (H. W. Charnley)

Bernice P. Bishop Museum, Honolulu, Hawaii (G. Samuelson)

Barry P. Moore, Canberra City, Australia
Naturhistorisches Museum, Basel, Switzerland (W. Wittmer)
Biosystematics Research Institute, Ottawa, Canada (A. Smetana)

U.S. Dept, of Agriculture, Sacramento, CA (F. G. Andrews)

California Academy of Sciences, San Francisco, CA (D. Kavanaugh)
Carnegie Museum of Natural History, Pittsburgh, PA (G. Wallace)

Field Museum of Natural History, Chicago, IL (H. Dybas)

Cornell University, Ithaca, NY (L. L. Pechuman)

Dayton Museum, Ohio (A. J. Koestner)

Department of Scientific and Industrial Research, Auckland, N.Z. (J.
Watt)

Daniel K. Young, E. Lansing, MI

U.S. Dept, of Agriculture, Gainesville, FL (R. Woodruff)

University of Georgia, Athens, GA (C. L. Smith)

Museo Civico di Storia Naturale “G. Doria”, Genoa (R. Poggi)

Museum d’Histoire Naturelle, Geneva, Switzerland (I. Lobl)

Gary L. Peters, Corvallis, OR

Harry J. Lee, Fairview Park, OH

Iowa State University, Ames IA (R. Miller)

Instituto Nacional de Pesquisas de Amazonia, Manaus, Brazil (N. D.
Penny)

Illinois State Natural History Survey, Urbana IL (M. Sanderson)

Indiana University, Bloomington IN
Karl Stephan, Tucson, AZ
Kagoshima University, Japan

Los Angeles County Natural History Museum, CA (C. L. Hogue)

Lincoln College, Canterbury, N.Z., (R.M. Emberson)

Rijksmuseum von Natuurlijke, Historic, Leiden, Netherlands (J. Krikken)
Academy of Sciences, Leningrad USSR (O. Kryzhanovskij)

Louisiana State University, Baton Rouge LA (J. B. Chapin)

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

LUN Zoological Institute, Lund, Sweden (R. Danielsson)

MAI Michael A. Ivie, Columbus, OH

MAY University of Puerto Rico, Mayaguez (J. Ramos)

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA

(J. Lawrence)

MN University of Minnesota, St. Paul MN (P. J. Clausen)

MNHB Museum fur Naturkunde der Humboldt-Universitat, Berlin, DDR (F.

Hieke)

MNHN Museum National d’Histoire Naturelle, Paris, France (A. Descarpentries)
MO University of Missouri, Columbia MO (W. R. Enns)

MRAC Musee Royal de l’Afrique Centrale, Tervuren, Belgium (P. Basilewsky)
MSU Michigan State University, E. Lansing MI

MZSP Museu de Zoologia da Universidade de Sao Paulo, Brazil (U.R. Martins)
NC North Carolina State University, Raleigh NC (D. A. Young)

NMNH U.S. National Museum of Natural History, Washington, D. C. (T. Erwin)
NMNZ National Museum of New Zealand, Wellington (R. G. Ordish)

NMW Naturhistorisches Museum Wien, Austria (F. Janczyk)

OK Oklahoma State University, Stillwater OK (W. A. Drew)

OS Oregon State University, Corvallis OR (G. L. Peters)

OSFS Oregon State Forest Sciences Collection, Corvallis OR
OSU Ohio State University, Columbus OH (C. A. Triplehorn)

OUA Ohio University, Athens, OH (H. Seibert)

PA Academy of Sciences, Philadelphia, PA (D. C. Rentz)

PK Paul Kittle, Southeast Missouri State University, Cape Girardeau

PU Purdue University, Lafayette, IN

QW Quentin Wheeler, Columbus, OH

RCG R. C. Graves, Bowling Green, OH

SATO Masataka Sato, Nagoya, Japan

SDA U.S. Dept, of Agriculture, Brookings. SD (V. M. Kirk)

SI Southern Illinois University, Carbondale, IL (J. E. McPherson)

TB Thomas Barr, University of Kentucky, Lexington KY
UCB University of California, Berkeley CA (J. A. Chemsak)

UCD University of California, Davis CA •

UD University of Delaware, Newark DE (P. P. Burbutis)

UI University of Illinois, Urbana IL (R. Selander)

UK University of Kansas, Lawrence KS (G. W. Byers)

UL University of Louisville, KY (C. V. Coveil)

UM University of Michigan, Ann Arbor, MI (I. J. Cantrall)

UN University of Nebraska, Lincoln NB (B. C. Ratcliffe)

UNH University of New Hampshire, Durham NH (D. Chandler)

UT Utah State University, Logan, UT (W. J. Hanson)

UVM University of Vermont, Zoology Department, Burlington, VT
UW University of Wisconsin, Madison WI (J. R. Baker)

VEN Universidad Central de Venezuela, Maracay (L. J. Joly)

VP Virginia Polytechnic Institute, Blacksburg, VA (M. Kosztarab)

WR William Rosenberg

Quaest. Ent., 1985,21 (1)

Bell and Bell

WRS Walter R. Suter, Carthage College, Kenosha, WI
WS Washington State University, Pullman WA (W. J. Turner)

GENUS RHYZODIASTES FAIRMAIRE 1895
Type species. — Rhyzodiastes parumcostatus Fairmaire 1868

Description. — Part I: 61-62. Most species have two spurs on each of the middle and hind tibiae, as stated in the
definition in Part I, but two species from Borneo each have only a single tibial spur.

This genus and Clinidium both have the striation strongly reduced and heterogeneous. The striae differ strongly in
depth, width, degree of punctation and pollinosity. Since striae disappear from both the disc and the margin of the elytron,
it would be quite confusing to refer to them by numbers, as we have in other genera. Accordingly, we designate each stria
with a name, and define it in terms of its spatial relationship with other parts of the elytron. The sutural stria is the most
medial one, closely paralleling the suture. The parasutural stria is the next one laterally. The intercalary stria is lateral to
the parasutural and medial to the sub-apical tubercle. In Rhyzodiastes it occurs only in Subgenus Rhyzotetrops. The
intratubercular stria is lateral to the parasutural and to the intercalary, if present. It can be identified by the fact that its
apex passes between the subapical and apical tubercles. The supramarginal stria is absent from Rhyzodiastes , but is
present in some subgenera of Clinidium. It is lateral to the intratubercular and dorsad to the marginal stria. The marginal
stria is the outermost stria visible in a dorsolateral view. It can be identified by the fact that its apex passes ventrad to the
apical tubercle, where it attains the suture in most species. The submarginal stria lies on the elytral epipleura, between the
marginal stria and the edge of the elytron. Posteriorly, it ends near the fifth or sixth abdominal sternum.

KEY TO SUBGENERA

1 Each compound eye divided into two ocelliform structures; elytron with

intercalary stria present Rhyzotetrops new subgenus, p. 6

V Compound eye entire, crescentic or hemispherical; elytron with intercalary

stria absent 2

2 (1') Paramedian grooves much closer together at middle of length than at base

or apex; outer carina much wider at middle of length than at either end;

tufts of minor setae present on antennal Segments VII-X

Rhyzoarca new subgenus, p. 8

2' Paramedian grooves not closer together at middle than at either end; outer
carina not much wider at middle than at either end; tufts of minor setae
present on Segments IV-X, V-X, or VI-X 3

3 (2') Inner carina of pronotum with lateral margin as distinct as medial margin;

inner carina abruptly separated from paramedian groove which is entirely

pollinose Rhyzodiastes sensu stricto, p. 54

3' Inner carina of pronotum with lateral margin sloped gradually into
paramedian groove; paramedian groove with pollinosity in most species
limited to its border with outer carina (R. pollinosus an exception) 4

4 (30 Eye enlarged, hemispheric: gena below eye with curved band of pollinosity;

basal margin of protonum with narrow strip of pollinosity

Rhyzostrix new subgenus, p. 48

4' Eye narrow, crescentic; gena below eye glabrous; basal margin of pronotum
not pollinose Temoana new subgenus, p. 1 1

SUBGENUS RHYZOTETROPS NEW SUBGENUS

Type species. — Rhyzodiastes ( Rhyzotetrops ) janus new species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Description. — Apical stylet of antenna acuminate; tufts of minor setae present on Antennal Segments VII-X;
compound eye divided into two ocellus-like structures, one directed anteriolaterally, the other posterio-laterally; clypeal
setae present; pronotum with median groove strongly dilated; inner carinae sloped gradually into paramedian grooves;
latter glabrous except for pollinose strip along medial margin of outer carina; latter with row of setae; outer carina curved,
rather narrow, of even width; elytron with intercalary stria present; intratubercular stria obsolete except for apex, which is
impressed; all femora with many long setae. This subgenus is restricted to Fiji.

Only one species is known.

Rhyzodiastes ( Rhyzotetrops ) janus new species
(Figs. 1,5,9, 10)

Type Material. — HOLOTYPE male, labelled: “Vitilevu, Fiji, 6 M.W. Nandarivatu, Mba, IX- 16-38, Coll. Y.
Kondo” (BPBM). PARATYPES one male, same data as holotype (BPBM); one female, labelled: “Nandarivatu, Viti
Levu, Fiji, IX-10-38, 3700’, rotten log, coll. E. G. Zimmerman” (BPBM); one female, same locality and collector but
dated “IX-6-38, 3600’, beating shrubbery” (BPBM); one female, labelled: “Navai, Fiji Isl. Mann” (MCZ); one female,
labelled: “Viti Levu, Fiji, Nadarivatu, W. M. Mann” (MCZ); one female, labelled: “Fiji, Viti Levu, Nadarivatu, 13-xi-74,
coll. B. P. Moore” (BPBM).

Description. — 5. 9-6. 3 mm. Tuft of minor setae very small on Segment VII of antenna; those of VIII-X larger;
basal setae of antennae on Segments VII-X; Segments I-IV with subapical pollinose bands; outer antennal segments nearly
spherical; Segment XI wider than Segment X, and over twice as long as latter.

Head slightly longer than wide; median lobe very short, transverse, its tip obtuse, far anterior to eyes; antennal lobe
glabrous, but divided by anterior pollinose extension of postantennal groove; temporal lobe 1.5 longer than wide, broadly
rounded medially, with broad pilose fringe across basal margin; orbital groove broad, distinct, but incomplete posteriorly,
its base just posterior to posterior eye; gena with vertical bar of pollinosity ending ventrad to space separating anterior and
posterior eyes (Fig. 10); one temporal seta.

Pronotum with length/greatest width 1.5, unusually large compared to elytra, more than 0.55 as long and nearly as
wide as elytra; widest near middle, sides rather weakly curved; apex truncate; base rounded; median groove broadly
dilated, forming about 0.2 of pronotal width; median groove entirely glabrous; anterior median pit near to anterior margin;
posterior median pit at basal 0.25 of length; both median pits pollinose, conspicuous; median groove posterior to posterior
median pit rather deep, but clearly shallower than remainder of pit; inner carinae rather narrow, sloped gradually into
paramedian groove except in basal 0.25, where separated from paramedian groove by vertical scarp; paramedian grooves
broad, glabrous except for narrow pollinose scarp bounding outer carina; basal impression conspicuous, glabrous except for
small circular pollinose impression at middle; basal impression bounded posteriorly by glabrous ridge which is continuation
of outer carina; outer carina curved, convex, narrow, of nearly even width, row of nine to 12 setae present in pollinose
median scarp of outer carina; marginal groove entirely absent; notopleural suture entirely pollinose; sternopleural groove
entirely absent.

Elytra rather short, sides nearly parallel; parascutellar pits large but widely separated from one another; base of elytra
without transverse pollinose band; Interval I flat; sutural stria fine, scarcely pollinose, with fine, widely separated
punctures; Interval II somewhat convex, sloped laterally; parasutural stria finely punctate, narrowly pollinose, slightly
deeper than sutural stria, curved medially at base, meeting sutural stria at apex; apical depression largely glabrous but
with posteriolateral pollinose strip; Interval III convex, its base bent medially, forming prominent angle; intercalary or
third stria deeper than parasutural stria, with moderately broad strip of pollinosity; Interval IV with medial margin
distinct but lateral one obsolete, continuous posteriorly with subapical tubercle; intratubercular stria incomplete, entirely
effaced in basal 0.33, for most of remainder represented by row of minute, widely spaced punctures, its apex impressed,
pollinose; apical tubercles swollen, broadly in contact at suture; marginal stria complete, pollinose, linear, becoming dilated
below apical tubercle; submarginal stria entire, extending to middle of Sternum V; intercalary stria with about 10 setae;
impressed apex of intercalary stria with two or three setae; medial face of apical tubercle with one or two setae; marginal
stria with continuous row of about 15 setae, these more closely spaced near apex. (Fig. 5)

Metasternum not sulcate; abdominal sterna with transverse sulci narrowly interrupted at midline; slight development
of lateral pit on Sternum IV in both sexes (Fig. 9); tibia slender, posterior spurs slightly smaller than anterior ones; male
with anterior tibia not dentate nor tuberculate; male trochanters not modifed; calcars small, acutely pointed.

The presence of an intercalary stria is unique within the genus, as is the strange divided eye.
A similar eye has evolved independently in Clinidium (5. str.) beccarii and its relatives.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

SUBGENUS RHYZOARCA NEW SUBGENUS

Type species. — Rhyzodes montrouzieri Chevrolat 1875

Description. — Tufts of minor setae present on antennal Segments VII-X; clypeal setae absent; eyes entire,
narrowly crescentic; temporal, pronotal, elytral setae entirely absent; pronotum relatively broad, subquadrate, with lateral
margins slightly curved; base, apex truncate; median groove linear or absent; anterior median pit absent; posterior median
pit small, shallow or absent; paramedian grooves closest together at middle of length, strongly curved; each paramedian
groove with large pilose pit at anterior and posterior end; disc of pronotum depressed between anterior lateral pits; outer
carina oval, broad at middle, tapered both anteriorly and posteriorly; marginal, submarginal grooves absent; intercalary
stria absent; metasternal sulcus absent in R. montrouzieri , R. proprius, ventral surface of R. burnsi not studied; female
with shallow, lateral pits on abdominal Sternum IV; male front femur with ventral tooth; calcar small, acutely pointed.

This subgenus occurs in Australia, New Zealand, and New Caledonia.

Phylogeny. — Of the three species, R. proprius and R. burnsi , of New Zealand and
Australia, respectively, are clearly more closely related than either is to R. montrouzieri of
New Caledonia.

KEY TO SPECIES

1 Apex of intratubercular stria impressed; median groove of pronotum

absent; median lobe of head very short

R. montrouzieri (Chevrolat), p. 8

1' Apex of intratubercular stria obsolete, subapical and apical tubercles thus
not separated; median groove of pronotum linear but distinct; median lobe
of head longer 2

2 (1') Sutural stria absent except for extreme base; parasutural stria impressed,

impunctate; posterior median pit absent R. proprius (Broun), p. 9

2' Sutural stria entire, impressed, coarsely punctate; parasutural stria
coarsely punctate; posterior median pit small but distinct
R. burnsi (Oke), p. 9

Rhyzodiastes ( Rhyzoarca ) montrouzieri (Chevrolat 1875) NEW COMBINATION

(Figs. 2, 8)

Rhyzodes montrouzieri Chevrolat 1875: 182.

Rhyzodiastes montrouzieri (Chevrolat) Bell and Bell 1978

Type Material. — We have not located type material for this species. It is easily recognized from the description
and the type locality.

Description. — Length 5. 0-7. 5 mm. Median lobe of head short, its tip opposite anterior margin of eye; length of
eye about 0.8 of length of temporal lobe.

Pronotum relatively short, length/greatest width 1.35; anterior angles acute, slightly prominent; lateral margin not
sinuate anterior to hind angle; base of pronotum relatively narrow, its width about 0.8 of greatest width of pronotum;
median groove entirely absent; posterior median pit represented by shallow pit.

Elytron without basal pollinosity; sutural stria fine, linear, pollinose, impunctate, its base dilated, (Fig. 8); its apical
0.66 entirely effaced; parasutural stria entire, deep, impressed, pollinose; intratubercular stria complete, impressed,
impunctate, pollinose, its apex separating apical and subapical tubercles; marginal stria represented by row of minute
punctures, its apex not impressed; hind calcar of male acute, its dorsal margin straight.

This species is easily recognized by the short median lobe of the head, the impressed apex of
the intratubercular stria, and the absence of the median groove of the pronotum.

Distribution. — Confined to New Caledonia. We have seen the following specimens: 13 males,
five females, Col des Roussettes, 450-550 m., 4-6-II-63, G. Kuschel, C. Yoshimoto, J. L. Gressitt (BPBM): one female,
Hanna, Foret de Thi, VII-16-1958, B. Malkin (CNHM); six males, three females, Mt. Koghi, Foret de Thi, 530 m.,
8-III- 1 96 1 , J. Sedlacek (BPBM); five males, four females, Noumea, Aug. 27-1944, Wilfred Crabb (NMNH); one male.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Mt. Chapeau, Gendarme rain forest, 1 3-VIII- 1 944, J. C. Herron (OUA)

Rhyzodiastes (Rhyzoarca) proprius (Broun 1880) NEW COMBINATION

(Figs. 3, 6)

Rhysodes proprius Broun 1880: 216.

Rhysodes probius Lewis 1888 (error).

Clinidium ( Rhyzodiastes ) proprium (Broun) Grouvelle 1903.

Rhyzodiastes proprius (Broun) Bell and Bell 1978.

Type Material. — Not studied. According to the original description, the type locality is Parua, New Zealand,
and there were three syntypes.

Description. — Length 6.0-8.0 mm. Median lobe of head elongate, its tip posterior to eye; eye small, about 0.5 of
length of temporal lobe.

Pronotum moderately long, length/greatest width 1.45; anterior angles obtuse; lateral margin shallowly sinuate
anterior to hind angles; base of pronotum very broad, scarcely narrower than greatest width; median groove fine,
inconspicuous, slightly abbreviated at base; posterior median pit absent.

Elytron without basal pollinosity; sutural interval of most specimens with elongate, very finely pollinose depression just
posterior to middle of elytron, in some of the smallest specimens this depression entirely absent; sutural stria represented
only by scarp-like basal portion, remainder entirely absent; parasutural stria pollinose, impunctate, deep, entire, its middle
0.33 slightly dilated; intratubercular stria impressed, impunctate for most of its length, its apex obsolete, not separating
apical and subapical tubercle; marginal stria absent except for short impressed apical portion which is ventrad to apical
tubercle (Fig. 6); hind calcar of male with its dorsal side convex, its apex a small but sharp point.

This species is similar to the next, but is easily separated by the absence of the sutural stria.
Distribution. — Confined to the North Island of New Zealand. We have studied the

following Specimens: one female, Auckland, coll. E. S. Gourley, 1970 (DSIR); one female, Auckland, Orere Bush,
rotten log, 10-1-1957 (DSIR); one female, Bayswater, 9-13, #381, T. Broun colln. (DSIR); one female, Clevedon, under
log, 2-4-1956, coll. J. C. Watt (DSIR); one male, one female, Huia, Auckland, ex rotten kauuka stump, 4-2-66, coll. J. C.
Watt (DSIR); two females, Little Barrier, 1913, 117, coll. H. Swale (BMNH); one male, one female, Pollok, Auckland,
coll. P. & M. Johns, 7-1-1964 (LCC); one male, one female, Rawhiti, forest remnant. Bay of Islands, 4-1-1969, coll. K. A.
J. Wise (AIM); one female, Spirits Bay, Waipuna Stream, 9-XI-67, coll. J. I. T. and J. McB., litter (DSIR); one male,
Tiki-Tiki, 18-1-63 (NMNZ); one female, Waitakere, Waitemata Co., C. E. Clarke colln. (AIM); one female, Whangarei,
18-20-3-31, coll. E. S. Gourley (DSIR); one male, one female, Whangarei Heads, colln. C. E. Clarke (AIM); four males,
Whangarei, Pukerui Hills, 21-11-44, coll. B. Given (DSIR); one male, Whangarei, Three Mile Bush, 24-11-44, coll. B.
Given (DSIR); one male, Whangarei, Western Hills, 13-IX-1956, coll. R. A. Crowson (CAS); one male, two females,
Whangarei, Whau Valley, 1 1-8-28, coll. Fairburn (DSIR).

Rhyzodiastes ( Rhyzoarca ) burnsi (Oke 1932) NEW COMBINATION

(Figs. 4, 7)

Rhysodes burnsi Oke 1932: 148-149.

Rhyzodiastes burnsi (Oke) Bell and Bell 1978.

Type Material. — HOLOTYPE, female, from AUSTRALIA: New South Wales, Mt. Wilson, in log with ants,
coll. C. Oke. We have not studied the type, but have seen a good enlarged photograph of it, kindly sent by B. Moore.

Description. — (Based on the original description and the photograph.) Length 7 mm. (Chaetotaxy not studied.)
Median lobe of head elongate, its tip opposite posterior margin of eye; eye larger than in R. proprius-, temporal lobes more
rounded posteriorly than in latter species; pronotum moderately long, length/greatest width about 1.4; front angles obtuse;
lateral margin shallowly sinuate anterior to hind angles; base of pronotum very broad, only slightly narrower than greatest
width; median groove distinctly impressed; posterior median pit distinct.

Sutural interval without pollinosity but second interval with small pollinose spot at apical fourth (Fig. 7); sutural stria
and parasutural striae both distinctly impressed, coarsely punctate; intratubercular stria shallowly impressed, coarsely
punctate; its apex obsolete, not separating subapical and apical tubercles; marginal stria absent; according to original
description, metasternum is sulcate.

The well-developed sutural striae and the coarse punctures of the sutural and parasutural
striae separate this species from R. proprius. Oke suggested that the enlarged anterior lateral
pits of the pronotum are trichomes and that the species might be myrmecophilous. This has
neither been confirmed nor disproven. If true of this species, it is probably true of the entire

Quaest. Ent., 1985, 21 (1)

Plate 1. Figs. 1, 5, 9, 10. Genus Rhyzodiastes, new Subgenus Rhyzotetrops R. (R.) janus new species. Fig. 1, Head and
pronotum, dorsal aspect; Fig. 5, Left elytron, dorsal aspect; Fig. 9, Metasternum and abdomen, left half; Fig. 10, Head,
lateral aspect. Figs. 2-4, 6-8. Genus Rhyzodiastes , new Subgenus Rhyzoarca. Figs. 2-4, Head and pronotum, dorsal
aspect; Fig. 2, R. (R.) montrouzieri (Chevrolat); Fig. 3, R. (R.) proprius (Broun) (drawn from photograph); Fig. 4, R. (R.)
burnsi (Oke); Figs. 6-8, Left elytron, dorsal aspect; Fig. 6, R. (R.) proprius (Broun); Fig. 7, R. (R.) burnsi (Oke); Fig. 8,
R. (R.) montrouzieri (Chevrolat).

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

subgenus.

SUBGENUS TEMOANA NEW SUBGENUS
Type species. — Clinidium spissicorne Fairmaire 1895.

Description. — Apical stylet of antenna present, though minute in some species; tufts of minor setae present on
antennal Segments IV-X or V-X; clypeal setae present; compound eye narrow, crescentic; genae glabrous ventrad to eye;
inner carinae of pronotum sloping gradually to paramedian groove; pollinosity on most species limited to border between
paramedian groove and outer carina (more extensive in R. pollinosus)\ paramedian grooves straight or slightly curved;
outer carina not greatly broadened at middle; elytron with intercalary stria absent.

This very large subgenus is most similar to Rhyzostrix, but lacks the enlarged eyes, the
basal pollinosity of the pronotum and the genal pollinosity of the latter subgenus. In addition,
all species of Rhyzostrix have coarsely punctate elytral striae. In Temoana, most species have
the striae impunctate or nearly so, but R. sulcicollis is an exception, having strial punctures as
coarse as those of Rhyzostrix.

Temoana ranges from the Caroline Islands, the Solomon Islands, and Australia westward to
the Andaman Islands, eastern India, and Formosa.

Phylogeny. — We divide this large and complex subgenus into seven species groups as
follows:

I. singulars group - antennal tufts commence on Segment V; temporal setae
present; orbital groove complete. Seven species, Australia, Solomon
Islands, New Guinea, New Britain, Celebes, Formosa, Sumatra.

II. mishmicus group - antennal tufts commence on Segment V; temporal setae
absent; orbital groove reduced or absent. Three species, southeast Asia.

III. sulcicollis group - antennal tufts commence on Segment V; marginal stria
not impressed. Three species, Carolina and Molucca Islands.

IV. pollinosus group - antennal tufts commence on Segment V; elytra
extensively pollinose, with narrow raised carinae; three or more temporal
setae. One species, Caroline Islands.

V. myopicus group - antennal tufts begin on Segment IV; median groove of
pronotum linear. Five species, Malay Peninsula, Borneo.

VI. gestroi group - antennal tufts commence on Segment IV; median groove of
pronotum moderately dilated. Three species, Sumatra, Andaman Islands.

VII. fairmairei group - antennal tufts begin on Segment IV; median groove very
strongly dilated. Three species, southeast Asia.

The interrelationships among these groups are not clear. The absence of an antennal tuft
from Segment IV appears to us to be a plesiomorphic (primitive) character in the singularis
and sulcicollis groups, since a low number of tufts (Segments VII-X) marks Subgenus
Rhyzotetrops , which is the only Subgenus of Rhyzodiastes to retain the intercalary stria, and
hence can be viewed as the most primitive Rhyzodiastes. In the related genus Clinidium , the
least modified subgenera, Arctoclinidium and Mexiclinidium also have antennal tufts only on
Segments VII-X, while in the most advanced Subgenus, Clinidium s. str., most species have an
increased number of tufts. It cannot be guaranteed, however, that the number of tufts has never
decreased. In particular, the mishmicus group, without a tuft on Segment IV, shows close
resemblances with the myopicus group which has such a tuft. The two groups are sympatric,
and it is entirely possible that they are related, the tuft having secondarily been lost in the

Quaest. Ent., 1985,21 (1)

Bell and Bell

former group. If this is true, then the mishmicus, myopicus, gestroi, and fairmairei groups
might represent a single phyletic line, embracing all the species west of Wallace’s Line. The
singularis group is bound together mainly by characters which could be considered to be
plesiomorphic, and perhaps it is not a true phyletic unit. R. indigens , which we provisionally
place in the singularis group, is really very similar to R. bonsae of the gestroi group, and is
perhaps an additional species which has secondarily lost the tuft from Segment IV. The
monotypic pollinosus group is enigmatic. The pollinose surface and carinate intervals set it
apart from all other Temoana. It appears superficially to be isolated, but it might be an
offshoot of the sulcicollis group which has undergone extensive modification.

KEY TO SPECIES

2 (1)

3 (2')
3'

4 (3)
4'

5 (40
5'

6 (30
6'

7 (6)
r

8 (70
8'

9 (60
9'

10 (9)
10'

11 (90

Antennal tufts present on Segments V-X (absent from Segment IV) 2

Antennal tufts present on Segments I V-X 15

Elytral intervals narrowly carinate; areas between carinae entirely
pollinose; orbital groove with one to three setae {pollinosus group)

R. pollinosus Bell and Bell, p. 14

Elytral intervals not narrowly carinate; pollinosity limited to narrow striae;
orbital groove with one seta or without (in one species, with one additional

temporal seta remote from orbital groove) 3

Marginal stria of elytron not impressed {sulcicollis group) 4

Marginal stria impressed throughout its length 6

Sutural stria of elytron impressed for basal 0.66 of its length; parasutural
stria with several setae R. raffrayi (Grouvelle), p. 15

Sutural stria not impressed, represented by row of coarse punctures or

entirely absent; parasutural stria without setae 5

Sutural and marginal striae each represented by row of coarse punctures;

marginal groove of pronotum present R. sulcicollis (Grouvelle), p. 24

Sutural, marginal striae absent; marginal groove of pronotum absent . .

R. maritimus Bell and Bell, p. 25

Orbital groove incomplete or absent; temporal seta absent {mishmicus

group) 7

Orbital groove complete; one or two temporal setae {singularis group) 9

Orbital groove present, abbreviated at middle of eye

R. mishmicus (Arrow), p. 32

Orbital groove absent 8

Metasternum sulcate R. waterhousei (Grouvelle), p. 33

Metasternum not sulcate R. preorbitalis new species, p. 34

Sutural stria absent or represented only by a few punctures 10

Sutural stria impressed 11

Median groove of pronotum obsolete; parasutural stria setose

R. singularis (Heller), p. 27

Median groove impressed; parasutural stria without setae

R. guineensis (Grouvelle), p. 26

Two temporal setae present; one in center of lobe, one on orbital groove

R. bipunctatus new species, p. 28

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

1 1' One temporal seta in orbital groove 12

12 (IT) Temporal lobes convergent posteriorly

R. confer gens new species, p. 3 1

12' Temporal lobes not convergent posteriorly 13

13 (12') Outer carina of pronotum strongly narrowed anteriorly, pronotum without

distinct front angles; outer antennal segments twice as wide as long,

cylindrical R. indigens new species, p. 3 1

13' Outer carina only slighly narrowed anteriorly, truncate at apex; pronotum
with distinct front angles; outer segments sphaeroid, about 1.5 wider than
long 14

14 (13') Apex of pronotum slightly narrower than base; median groove dilated, as

wide as anterior median pit; antennal Segments VIII-X with basal setae

R. rimoganensis (Miwa), p. 28

14' Apex of pronotum broader than base; median groove nearly linear, much
narrower than anterior median pit; antenna without basal setae
R. mirabilis (Lea), p. 30

15 (1') Median groove linear between median pits {myopicus group) 16

15' Median groove dilated between median pits 21

16 (15) Parasutural stria straight to base; outer carina broad, truncate anteriorly

R. myopicus (Arrow), p. 37

16' Parasutural stria bent medially, crossing base of Interval II; outer carina

narrow, not truncate anteriorly 17

17 (16') Preapical tubercles not prominent, widely separated from one another;

apical tubercle in lateral view not separated from preapical by deep notch 18

17' Preapical tubercles prominent, tooth-like, separated from one another by
width of sutural interval or less; apical tubercle in lateral view separated
from preapical tubercle by deep notch 20

18 (17) Pronotum without marginal groove; middle, hind tibiae each with two

equal spurs, without curved apical process

R. vadiceps new species, p. 38

18' Pronotum with complete, pollinose marginal groove; middle, hind tibiae

each with one spur, plus curved apical process laterad to spur 19

19 (18') Temporal seta present; basal setae of antenna absent; ventral surfaces of

femora of male tuberculate R. patruus new species, p. 39

19' Temporal seta absent; basal setae of antenna present; ventral surface of
femora of male not tuberculate R.frater (Grouvelle), p. 38

20 (17') Apical tubercles contiguous; head as broad as long

R. bifossulatus (Grouvelle), p. 40
20' Apical tubercles widely separated from one another; head 1.5 longer than
broad R. denticauda new species, p. 41

21 (15') Median groove only moderately dilated, narrower than anterior median pit

(gestroi group) 22

21' Median groove very broadly dilated {fairmairei group) 24

22 (21) Outer carina of pronotum broad, flat, sloped laterally; temporal lobes

strongly convergent posteriorly R. gestroi (Grouvelle), p. 42

22' Outer carina very narrow, strongly cariniform; temporal lobes not

Quaest. Ent., 1985,21 (1)

Bell and Bell

23 (22')
23'

24 (21')
24'

25 (240
25'

26 (250
26'

convergent posteriorly

Temporal lobes divergent posteriorly; cauda of elytra absent

R. propinquus new species, p. 43
Temporal lobes evenly rounded medially; cauda of elytra distinct

R. bonsae new species, p. 43
Median groove of pronotum dilated at middle, resembling keyhole;
temporal seta absent R.fairmairei (Grouvelle), p. 44

Median groove of pronotum long-oval; temporal seta present

Metasternum sulcate; spurs of middle and hind tibiae equal; sutural,

parasutural striae, Interval IV, apical tubercle setose

R. spissicornis (Fairmaire), p. 45

Metasternum not sulcate; tibial spurs very unequal; elytral setae limited to

parasutural stria or absent

Parasutural stria setose; deep portion of median groove six times longer
than wide R.fossatus new species, p. 47

Parasutural stria not setose; deep portion of median groove five times
longer than wide R. alveus new species, p. 46

THE POLL1NOSUS GROUP

The single species in this group differs from all other species of Temoana in having the
elytron almost entirely pollinose except for three narrow glabrous carinae, and in having in
most specimens more than one seta in the orbital groove.

Rhyzodiastes ( Temoana ) pollinosus Bell and Bell 1981 NEW COMBINATION

(Figs. 11, 19)

Rhyzodiastes pollinosus Bell and Bell 1981: 61-63.

Type Material. — HOLOTYPE male (CAROLINE ISLANDS), labelled: “Yap Group; Yap Island, Jul-Au 50,
R. J. Goss” (BPBM). PARATYPES one male, same data as holotype; two females, one male. Yap Group, Gagil District;
one male. Yap Group, Map I; two males, one female (on same point mount), Yap Group, Ruming I. All paratypes
labelled: “July-Au 50, R. J. Goss” (BPBM).

Description. — Length 5. 1-7.8 mm. Tufts of minor setae present on Segments V-X; median lobe of head short,
broad, its tip anterior to eye; medial margin of temporal lobe curved; orbital groove very broad, pollinose, one to three
temporal setae present in orbital groove.

Pronotum elongate, oval; length/greatest width 1.45; widest at basal 0.33; sides oblique anteriorly, evenly narrowed to
apex, distinctly narrowed to base; hind angles very obtuse; base strongly oblique, forming obtuse angle at midline; pronotal
setae absent; median groove relatively broad, closed anteriorly, open posteriorly; pollinose; paramedian groove very broad,
sloped gradually to inner carina medially; outer carina curved, narrow, of even width; marginal groove not visible in dorsal
view, not impressed, but represented by complete strip of pollinosity.

Elytron without distinct striae except for traces of sutural stria; each elytron with three narrow raised glabrous
carinae, separated by broad pollinose areas (evidently representing dilated striae); elytron with many setae, about five
medial to inner carinae; 10-12 between inner and second carinae; 12-14 between second and third carinae, and 15-20
between third carina and margin (Fig. 19); metasternum without median sulcus; Sterna III-V each with pair of broad
pollinose transverse grooves; lateral pit in Sternum IV, slight in male, greatly enlarged in female; male with front and hind
trochanters pointed; male anterior femur without ventral tooth; calcars of male pointed at apex, with dorsal margin sinuate
or notched.

The narrow glabrous carinae on an otherwise pollinose elytron separate this species from all
other members of the subgenus. The elytral carinae suggest Rhyzodiastes s. str. of South
America, but the gradual lateral slope of the inner carina contrasts with the sharply defined
lateral margin of the latter subgenus.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Distribution. — Caroline Islands. Recorded from Yap, Palau, and Ulithi. Detailed records
are in Bell and Bell, 1981.

THE SULCICOLLIS GROUP

This group is characterized by the reduction of the marginal stria of the elytron, which is
represented by a row of punctures or is entirely absent. The tufts of minor setae occur on
antennal Segments V-X. The group is known from the Moluccas and the Caroline Islands. R.
sulcicollis and R. maritimus appear closely related. Shared characters include: the outer carina
of the pronotum is broad; the median groove is narrow and is abbreviated both anteriorly and
posteriorly; the sutural stria is not impressed; the parasutural stria lacks setae. R. raffrayi is
much more distantly related; as the outer carina of the pronotum is narrow and curved, the
median groove is broader and is not abbreviated; the sutural stria is impressed for 0.67 of its
length, and the parasutural stria has setae. All these characters are probably plesiomorphic,
and R. raffrayi may be little modified from the common ancestor of the group. The two
remaining species may have evolved from beetles similar to R. raffrayi which rafted to the
Caroline Islands in the Equatorial Countercurrent. R. maritimus appears to have derived from
populations from the central Carolines, which subsequently became R. sulcicollis , rather than
representing a separate invasion from the Moluccas. In R. pollinosus, the form of the pronotum
strongly suggests that of R. raffrayi , suggesting that the former species might be a highly
modified offshoot of the latter species, and representing an independent invasion of the western
Carolines.

Rhyzodiastes ( Temoana ) raffrayi Grouvelle 1895a NEW COMBINATION

(Fig. 12)

Rhyzodiastes raffrayi Grouvelle 1895a: 158.

Clinidium raffrayi (Grouvelle) Grouvelle 1903.

Rhyzodiastes raffrayi (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “MOLUQUES: Gilolo, Raffray & Maindron, 78” (MNHN).
This island is now known as Halmahera.

Description. — Length 5.3 mm. Antennal stylet short, pointed; tufts of minor setae present on Segments V-X;
basal setae of antennal segments absent; Segment I with distinct apical pollinose band; Segments II-X without pollinosity;
head longer than wide; median lobe pointed, its tip opposite interior third of eye; postclypeal groove connected to frontal
groove; medial margin of temporal lobe rather evenly curved; temporal lobes well separated from one another, coming
closest together opposite posterior margin of eye; orbital groove entirely absent; one temporal seta present.

Pronotum only moderately elongate; length/greatest width 1.45; sides strongly curved, widest near middle; narrowed
at base, apex; apex truncate; base rounded; hind angles very obtuse; median groove deep, moderately narrow, anterior
median pit more dilated than posterior median pit; median groove deeply impressed to base; paramedian groove rather
broad, its base dilated into large basal impression; basal impression closed posteriorly by raised glabrous carina which is
continuous with outer carina; latter forms narrow, raised margin of uniform width, less than 0.2 of distance from
paramedian groove to midline at middle of pronotal length; pronotal setae absent; marginal groove shallow, complete,
visible only in lateral view.

Elytra elongate, slightly narrowed anteriorly; sutural stria fine, shallow, punctate; its apical third effaced; parasutural
stria deep, complete, impunctate, its base bent medially to medial angle of scarp; intratubercular stria deep, impunctate,
complete; marginal stria not impressed, represented by row of punctures, effaced in basal and apical thirds; apical portion
of marginal stria impressed below apical tubercle; subapical and apical tubercles elevated; latter contacting one another at
midline; parasutural stria with three setae in apical half; intratubercular stria with one or two setae near apex; apex of
marginal stria with several setae; hind trochanter of male pointed; hind calcar of male bluntly pointed.

The above description is incomplete because the holotype is damaged, with the front legs
missing. The female is unknown. The form of the pronotum is distinctive in this species, as is
the combination of a reduced marginal stria with an impressed sutural stria which is 0.66

Quaest. Ent.. 1985,21 (1)

Bell and Bell

Plate 2. Figs. 1 1-21. Genus Rhyzodiastes, new Subgenus Temoana. Figs. 1 1-18. Head and pronotum, dorsal aspect; Fig.
11, /?. (T.) pollinosus Bell and Bell; Fig. 12, R. (T.) raffrayi (Grouvelle); Fig. 13, R. (T.) sulcicollis (Grouvelle); Fig. 14,
R. ( T .) maritimus Bell and Bell; Fig. 15, R. (T.) guineensis (Grouvelle); Fig. 16, R. (T.) singularis (Heller); Fig. 17, R.

( T .) bipunctatus new species; Fig. 18, R. (T.) rimoganensis (Miwa); Fig. 19, Left elytron, dorsal aspect, R. (T.i pollinosus
Bell and Bell. Fig. 20, Hind tibia, male R. (T.) guineensis (Grouvelle). Fig. 21, Sternum VI, R. (T.) bipunctatus new
species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Eni., 1985, 21 (1)

Bell and Bell

Plate 3. Figs. 22-33. Genus Rhyzodiastes, new Subgenus Temoana. Figs. 22-29, Head and pronotum, dorsal aspect; Fig.
22, R. (T.) mirabilis (Lea); Fig. 23, R. (T.) indigens new species; Fig. 24, R. (T.) convergens new species; Fig. 25, R. (T.)
waterhousei (Grouvelle); Fig. 26, R. (T.) mishmicus (Arrow); Fig. 27, R. (T.) myopicus (Arrow); Fig. 28, R. (T.) vadiceps
new species; Fig. 29, R. (T.) preorbitalis new species; Fig. 30, Sternum VI, R. (T.) convergens new species; Figs. 31-33,
Sterna IV-VI, right half; Fig. 31, R. (T.) preorbitalis new species, female; Fig. 32, R. (T.) myopicus (Arrow), female; Fig.
33, R. (T.) vadiceps new species, male.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent., 1985,21 (1)

Bell and Bell

Plate 4. Figs. 34-45. Genus Rhyzodiastes, new Subgenus Temoana. Figs. 34-39, Head and pronotum, dorsal aspect; Fig.
34, R. (T.) frater (Grouvelle); Fig. 35, R. (T.) patruus new species; Fig. 36, R. (T.) bifossulatus (Grouvelle); Fig. 37, R.
(T.) denticauda new species; Fig. 38, R. (T.) gestroi (Grouvelle); Fig. 39, R. (T.) propinquus new species; Figs. 40-41,

Hind tibia, male; Fig. 40, R. (T.) frater (Grouvelle); Fig. 41, R. (T.) patruus new species; Figs. 42-43, Left elytron, apex,
dorsal aspect; Fig. 42, R. (T.) frater (Grouvelle); Fig. 43, R. (T.) patruus new species; Figs. 44-45, Elytra, posterior aspect;
Fig. 44, R. (T.) bifossulatus (Grouvelle); Fig. 45, R. (T.) denticauda new species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent.. 1985, 21 (1)

Bell and Bell

Plate 5. Figs. 46-50, 52-54. Genus Rhyzodiastes, new Subgenus Temoana. Figs. 46-51, Head and pronotum, dorsal
aspect; Fig. 46, R. (T.) bonsae new species; Fig. 47, R. (T.) fairmairei (Grouvelle); Fig. 48, R. (T.) spissicornis
(Fairmaire); Fig. 49, R. (T.) alveus new species; Fig. 50, R. (T.) fossatus new species; Figs. 52-53, Prothorax, left lateral
aspect; Fig. 52, R. (T.) alveus new species; Fig. 53, R. (T.) fossatus new species; Fig. 54, Left elytron, dorsal aspect, R. (T.)
fossatus new species. Figs. 51, 55-58. Genus Rhyzodiastes , new Subgenus Rhyzostrix. Fig. 51, R. (R.) davidsoni new
species; Figs. 55-58, Sterna III— VI, right half, female; Fig. 55, R. (R.) davidsoni new species; Fig. 56, R. (R.) nitidus new
species; Fig. 57, R. (R.) menieri new species; Fig. 58, R. (R.) maderiensis (Chevrolat).

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent., 1985, 21 (1)

Bell and Bell

complete.

Rhyzodiastes ( Temoana ) sulcicollis (Grouvelle 1903) NEW COMBINATION

(Fig. 13)

Clinidium sulcicolle Grouvelle 1903: 137-138.

Rhyzodiastes sulcicollis (Grouvelle) Bell and Bell 1981.

Type Material. — According to Grouvelle (1903), in the Oberthiir collection. We did not find it in our visits to
the MNHN, but it may be in recently discovered Oberthiir material which we have not yet studied. Grouvelle gave the
locality as “Isles Carolines: Hogolu” an obsolete name for Truk.

Description (abridged from Bell and Bell 1981). — Length 4.0-6.5 mm. Tufts of minor setae on
Segments V-X; basal setae present on Segments V-X; Segment I with apical pollinose band; Segment II with trace of one;
remaining segments without pollinosity; head slightly longer than wide; median lobe short, its tip acute, entirely anterior to
eyes; median lobe narrowly connected laterally to antennal lobe, separating frontal groove from post-clypeal groove; latter
forming isolated oval impression; temporal lobes rather narrowly separated at middle, forming obtuse median angles just
posterior to posterior margin of eye; orbital groove entirely absent; small pollinose preorbital pit present; temporal setae
absent; mentum with four prelabial and two labial setae.

Pronotum elongate, narrow, length/greatest width 1.65; widest near middle; lateral margin feebly curved, narrowed at
apex, base; apex truncate, base rounded; hind angles very obtuse; pronotal setae absent; median groove very narrow, linear,
its margins finely pollinose; groove closed at both ends, both median pits distinctly wider than groove, both removed from
ends of groove, groove represented by shallow impressions anterior to anterior median pit and posterior to posterior median
pit; posterior median pit equidistant from middle of pronotum and pronotal base (shallow median depression posterior to it
looking like a second pit); paramedian groove narrow, posterior end with small, deep, punctiform basal impression;
pollinosity of paramedian groove restricted to very narrow strip along lateral margin; inner carina with well-defined lateral
margin, nearly straight, wider than paramedian groove; outer carina 0.66 as broad as inner carina at middle, curved,
slightly tapered anteriorly; marginal groove entire, finely pilose, shallow except at posterior end.

Elytron slightly narrowed anteriorly; sutural stria not impressed, represented by row of very coarse punctures;
parasutural and intratubercular striae deeply impressed; lateral margin of each higher than medial margin, suggesting a
carina; parasutural and intratubercular striae uniting posteriorly; marginal striae not impressed except near apex;
represented by row of very coarse punctures in middle of elytron, entirely effaced anteriorly and posteriorly; apex of
marginal stria with four to six setae; elytron otherwise without setae; abdominal Sterna III-V each with pair of pollinose
transverse sulci which are narrowly separated in midline, each with conspicuous puncture at medial end; a similar pair of
pits but no transverse sulci on Sternum II; lateral ends of sulci of Sterna IV and V forming enlarged pits in both sexes,
larger in female than in male; Sternum VI with submarginal groove and two pairs of anteriolateral pits; male without
ventral tooth on anterior femur; trochanters of both sexes rounded; calcars small, pointed. The male genitalia of this
species have been illustrated by Bell and Bell (1978).

The coarse punctures of the sutural stria, which is not impressed, separate this species from
all other members of the subgenus.

Bell and Bell (1981) discussed a form from the Islands of Param, Tol, and Dublon, which
might be a separate species. In this form, the female is 4.0-5. 0 mm. long, and lacks a tubercle of
the sixth abdominal sternum. Females of the nominate form are 5.0-6. 5 mm. long, and have a
tubercle on the sixth sternum. This larger form coexists with the small one on the three islands
named above, and is found in many additional islands. There is a male specimen, from Tol,
which might belong to the dwarf form. It is only 4.0 mm. long, and has a pollinose spot near the
center of the sixth sternum. All other males are larger, and have at least a trace of a tubercle at
the middle of the sixth sternum. More collections are necessary to establish that the small male
is correctly associated with the small female, and that the small form is a distinct species.

Distribution. — Central Carolines, on the high islands of Truk (Dublon, Moen, Tol, Param),
also on the low island of Pis, in the barrier reef of Truk, and on the atolls of Satawal, Nama,
and Woleai (Utagal Island). Bell and Bell (1981) consider it likely that the records from
barrier islands and atolls result from accidental introduction by man. Bell and Bell (1981) give
detailed locality records.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Rhyzodiastes ( Temoana ) maritimus Bell and Bell 1981 NEW COMBINATION

(Fig. 14)

Rhyzodiastes maritimus Bell and Bell 1981: 66-67.

Type Material. — HOLOTYPE female, labelled: KUSIAE, Mutunlik, 22 m. 1-31-53, J.F.G. Clarke (BPBM).
PARATYPES one female, same locality and collector as holotype; one female, KUSIAE: Mt. Matanta, 180 m., 11-12-53,
J.F.G. Clarke, “decaying Hibiscus tiliaceus ” (BPBM).

Description (abridged from Bell and Bell 1981). — Length 4. 3-6. 2 mm. Tufts of minor setae present on
Segments V-X; basal setae present on Antennal Segments V-X; Segment I with apical pollinose band which is interrupted
ventrally; Segment II with trace of pollinosity dorsally; pollinosity otherwise absent from antenna; head distinctly longer
than wide; median lobe rather long, ending in acute point opposite middle of eye; median lobe connected laterally to
antennal lobe, separating postclypeal groove from frontal groove; postclypeal groove forming oval depression; temporal
lobes converging posteriorly, closest together posterior to eyes, where they form rounded medial angles; orbital groove Fine,
linear, complete, extending to occiput; temporal seta absent; eye very narrow, crescentic, smaller than in related species;
mentum with four prelabial and two postlabial setae.

Pronotum elongate, oval; length/greatest width 1.48, widest near middle, lateral margins distinctly curved, base
slightly wider than apex; apex truncate, base rounded; hind angles very obtuse; pronotal setae absent; median groove very
Fine, linear, abbreviated both anteriorly and posteriorly, ending anteriorly at anterior median pit which is separated from
anterior margin by more than its own length; median groove ending posteriorly at posterior median pit, which is closer to
middle of pronotum than to base of pronotum; paramedian groove linear, curved, pollinose, ending posteriorly at basal
impression, which is closed posteriorly; inner carina broad, flat, scarcely cariniform; outer carina about 0.5 as broad as
inner one, of equal width throughout, curved; marginal groove entirely absent.

Elytron slightly narrowed anteriorly; sutural stria entirely absent; parasutural and intratubercular striae complete,
impressed. Finely punctate, pollinose; lateral margin of each stria much higher than medial margin, suggesting a carina;
parasutural and intratubercular striae uniting posteriorly; marginal stria entirely absent except for a short impressed part
near apex, which contains four to six setae; elytral setae otherwise absent; abdominal Sterna III-V each with transverse
pollinose band which is narrowly interrupted at midline; Sternum II with pair of pollinose spots; Sternum VI with pair of
transverse pollinose bands anteriorly, and an entire submarginal pollinose band posteriorly; female with lateral pit in
Sternum IV (male unknown); midline of abdomen slightly carinate; femora entirely devoid of pollinosity and setae;
trochanters and coxae glabrous.

This species is similar to R. sulcicollis, but is easily separated by the presence of the orbital
groove, and the absence of the marginal groove of the pronotum and of the sutural and
marginal striae.

Distribution. — Known only from Kusiae, in the eastern Caroline Islands.

THE SINGULARIS GROUP

In this group, the tufts commence on Antennal Segment V, the orbital groove and marginal
striae are complete, and temporal setae are present. Six species are known, from Australia and
the Solomon Islands, west to Celebes, Formosa, and Sumatra. In all species in which the male
is known, the anterior and posterior trochanters are pointed in the male.

Phytogeny. — R. guineensis of New Guinea and R. singularis of Celebes appear to form a
line apart from the other species. Among the characters shared by the two species are an
obsolete median groove, an obsolete sutural stria and a last visible abdominal sternum with a
narrow sub-marginal groove which is well separated from the transverse grooves. The male of
R. guineensis differs from other known males in the form of the middle and hind tibiae, and in
the presence of a tooth on the ventral margin of the anterior femur. The male of R. singularis is
unknown, so it is uncertain whether or not these characters occurred in the common ancestor of
it and R. guineensis.

In the remaining five species, the median prothoracic groove is deeply impressed, as is the
sutural stria. In the male, the middle and hind tibiae are not thickened above the calcars, and
the anterior femur of the male is not toothed ventrally but is tuberculate in most species. The
submarginal groove of Sternum VI is expanded and joined to the transverse groove, nearly

Quaest. Ent.. 1985, 21 (1)

Bell and Bell

completely enclosing a diamond-shaped central glabrous area.

R. indigens of Sumatra is a puzzling species. On one hand, it resembles R. bonsae of the
gestroi group, also from Sumatra. It might be a relative of the latter species which has
secondarily lost the tuft of minor setae on Segment IV. On the other hand, it is close to R.
mirabilis of Australia (singulars group) except for the shape of the pronotum.

The four remaining species can be grouped in two pairs: R. bipunctatus of Guadalcanal and
R. rimoganensis of Taiwan have many setae in the parasutural stria, and well developed basal
setae on some of the outer antennal segments, and have at least some of the transverse sulci of
the abdomen continuous across the midline; R. mirabilis of Australia and R. convergens of
New Britain, have the parasutural stria with at most one seta, and lack basal setae on the
antennal segments, while the transverse grooves are broadly separated at the midline.

Rhyzodiastes ( Temoana ) guineensis (Grouvelle 1903) NEW COMBINATION

(Figs. 15, 20)

Clinidium guineense Grouvelle 1903: 138-139.

Rhyzodiastes guineensis (Grouvelle) Bell and Bell 1978

Type Material. — LECTOTYPE (here designated) male, labelled: “Nuova Guinea, Fly River, L. M. D’Albertis,
1876-77” (GEN). PARALECTOTYPES 1 1 males, 1 1 females, same data as lectotype (GEN). In the original description,
mention is also made of a specimen collected at Sattelberg by Biro, located in the Budapest Museum. We have not studied
this specimen and cannot testify it is identical to the series from the Fly River. A male specimen in the BMNH is labelled
“co-type. New Guinea 1901.267, N.J. Gella or Golla”. This specimen is not listed in the original description and is
probably incorrectly labelled as a co-type.

Description. — Length 6. 3-8.0 mm. Antennal stylet small, acute, tufts of minor setae present on Segments V-X;
Antennal Segments IX and X with basal setae; Segment I pollinose dorsally; Segment II with broken pollinose ring;
Segment III with traces of pollinosity; head almost twice as long as wide; median lobe rhomboid, its apex acute, opposite
middle of eye; median lobe separated from antennal lobe; latter shining, glabrous; temporal lobe 2.5 longer than wide;
frontal space unusually long, wide; medial margins nearly straight; temporal lobe with rather broad fringe of pilosity on
posterior margin and posterior 0.5 of medial margin; temporal lobes actually somewhat convergent posteriorly, but this is
inconspicuous because it is concealed by the fringe; frontal grooves deep; postantennal grooves deep, entire, narrow; orbital
groove deep, narrow, nearly straight, margin with postorbital pilosity shortly behind eye; temporal lobes separated by more
than width of one of them; temporal lobe flat, shining, impunctate; eye narrow, crescentic, about twice as long as wide,
0.66 of length of temporal lobe; one temporal seta located in orbital groove just posterior to eye; genae glabrous.

Pronotum moderately long, length/greatest width 1.46; widest slightly behind middle, sides curved, base narrowed,
apex more strongly narrowed; median groove very Fine, linear between the pits; anterior median pit enlarged, tear-drop
shaped, apical; posterior median pit at basal 0.25 of length, median groove posterior to it widened, shallow; median groove
entirely glabrous; inner carinae together forming convex, glabrous surface, sloping laterally into paramedian groove; latter
distinct, its floor glabrous, its lateral boundary (medial scarp of outer carina) pollinose; basal impression small, oblique,
closed posteriorly by flat glabrous elevation; outer carina narrowed anteriorly, broadened posteriorly, its base rounded,
apex very narrowly truncate; pollinosity of its medial margin attaining hind angle, curving into marginal groove; marginal
groove complete, linear, not visible in dorsal view; pronotal setae absent, sub-marginal groove absent; propleuron
iridescent; anterior part of notopleural suture pollinose; sternopleural groove absent; pleural groove represented by reduced
pit.

Elytra rather narrow, scutellar pits very large, surrounded by pollinosity which meets at midline anterior to pits,
forming triangle, tapering posteriorly to point at base of remnant of sutural stria, connected laterally to transverse strip
which reaches base of parasutural stria; sutural stria reduced to short, medially-directed scarp in basal 0.2 of length; first
and second intervals thus scarcely distinct; in some specimens a line of Fine punctures represents more posterior part of
sutural stria in middle 0.33 of elytron; parasutural stria straight, complete, forming pollinose, medially-directed scarp;
Interval III nearly flat, facing dorsolaterally; basal pollinosity of elytron broadly interrupted opposite base of Interval III;
apex of Interval III forming subapical tubercle which is not at all swollen; intratubercular stria linear, pollinose, impressed,
its base disappearing into humeral pollinose area of elytron; apex of intratubercular stria merging with broad pollinose
band across anterior walls of apical tubercles; marginal stria impressed, linear, pollinose, complete; apical tubercle
impunctate, slightly swollen; elytron entirely without setae; metasternum not sulcate; transverse sulci of abdominal sterna
narrowly separated at midline; female with lateral pits in Sternum IV; tibial spurs of middle and hind legs slightly unequal;
male with trochanters I, III pointed; anterior femur of male with ventral tooth; calcars with distinct shoulder tooth,
bounded above by rounded emargination (Fig. 20); tibia above emargination greatly thickened.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

The greatly reduced sutural stria separates this species from all others excepting R.
singulars. From the latter species it differs in having the subapical tubercle narrow, in lacking
elytral setae, and in having the median groove of the pronotum impressed, though narrow.
Distribution. — New Guinea. In addition to the type material, we have seen one female

Specimen, labelled: “NEW GUINEA, Orio, 145’, Purari River, Oct. 7, 1967” (MCZ) and one female, labelled:
“Humboldt B., N. Guinea, Doherty” (MNHN). This locality was later called Hollandia.

Rhyzodiastes ( Temoana ) singularis (Heller 1898) NEW COMBINATION

(Fig. 16)

Clinidium singulare Heller 1898: 3.

Rhyzodiastes singularis (Heller) Bell and Bell 1978.

Type Material. — HOLOTYPE female, labelled: “S. CELEBES, Lompa-Battau, 3000’, Marz, 1896, H.
Fruhstorfer, ex museo W. Rothschild, 1899” (MNHN).

Description. — Length 7.0 mm. Antennal stylet small, acute; antennal segments entirely without basal setae; all
antennal segments with pollinose rings (these broken on Segments IX, X), tufts of minor setae on V-X; head only slightly
longer than wide, median lobe hastate, rather short, its apex acute, opposite middle of eye; antennal lobe entirely pollinose;
temporal lobe 1.5 longer than wide; frontal space broad, its anterior 0.5 glabrous except for linear median strip; frontal
grooves deep but glabrous; medial margins of temporal lobes broadly curved, closest together opposite middle of eyes;
temporal lobe with fringe of pilosity on posterior margin; frontal grooves deep, glabrous; orbital groove complete though
very narrow opposite middle of eye; temporal lobe convex, shining, impunctate; eye small, crescentic, about 0.5 length of
temporal lobe, its length three times its width; one temporal seta present, in orbital groove midway between hind margin of
eye and occipital angle; genae glabrous.

Pronotum moderately long, length/greatest width 1.44; widest behind middle, sides curved, base and apex both only
slightly narrowed; median groove obsolete, scarcely visible; anterior median pit large, tear-drop shaped, apical; posterior
median pit large, occupying basal 0.15 of length, constricted at 0.5 of its length, open posteriorly; inner carinae fused to
form broad convex glabrous surface, sloping laterally into paramedian groove; bases of inner carinae form lobes on either
side of posterior median pit; margins of lobes fringed with pollinosity; basal impressions oblique, punctiform, pollinose,
bounded posteriorly by flat glabrous ridges; paramedian groove bounded laterally by steep, pollinose, slightly undulated
scarp; outer carina only slightly narrowed anteriorly, medial margin nearly straight, lateral one feebly curved; pollinosity
of its medial margin attaining hind angle, curved into marginal groove; marginal groove complete, linear, scarcely visible
in dorsal view; submarginal groove absent; pronotal setae absent; pollinose pit present at anterior end of notopleural suture;
sternopleural groove absent; pleural groove oblique, narrow.

Elytron moderately narrow; pilose area occupying lateral part of base of sutural interval, nearly concealing scutellar
pits; sutural stria almost absent, in basal 0.2 represented by medially-directed scarp, from there to middle of length barely
traceable as a shallow impression; in posterior 0.5 of elytron entirely invisible; First and second intervals thus not distinct;
parasutural stria straight, complete, forming pollinose, medially-directed scarp; basal transverse pollinosity of elytron
entire, not interrupted opposite base of Interval III, Interval III convex, its apex forms strongly swollen subapical tubercle,
medial margin of latter (apex of parasutural stria) abruptly sinuate; intratubercular stria impressed, linear, entire, its apex
merging with broad pollinose band across anterior wall of arpical tubercle; Interval IV continuous with apical tubercle,
latter not swollen; marginal stria complete, linear, impressed, pollinose; parasutural stria with at least three or four setae
near base perhaps with complete row of very small setae (only basal punctures visible in holotype, but it appears that setae
may have broken off); intratubercular stria with one big basal seta; apical tubercle with several small setae; marginal stria
with several setae in apical portion; metasternum not sulcate; transverse sulci of abdominal Sterna III-VI widely separated
medially; female with rather small lateral pit in Sternum IV; Sternum VI with basal transverse sulci broadly separated
from submarginal groove; middle of Sternum VI evenly convex.

This species is separated from all other except R. guineensis by the great reduction of the
sutural stria. It differs from the latter in having well-developed elytral setae, in having the
medial margin of the subapical tubercle strongly swollen, and in having the median groove of
the pronotum almost absent.

Quaest. Ent., 1985,21 (1)

Bell and Bell

Rhyzodiastes ( Temoana ) bipunctatus new species
(Figs. 17,21)

Type Material. — HOLOTYPE male, labelled: “SOLOMON ISLANDS: Gaudalcanal, Mt. Austen, 18/4.1963,
P. Greenslade, 5401, B.M. 1966-477” (BMNH). PARATYPES one male, same data as holotype except dated 19-9-1962
(BMNH); one female, labelled: “SOLOMON ISLANDS, Gaudalcanal, Ngalim Mtn., 8/8,1963, P. Greenslade, 8383”
(BMNH). There is another male, missing head and pronotum, that is mounted on the same pin as this specimen but it was
not made a paratype.

Description. — Length 5.0-7.9 mm. Antennal stylet short, conical; tufts of minor setae on Segments V-X; antennal
Segments VI-X with basal setae; antennal Segments I-X with pollinose rings (those of IX, X more or less broken between
the setae); head longer than wide; median lobe short, triangular, its apex pointed, opposite anterior margin of eye; antennal
lobe entirely pollinose; frontal space broad, parallel-sided pollinose; frontal grooves broad, deep, pollinose; temporal lobe
2.5 times longer than wide, entirely broadly fringed with pilosity; orbital groove broad, deep, continuous; two temporal
setae present, one located in orbital groove near posterolateral angle of temporal lobe, the other arising from conspicuous
puncture in glabrous part of temporal lobe well posterior to eye; eye narrow, crescentic, less than 0.5 length of temporal
lobe; posterior 0.5 of gena pilose.

Pronotum moderately elongate, length/greatest width 1.50, widest at middle, sides distinctly curved; both ends
narrowed, the apex more than the base; median groove deep, middle 0.33 sublinear; anterior median pit apical, dilated;
posterior median pit appearing double, anterior portion at basal 0.33 of pronotum, this separated by constriction from
posterior part which is almost as deep, and which reaches base of pronotum; median groove and pits pollinose; inner carina
convex, glabrous, rather broad, its lateral margin more distinct from paramedian groove than in most species of Temoana;
inner carina and paramedian groove without evident microsculpture; paramedian groove bounded laterally by broad
pollinose strip along nearly vertical medial scarp on outer carina; basal impression oblique, pollinose, about 0.20 of length
of pronotum; outer carina broad, its medial margin only slightly curved, its lateral margin more strongly so, therefore outer
carina is widest at middle and tapered at both ends; marginal groove marked by complete strip of pollinosity, but little
impressed, scarcely visible in dorsal view, but well-marked in lateral view; submarginal groove absent; pronotal setae
absent; notopleural suture with pollinosity in anterior 0.5; sternopleural groove absent; pleural groove impressed, pollinose,
its ventral 0.5 linear, its dorsal 0.5 expanded into a pit.

Elytra rather narrow, without a caudal lobe; scutellar pits present, but inconspicuous, lying within transverse band of
pilosity which extends entirely across base of elytra; sutural interval flat; sutural stria with basal 0.25 glabrous, middle
portion with pollinose strip, apex recurved for short distance at apical 0.25 of elytron; its apex not joining parasutural stria;
Interval II nearly flat, sloped laterally; parasutural stria impunctate, broad, pollinose, its lateral wall a medially-directed
scarp; pollinosity of parasutural stria continuing posteriorly, where it merges with that of intratubercular stria, and
combined strip continues to midline along anterior slope of apical tubercle; Interval III convex, its apex forming short
subapical tubercle which is scarcely dilated; tips of subapical tubercles separated by 3.5 times width of one of them;
intratubercular stria impressed, pollinose, dilated; Interval IV broad, nearly flat, continuous with apical tubercle; latter
moderately swollen; marginal stria complete, its base expanded, middle part (from basal 0.16 to middle) very fine, linear;
apical part deeper, curving below apical tubercle; marginal stria entirely pollinose; submarginal stria impressed, ending
opposite Sternum V of abdomen; sutural stria without setae (but apical impression without setae or with one or two setae
in line with sutural stria); parasutural stria with complete series of seven to nine setae; intratubercular with two to four
setae in apical 0.5; marginal stria with six to eight setae in apical 0.5; apical tubercle with three or four setae.

Metasternum with median sulcus; abdominal Sterna III-V each with dilated pollinose transverse sulci; that on
Sternum III continuous; that of IV either continuous or very narrowly interrupted at midline; that of V distinctly but
rather narrowly interrupted; female with large, deep lateral pit in Sternum IV: Sternum VI with short triangular
transverse sulci broadly joined to greatly dilated marginal groove, partly isolating glabrous discal area (Fig. 21); tibial
spurs of middle and hind legs decidedly unequal; male with anterior and posterior trochanters pointed; anterior femur of
male with ventral side tuberculate; both calcars are distinctly angulate proximally, separating basal transverse margin
from oblique anterior margin, latter sloped to acute point; hind calcar larger than middle one.

This species is unique within the subgenus in having a setiferous puncture on the disc of the
temporal lobe, in addition to the usual one in the orbital groove. Otherwise it is closest in form
to R. rimoganensis (Miwa), but differs from the latter in having the median groove of the
pronotum less dilated and in having the antennal lobe entirely pollinose.

Rhyzodiastes ( Temoana ) rimoganensis (Miwa 1934) NEW COMBINATION

(Fig. 18)

Clynidium ( Rhyzodiastes ) rimoganense Miwa 1934: 256-257 (note misspelling of generic name).

Rhyzodiastes rimoganensis (Miwa) Bell and Bell 1978.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Type Material. — HOLOTYPE, female. According to Miwa, from TAIWAN: Taihoku Province, Rimogan; coll.
K. Obayashi, March 25, 1933. We have not studied the holotype, but have studied several specimens from Taiwan which
agree closely with the original description and figure.

Description. — Length 5. 5-6. 9 mm. Antennal stylet short, conical, acute; tufts of minor setae on Segments V-X;
basal setae present, though sparse on Segments VIII-X; Segment I pollinose dorsally; antennae otherwise without
pollinosity; head slightly longer than wide; median lobe rather short, triangular, its apex acute, opposite middle of eye;
antennal lobe glabrous, shining, well separated from median lobe; frontal space rather narrow anteriorly, becoming broad
posteriorly; frontal grooves rather narrow, pollinose; temporal lobe 1 .5 longer than wide; medial margins strongly curved,
so that at middle, temporal lobes are separated by less than 0.5 of width of one of them; temporal lobe fringed with pilosity;
orbital groove narrow but complete, angulate opposite posterior margin of eye; one temporal seta present, in orbital groove,
posterior to eye; eye narrow, crescentic, about 0.75 of length of temporal lobe; genae glabrous, posterior face of temporal
lobe pilose.

Pronotum moderately elongate, length/greatest width 1.57, widest at middle, sides distinctly, evenly curved; both base
and apex distinctly narrowed; median groove moderately dilated, as broad as median pits; anterior median pit wider than
posterior median pit, so margins of median groove slightly convergent posteriorly; posterior median groove displaced
anteriorly, its anterior end at 0.20 of pronotal length anterior to pronotal base; median groove deep posterior to posterior
median pit, but narrowed, its sides glabrous; median groove in and between median pits glabrous medially, but with lateral
scarps pollinose; inner carina convex, its lateral margin sloping gradually into paramedian groove; paramedian groove
bounded laterally by pollinose strip on vertical medial scarp of outer carina; medial margin of outer carina curved, slightly
undulating; basal impression very small, narrowly closed posteriorly, connected to margin by strip of pollinosity; outer
carina moderately narrow, tapered anteriorly; marginal groove fine, impressed, linear, complete; visible in dorsal view as is
margin laterad to it; submarginal groove nearly complete, 0.95 of length of pronotum, becoming finely pollinose
posteriorly; pronotal setae absent; notopleural suture not pollinose; sternopleural groove absent; pleural groove oblique;
ventral surface of prothorax opalescent.

Elytra moderately elongate; elytral cauda absent; elytron with pilosity around scutellar pits, extending laterally to base
of Interval II, but broadly interrupted in Interval III, and represented laterally only by small pilose area at base of
marginal stria; sutural interval nearly flat; sutural stria impressed, faintly punctate, pollinose, finer than other striae, its
apex slightly out-curved at apical 0.15 of elytron (in some specimens joining parasutural stria); Interval II nearly flat,
sloped laterally; parasutural stria deep, its lateral wall a medially directed scarp, its extreme base curved medially,
posterior its pollinosity combining with that of intratubercular stria, and the combined strip continuing across anterior face
of apical tubercle to suture; Interval III nearly flat, its apex becoming slightly convex, forming subapical tubercle; latter
scarcely dilated; subapical tubercles separated by 3.5 times width of one of them; intratubercular stria impressed,
pollinose, dilated; Interval IV flat, continuous with apical tubercle; latter moderately swollen; marginal stria entire,
impunctate, rather deep; submarginal stria impressed, ending opposite base of Sternum VI; sutural stria without setae;
parasutural with two to seven setae; intratubercular stria with one seta near apex; marginal stria with several setae near
apex; stria with two or three setae near apex; apical tubercle without setae.

Ventral surface of pterothorax and abdomen opalescent; metasternum not sulcate; male with flattened,
microsculptured median area on abdominal Sterna I, II; abdominal Sterna III-V with pollinose transverse sulci, these
entire on III, IV, narrowly interrupted at midline in V; female with lateral pits on Sternum IV; Sternum VI with triangular
transverse sulci broadly joined to greatly dilated marginal groove, nearly isolating rhomboid glabrous area, latter in both
sexes with pair of tubercles; tibial spurs of middle, hind legs unequal; male with front, hind trochanters pointed; anterior
femur of male tuberculate ventrally; middle calcar small, pointed, acute; hind calcar small, obtuse. (Description of male
characters taken from specimens from Nakanoshima.)

This species differs from R. bipunctatus in lacking a setiferous puncture in the middle of the
temporal lobe and in having distinct submarginal groove of the pronotum. It differs from R.
mirabilis and R. convergens in having basal setae on the antennae and more than one seta in
the parasutural stria. The tubercles on Sternum VI are a unique but inconspicuous character of
this species.

Distribution. — We have studied a female specimen from Taiwan: Puli (Hori), July 1954
“native collector” (BPBM). We tentatively assign to this species a series of 15 specimens from
RYUKYU ISLANDS: Nakanoshima, Is. Tokara, 5 July, 1960, M. Sato leg. These do not
appear to differ in form from the Taiwan specimen, except that the latter has seven setae in the
parasutural stria, while the Nakanoshima specimens have two or three (or, unilaterally, one).
However, we have not seen males from Taiwan, so it is possible that the populations from the
two islands are not conspecific.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Rhyzodiastes ( Temoana ) mirabilis (Lea 1904) NEW COMBINATION

(Fig. 22)

Rhysodes mirabilis Lea 1904: 80-81.

Rhyzodiastes mirabilis (Lea) Bell and Bell 1978.

Type Material. — According to the original description, from Cairns, Queensland, Australia. We have not
studied it, but have studied an enlarged photograph of it, kindly sent by Barry Moore. It is a female.

Description. — Length 6.5-7.0 mm. Antennal stylet short, slightly flattened; tufts of minor setae present on
Segments V-X; basal setae entirely absent from antenna; Segments I-X each with apical pollinose band; head as wide as
long; median lobe triangular, pointed posteriorly, its apex opposite middle of eye; antennal lobe largely pollinose, but with
small isolated frontal boss; frontal space rather narrow, its smallest diameter about 0.33 of width of temporal lobe; frontal
grooves narrow, entirely glabrous; temporal lobe 1.8 longer than wide, medial margins strongly curved so that at middle
temporal lobes are separated by about 0.33 times width of one of them; temporal lobe fringed with pilosity; orbital groove
narrow, complete, sinuate posterior to eye; temporal seta present, on medial margin of orbital groove posterior to eye; eye
narrowly crescentic, about 0.67 as long as temporal lobe; genae glabrous, posterior face of temporal lobe pilose.

Pronotum moderately elongate, length/greatest width 1.56; widest at middle, sides distinctly, evenly curved; base
distinctly narrowed; apex less narrowed than base; median groove deep, narrowed between median pits, which are small;
posterior median pit at basal 0.33 of length; groove posterior to posterior median pit as deep as remainder of groove,
deepened at pronotal base to form a secondary posterior median pit; inner carinae together convex, sloping laterally into
paramedian groove; paramedian groove bounded laterally by vertical pollinose scarp on medial margin of outer carina;
medial margin of outer carina evenly curved; basal impression very small, triangular, its posterior margin pollinose (so
impression appears open posteriorly); outer carina broad, 0.5 as wide as inner one bounded posteriorly by pollinosity which
reaches hind angle and connects to marginal groove; marginal groove visible in lateral but not in dorsal view, rather broad,
0.33 as wide as outer carina, shallow, pollinose; submarginal groove absent; pronotal setae absent; sternopleural groove
absent; pleural groove oblique; notopleural suture pollinose; prosternum with anterior margin narrowly pollinose, narrowly
interrupted at midline.

Elytra moderately broad, without caudal lobe; each elytron with prominent parascutellar pit at base of sutural stria;
these pits situated relatively far from one another, separated by glabrous area; elytron with complete transverse strip of
pollinosity at base, not interrupted opposite Interval III; Interval I broad, flat; sutural stria straight, impressed, pollinose,
its apex slightly recurved; Interval II nearly flat, sloped laterally; parasutural stria impressed, straight, becoming slightly
broader posteriorly, pollinose, its apex joined to intratubercular stria, and combined pollinosity continued posteriorly along
anterior face of apical tubercle to suture; Interval III nearly flat, its apex convex, suddenly dilated, forming preapical
tubercle; preapical tubercles separated by 3.5 times width of one of them; intratubercular stria impressed, pollinose;
Interval IV flat, continuous with apical tubercle; latter scarcely dilated; marginal stria impressed, narrow, slightly dilated
posteriorly, submarginal stria reaching base of Sternum V; sutural, parasutural and intratubercular striae without setae;
preapical tubercle with five or six setae; marginal stria with five to seven setae near apex.

Ventral surface not opalescent; metasternum not sulcate; abdominal sterna with pollinose transverse sulci which are
interrupted at midline (broadly so except for Sternum III of female which is very narrowly interrupted); both sexes with
rather small lateral pits on Sternum IV; Sternum VI with dilated submarginal groove which connects anteriorly to
transverse sulci, nearly isolating rhomboid glabrous area; latter not tuberculate; spurs of middle and hind tibiae equal;
male with anterior, posterior trochanters pointed; anterior femur of male tuberculate ventrally; middle and hind calcars
with slight “shoulder” angle on dorsal margin, calcars triangular, their apices obtuse.

The rounded temporal lobes of this species suggest R. rimoganensis , but it differs from the
latter species in lacking basal setae on the antennae, and lacks a submarginal groove on the
pronotum. It also has the elytral setae much more restricted, and has a small, isolated glabrous
spot on each antennal lobe. The shape of the pronotum, wider at apex than base, is distinctive
and separates it from R. indigens.

Distribution. — Restricted to Queensland. We have seen specimens from the following

localities: one male, one female, Shipton’s Flat (south of Cooktown), June, 1958, coll. Darlington (MCZ); one male.
Upper Little Mulgrave, 3-VIII-69, coll. James Tobler (CAS); one female, labelled; “Queensl. Myoberg” (LUN); one
male, one female, labelled “N. Queensland, Redlynch, 1 2-20- VIII- 1 938, R. G. Wind” (BMNH). The female of this pair is
in all respects R. mirabilis except for a deep median pit in Sternum VI. Whether or not this is an anomoly will depend on
the study of more specimens.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Rhyzodiastes ( Temoana ) indigens new species
(Fig. 23)

Type Material. — HOLOTYPE male, labelled: “SUMATRA, Si Rambe, XII-90-III-91, E. Modigliani’, (GEN).
PARATYPE one male, two females, same label as holotype (GEN). The female is labelled as a syntype of R. gestroi, but
is not conspecific with it, and does not have convergent temporal lobes, as specified in the original description of the latter
species.

Description. — Length 8. 0-9.0 mm. Antennal stylet short, conical; tufts of minor setae on Segments V-X;
Segments I-X each with subapical pollinose ring; basal setae of Segments IX, X one or two or absent; head slightly longer
than wide; median lobe triangular, tip pointed, opposite anterior end of eye; frontal grooves rather wide, deep, glabrous;
temporal lobe more than two times longer than wide; median margins curved, posteriorly oblique, slightly divergent;
temporal lobe fringed posteriorly and on posterior 0.5 of medial margin with pilosity; orbital groove complete; one
temporal seta in orbital groove posterior to eye; eye narrowly crescentic, about 0.67 of length of temporal lobe; genae
glabrous, posterior face of temporal lobe pollinose.

Pronotum short, length/greatest width 1.37, widest near middle, sides curved; base moderately narrowed, apex very
strongly narrowed, median groove deep, anterior 0.5 as wide as anterior median pit, constricted posterior to middle, then
broadened to posterior median pit; groove posterior to posterior median pit as deep as at middle; inner carinae together
convex, sloped laterally to paramedian groove; paramedian groove bounded laterally by vertical pollinose scarp on medial
margin of outer carina; latter evenly curved; basal impression small, triangular, open posteriorly; outer carina broad, 0.5 as
wide as inner one at middle, strongly narrowed anteriorly, extreme apex pollinose, marginal groove visible in dorsal view;
submarginal groove absent; pronotal setae absent; sternopleural groove absent; pleural groove oblique, notopleural suture
glabrous.

Elytron moderately broad, without caudal lobe; elytron with basal pollinosity interrupted at Interval III; Interval I
broad, slightly convex, sutural stria straight, impressed, pollinose, apex slightly recurved; Interval II nearly flat, sloped
laterally; parasutural stria impressed, straight, pollinose; apex joined to intratubercular stria; Interval III, raised above
level of Interval II, nearly flat; preapical tubercle inflated; preapical tubercles separated by 1.5 width of one of them;
intratubercular stria impressed, pollinose; Interval IV flat, continuous with apical tubercle; latter inflated; marginal stria
impressed, not dilated posteriorly; submarginal stria reaching base of Sternum V; parasutural stria with one or two setae
near apex; intratubercular stria with one seta near apex; marginal stria with four setae near apex.

Ventral surface not opalescent; metasternum not sulcate; abdominal sterna with transverse sulci broadly interrupted in
midline in both sexes; each sulcus with prominent medial, lateral pit; female with large lateral pit on Sternum IV; Sternum
VI with submarginal sulcus widely separated from transverse sulci; middle, hind tibiae with spurs nearly equal; male with
anterior, posterior trochanters pointed; ventral surface of anterior femur of male with many small tubercles; tibiae thick;
middle calcar triangular, small, acute; hind calcar larger, acute, dorsal margin convexly curved.

Among members of the singularis group, this species comes closest to R. mirabilis. It differs
from the latter in having a much shorter pronotum which is strongly narrowed anteriorly. In
appearance it comes close to R. bonsae in the gestroi group, but the latter species has the outer
carina of the pronotum shallowly concave and the tufts of minor setae on the antenna beginning
on Segment IV.

Rhyzodiastes ( Temoana ) convergens new species
(Figs. 24, 30)

Type Material. — HOLOTYPE male, labelled: “New Britain, Gisiluve, Nakanai Mts., 1050 m., July 26, 1956,
coll. E. J. Ford, Jr.” (BPBM). PARATYPES two males, two females, same data as holotype (BPBM); two males, one
female, same data as holotype but dated July 25, 1956 (BPBM).

Description. — Length 6. 2-7. 2 mm. Antennal stylet short, acuminate; tufts of minor setae present on Segments
V-X; basal setae of antennae entirely absent; head distinctly longer than wide; median lobe short, broad, at widest point
0.33 of width of head, its apex opposite anterior end of eye; parafrontal boss rather large, narrowly separated from
antennal rim; frontal grooves rather broad, deep, pollinose; temporal lobe about two times longer than broad, medial
margins shallowly emarginate, margins divergent posterior to median lobe; then convergent, shallowly sinuate to occipital
angles; latter separated by about 0.20 of width of head; medial margin of temporal lobe with fringe of very fine pollinosity;
posterior margin with fringe of pilosity; orbital groove not quite complete, ending posteriorly at temporal seta, not quite
attaining basal pilosity; orbital groove barely sinuate posterior to eye; one temporal seta present, on orbital groove near to
posterior margin of temporal lobe; eye narrowly crescentic in lateral view, 0.67 as long as temporal lobe; genae glabrous;
posterior face of temporal lobe pilose.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Pronotum elongate, length/greatest width about 1.67; widest at middle, sides distinctly curved; both base and apex
narrowed; median groove deep, narrow, parallel-sided, scarcely enlarged opposite median pits; posterior median pit at 0.16
of length; groove posterior to posterior median pit as deep as remainder of groove, but not forming distinct secondary
posterio-median pit; inner carinae together convex, sloping laterally into paramedian grooves; paramedian grooves broad,
bounded laterally by vertical pollinose scarp on medial margin of outer carina; medial margin of outer carina evenly
curved; basal impression oblique, sloped up gradually to flat, very finely pollinose ridge which closes it posteriorly; outer
carina broad, very convex, narrowed to both base and apex; pollinosity of median scarp connected to marginal groove at
hind angle; marginal groove visible in lateral but not in dorsal view; marginal groove linear; submarginal groove absent;
pleural groove oblique, impressed, rather narrow; notopleural suture pollinose; prosternum without pollinosity on anterior
margin.

Elytra moderately broad, without caudal lobe; elytra with small, widely separated parascutellar pits at bases of sutural
striae, with very little associated pilosity; basal pilosity of elytron narrowly interrupted opposite medial 0.5 of Interval III;
Interval I broad, flat; sutural stria impressed, very finely pollinose, its apex curved laterally, to meet parasutural stria;
Interval II convex; parasutural stria impressed, straight, becoming scarp-like posteriorly; Interval III nearly flat anteriorly,
becoming convex posteriorly, forming scarp on lateral face near apex; subapical tubercle swollen, its median margin
oblique, intratubercular stria impressed, linear anteriorly, becoming slightly dilated posteriorly, pollinose; Interval IV flat,
connected to apical tubercle, latter impunctate, somewhat swollen; a pore ventrad to apical tubercles on suture; marginal
stria fine, impressed, complete, pollinose; submarginal stria ends at apex of Sternum V; parasutural stria with one seta
posterior to middle; intratubercular stria with two setae in apical 0.2; marginal stria with three or four setae near apex.

Ventral surface not opalescent; metasternum with fine, incomplete medial sulcus limited to posterior 0.5; abdominal
sterna with pollinose transverse sulci which are broadly interrupted medially in both sexes; both sexes with lateral pits on
Sternum IV; Sternum VI with marginal groove connected anteriorly to transverse sulci, nearly isolating rhomboid glabrous
area (Fig. 30); latter not tuberculate; spurs of middle, hind tibiae equal; male with anterior, posterior trochanters pointed;
anterior femur of male neither dentate nor tuberculate ventrally; middle and hind calcars “shouldered” on dorsal side.

This species resembles R. mirabilis , but is more elongate and narrow, with the pronotum
more oval, and more narrowed at the apex. The medial sinuation on the temporal lobe is
characteristic, but is very small in some specimens.

THE MISHMICUS GROUP

This contains the only species from west of Wallace’s Line which have the tufts of minor
hairs beginning on Antennal Segment V, except for R. indigens. The species resemble those of
the singulars group except in having the orbital groove strongly abbreviated or absent and in
lacking temporal setae. All species lack pronotal and elytral setae. The anterior pits are
enlarged, rounded and conspicuous. The transverse sulci of the abdominal sterna are reduced.
The group ranges from Thailand to extreme eastern India. R. water housei and R. preorbit alis
appear more closely related to one another than to R. mishmicus. The latter species has a
distinct though reduced orbital groove and a long median lobe, while the two former species
have a preorbital pit rather than an orbital groove, and have a very short median lobe. On the
other hand, R. preorbitalis lacks the distinct median metasternal sulcus found in the other two
species. This suggests that the sulcus was acquired by the common ancestor of the group, and
then secondarily lost in R. preorbitalis.

Rhyzodiastes ( Temoana ) mishmicus (Arrow 1942) NEW COMBINATION

(Fig. 26)

Clinidium mishmicum Arrow 1942: 182-183.

Rhyzodiastes mishmicus (Arrow) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: ASSAM: “Mishmi Hills, Delai Valley, Chauliang, i-xii, 1936,
alt. 4840 ft.; (Miss) M. Steele BM 1937-324” (BMNH).

Description. Length 7.0 mm. Antennal stylet prominent, acuminate; basal setae of antennae restricted to lateral
surfaces, sparse, but present on Segments V-X; head as broad as long; median lobe triangular, long, its apex opposite
posterior 0.5 of eye; frontal space very narrow, scarcely wider than one frontal groove; frontal grooves long, broad, deep,
pollinose; length of temporal lobe 1.5 greater than its width; medial margin of temporal lobe evenly, convexly curved; inner

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

and posterior margins of temporal lobe broadly fringed with pilosity; orbital groove present, shallow, extending posteriorly
to end even with middle of eye; eye short, crescentic, about 0.33 as long as temporal lobe.

Pronotum short, broad, length/greatest width 1.25; widest near middle, sides curved; apex strongly narrowed, base
moderately so; median groove fine, linear; both median pits displaced towards middle of pronotum; anterior median pit in
oval depression; median groove distinct but shallow posterior to posterior median groove; inner carina convex, glabrous, its
lateral margin sloped gradually into paramedian groove; lateral margin of paramedian groove bounded by very narrow,
inconspicuous strip of pollinosity on medial scarp of outer carina; outer carina broad, its width at middle about 0.67 of
width of inner carina at same level; outer carina strongly tapered anteriorly, moderately so posteriorly; basal impressions
deep but narrow, about 0.33 as wide as posterior part of inner carina; marginal groove narrow, distinct, visible in dorsal
view; submarginal groove absent.

Elytra short, rather broad, their sides parallel; sutural interval pollinose at base; sutural stria fine, its apical fifth
obsolete; parasutural stria complete, its base bent medially to reach base of sutural stria; intratubercular stria deeper than
the others, complete; subapical tubercle somewhat elevated, its apex rounded; marginal stria fine, complete, impressed;
apical tubercles small, contiguous, marginal stria with three setae below apical tubercle; elytral setae otherwise absent.

Metasternum with complete, deep, dilated median sulcus; male with triangular lateral pits on abdominal Sterna III-V,
that of IV deeper than the other; pits not extended medially to form transverse sulci; Sternum VI with complete marginal
groove; female unknown; middle and hind tibiae each with spurs equal; male with large, distally-directed ventral tooth on
anterior femur and deep lateral groove on anterior femur; all trochanters rounded distally; calcars small; calcar of middle
leg with apex level with bases of spurs; that of hind leg with apex raised well above level of spurs.

The presence of a short orbital groove, the short, broad form of the body, the fine median
groove of the pronotum, and the elongate median lobe of the head easily distinguish this species
from the other members of the group. R. myopicus, in the myopicus group, is superficially
similar in appearance, but has the tufts of minor hairs beginning on Segment IV of the
antennae, the orbital groove entirely absent, tibial spurs strongly unequal, and the abdominal
sterna with prominent transverse sulci.

Rhyzodiastes ( Temoana ) waterhousei (Grouvelle 1910) NEW COMBINATION

(Fig. 25)

Clinidium ( Rhyzodiastes ) waterhousei Grouvelle 1910: 326-327.

Rhyzodiastes waterhousei (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE female, labelled: “BIRMAH: Ruby Mines (coll. Doherty) 64626 Fry Coll.
1905.100” (BMNH)

Description. — Length 6.5 mm. Antennal stylet prominent, conical; basal setae numerous on Segments VII-X;
head as broad as long; median lobe triangular, short, its apex opposite anterior end of eye; anterior tentorial pits very large,
separated by less than the width of one of them; frontal space parallel-sided, elongate, its width about 0.33 of width of one
temporal lobe; frontal grooves narrow, shallow, temporal lobe 1.5 longer than wide; medial margin of temporal lobe almost
straight, forming obtuse angle with posterior margin; posterior margin of temporal lobe broadly fringed with pollinosity,
medial margin glabrous; orbital groove absent; preorbital pit present; eye narrowly crescentic, rather elongate, about 0.67
of length of temporal lobe.

Pronotum moderately long, length/greatest width about 1.36; widest near middle; base distinctly narrowed; apex
strongly so; median groove narrow, sublinear, deeper than in R. mishmicus\ both median pits displaced towards middle of
pronotum; anterior median pit in oval depression; median groove distinct but shallow posterior to posterior median pit;
inner carina convex, glabrous, its lateral margin sloped gradually into paramedian groove; basal impressions broad, deep,
distinctly wider than posterior part of inner carina; the base of latter consequently distinctly narrower than in R.
mishmicus ; lateral margin of paramedian groove bounded by narrow strip of pollinosity on medial margin of outer carina;
outer carina broad, its width near middle about 0.67 of width of inner carina at same level; outer carina distinctly
narrowed anteriorly, and posteriorly; marginal groove narrow, complete, visible in dorsal view; a shallow submarginal
groove present, visible only in lateral view.

Elytra rather short, the sides nearly parallel; sutural interval pollinose at base; sutural stria rather fine, complete, its
apex joining parasutural; sutural stria obsoletely punctate; Interval II distinctly convex; parasutural stria impunctate, more
deeply impressed than sutural stria, its base bent medially to reach base of sutural stria; Interval III distinctly convex;
intratubercular stria slightly less impressed than parasutural; subapical tubercle somewhat elevated, its apex rounded;
marginal stria impressed, fine, complete; apical tubercles small, contiguous; marginal stria with three or four setae below
apical tubercle; elytral setae otherwise absent.

Metasternum with complete, deep, linear median sulcus; abdominal sterna with short transverse sulci which are
dilated laterally and which have a small pit at medial end; transverse sulci separated medially by approximately 0.33 of
width of sternum; female with deep lateral pit on Sternum IV; middle and hind tibiae each with spurs equal; anterior

Quaest. Ent., 1985, 21 (1)

Bell and Bell

femur of female not angulate; male unknown.

This species is most similar to R. preorbitalis , but the latter species lacks the median sulcus
on the metasternum, has the anterior tentorial pits less enlarged, the transverse sulci of the
abdomen more poorly developed, and the anterior femur of the female is strongly angulate
ventrally. Rhyzodiastes vadiceps, in the myopicus group, is superficially similar to the two
preceding species, but has the tufts of minor setae beginning on Antennal Segment IV, and the
occiput largely glabrous and distinctly notched in lateral view.

Rhyzodiastes ( Temoana ) preorbitalis new species
(Figs. 29,31)

Type Material. — HOLOTYPE female, labelled: “THAILAND: E. slope Doi Sutep, 875-950 m., 15-VII-1962,
coll. E. S. Ross, D. O. Cavagnero” (CAS). PARATYPE one female, labelled: “Doi Sutep, Siam, Feb. 10, 1928, Coll. Alice
Mackie” (NMNH).

Description. — Length 6.8-7. 8 mm. Antennal stylet small, acute; basal setae numerous on Segments VII-X; head
as broad as long; median lobe short, triangular, its apex entirely anterior to eye; anterior tentorial pits moderately large,
separated by 1.5 times the width of one of them; frontal space parallel-sided, elongate, its width about 0.33 of width of one
temporal lobe; frontal grooves narrow, shallow, glabrous, temporal lobe 1.5 times longer than wide; medial margin of
temporal lobe almost straight, forming angle with posterior margin; posterior margin of temporal lobe rather narrowly
fringed with pilosity; medial margin glabrous; orbital groove absent; preorbital pit present; eye narrow, crescentic, less
than 0.5 of length of temporal lobe.

Pronotum rather elongate, length/greatest width about 1.42; widest near middle, sides curved; base moderately
narrowed; apex very strongly narrowed; median groove fine, linear, both median pits displaced towards middle of
pronotum; anterior median pit in long, oval depression; median groove represented by fine, shallow line posterior to
posterior median pit; inner carina convex, glabrous, sloped gradually to paramedian groove; basal impressions broad, deep,
distinctly wider than posterior part of inner carina; the base of latter narrowed as in R. waterhousei ; lateral margin of
paramedian groove bounded by narrow strip of pollinosity on medial margin of outer carina; outer carina rather broad, its
width at middle about 0.50 of width of inner carina at same level; outer carina distinctly narrowed anteriorly and
posteriorly; marginal groove fine, complete, visible in dorsal view; submarginal groove short, oblique, restricted to basal
0.20 of pronotum, visible only in lateral view.

Elytra rather short, their sides nearly parallel; sutural interval pollinose at base; sutural stria rather fine, complete,
impunctate; Interval II slightly convex; parasutural stria impunctate, more deeply impressed than sutural stria, its apex
bent medially to reach base of sutural stria; Interval III distinctly convex; intratubercular stria complete, slightly less
impressed than parasutural; subapical tubercle somewhat elevated, its apex rounded; marginal stria fine, scarcely
impressed at middle, becoming finely punctate and more distinctly impressed posteriorly, where it passes below apical
tubercle; apical tubercles small, contiguous; apex of marginal stria with three or four setae below apical tubercle; elytral
setae otherwise absent.

Metasternum with elongate median impression near posterior margin, but without median sulcus; abdominal sterna
with transverse sulci scarcely developed, each appearing as short medial extension from triangular lateral pit in Sterna III,
IV, V; female with lateral pits of IV deeper than the others (Fig. 31); middle and hind tibiae each with two equal spurs;
anterior femur of female strongly angulate ventrally; male unknown.

This species is closest to R. waterhousei , but differs sharply in lacking a median sulcus on
the metasternum, in having the anterior tentorial pits less enlarged, and in having the anterior
femur of the female strongly angulate ventrally.

THE MYOPICUS GROUP

The members of this group have the tufts of minor setae beginning on Segment IV of the
antenna, and the median groove of the pronotum linear. Some species, particularly R.
myopicus, resemble the mishimcus group in having a short, broad head, with the orbital groove
absent or very reduced. In others the head is elongate, and the orbital groove more distinct. All
species differ from the mishmicus group in having strongly developed transverse sulci on the
abdominal sterna, and in having a tuft of minor setae on antennal Segment IV. The gestroi and
fairmairei groups resemble the myopicus in the latter respect, but differ in having the median

denticauda

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Phylogenetic Diagram 1 . Reconstructed Phylogeny of species of Rhyzodiastes subgenus Temoana, Myopicus group.

Quaest. Ent., 1985,21 (1)

myopicus

Bell and Bell

groove of the pronotum dilated. Many of them have numerous elytral setae, while the myopicus
group lack such setae, except, in some species, near the apex of the marginal stria.

There are six species, two from the Malay Peninsula, three from Borneo, and one of
unknown range, but probably also from Borneo.

Phylogeny. — A possible phylogeny for the group is illustrated in Diagram 1. Species 1, the
hypothetical ancestor for the group, probably had the following characters: antennal stylet long;
Segment XI elongate, nearly cylindrical; basal setae present; antennal segments not thickened;
orbital groove, one temporal seta present; base of parasutural stria straight; preapical tubercle
not elevated; apical tubercles thickened, contiguous, elytral humeri not narrowed; setae present
in apex of marginal stria; tibiae slender; at least anterior femur of male with many tubercles on
ventral surface; middle, hind tibial spurs equal; male trochanters rounded at apex.

R. myopicus appears to be the sister species to Species 2, the hypothetical ancestor of the
five remaining species. Apomorphic features of R. myopicus include loss of the orbital groove,
the short, very flat head with strongly reduced eyes, and the strongly unequal tibial spurs.
Probable plesiomorphic features include base of parasutural stria straight; antennae and tibiae
slender; one temporal seta retained; setae in apex of marginal stria retained; elytral humeri not
narrowed; antennal Segment XI cylindrical; trochanters of male rounded. The broad anterior
truncation of the pronotum and outer carina are of uncertain significance.

Species 2 probably had the following apomorphic features: base of parasutural stria bent
sharply medially at base; humeral region of elytra strongly narrowed; antennae, tibiae strongly
thickened; antennal Segment XI compressed, short; at least hind trochanter of male pointed.
Like R. myopicus it probably retained a temporal seta and several subapical setae in the
marginal striae, although these have been lost in some of its descendants. Unlike R. myopicus it
had the pronotum narrowed anteriorly, with the outer carina not truncate at the apex.

We postulate three species descended from Species 2. Species 3, Species 4, and R. vadiceps.
Species 3 showed the following specializations: outer antennal segments with complete pollinose
rings; preapical tubercle elevated, tooth-like; apical tubercle bounded anteriorly by deep
transverse notch; basal setae of antennae entirely absent. Unspecialized features retained from
Species 2 included equal tibial spurs; one temporal seta; one seta in apex of marginal stria;
marginal groove present on pronotum, though shallow; male with trochanters 1 and 3 pointed;
and femora 1 and 3 tuberculate ventrally.

Species 4 showed striking specializations in the apex of the middle and hind tibiae, only one
tibial spur is present, while there is a curved apicolateral process. Other apomorphic characters
included loss of setae of the marginal stria. Possible plesiomorphic characters include retention
of basal setae on the antennae and of the marginal groove of the pronotum.

R. vadiceps shows the complete loss of the marginal groove of the pronotum as a
specialization. Possibly the elongate antennal stylet is also an apomorphic feature, as are loss of
marginal and temporal setae. Most of its other characters are plesiomorphic, for instance, the
retention of two equal tibial spurs. The antennal segments are less thickened than in the
descendants of Species 3 and 4. Of uncertain significance are the secondary sexual characters
of the male. This species and R. patruus are the only species in which all three pairs of femora
are tuberculate ventrally. If all were tuberculate in the male of Species 2, then the absence on
some legs in the remaining species can be interpreted as a secondary loss. Likewise, the loss of
the point on the anterior trochanter in R. vadiceps is probably secondary, as both anterior and
posterior trochanters are pointed in the remaining descendants of Species 2, with the possible
exception of R. denticauda, of which the male has not been collected.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Species 3 gave rise to R. bifossulatus and R. denticauda. R. bifossulatus is specialized by
the great elevation of the preapical tubercles, which are close together. Otherwise, it is probably
much like Species 3. It is the only member of the group besides R. vadiceps to retain setae in
the marginal stria, and shares with the latter species and R. patruus the retention of the
temporal seta. R. denticauda shows a unique specialization on the separation of the prominent,
subtruncate apical tubercles. It has also lost both temporal and elytral setae. The preapical
tubercles are less specialized than are those of R. bifossulatus , being further apart and
somewhat less elevated. Unfortunately the secondary sexual characters of the male are
unknown.

Species 4 gave rise to R. frater and R. patruus. R. frater has lost the temporal seta and the
ventral tubercles on the femora. A plesiomorphic feature is the retention of the basal setae of
the antenna. R. patruus has lost the basal setae. The tubercles and the projecting angle on the
hind tibia of the male are clearly apomorphic features, while the retention of ventral tubercles
on all femora of the male and the retention of a temporal seta are clearly plesiomorphic.

Rhyzodiastes ( Temoana ) myopicus (Arrow 1942) NEW COMBINATION

(Figs. 27, 32)

Clinidium myopicum Arrow 1942: 182.

Rhyzodiastes myopicus (Arrow) Bell and Bell 1978.

Type Material. — LECTOTYPE male, labelled: “FEDERATED MALAY STATES: Pahang, Cameron’s
Highlands, Bukit-Lendong, 5000 ft., May 21, 1931, coll. H. M. Pendlebury” (BMNH). PARATYPE According to Arrow
there was a second specimen, sex unstated, collected with the type. We have been unable to locate it. If it still exists, it is a
paralectotype.

Description. — Length 6. 3-7. 5 mm. Antennal stylet short, conical; antennal Segment XI longer than broad; sparse
basal setae on Segments IX, X; antennal pollinosity restricted to Segment I; head as broad as long, strongly flattened;
antennal lobe pollinose; median lobe somewhat cross-shaped with lateral lobe posterior to each tentorial pit; tip of median
lobe acute, posterior to middle of eye; frontal grooves very narrow, shallow, postantennal pit scarcely evident; medial
margin of temporal lobe curved; temporal lobe less than 1.5 longer than wide posterior margin; posterior 0.5 of medial
margin narrowly fringed with pilosity; orbital groove entirely absent, eye narrow, short, less than 0.33 of length of
temporal lobe; one temporal seta, near posteriormost point on temporal lobe; postorbit, genal lobe pilose.

Pronotum moderately elongate, length/greatest width 1.40; basal margin strongly curved; lateral margins rather
weakly curved; base, apex moderately narrowed; apex truncate, with distinct anterior angles; median groove Fine, linear;
median pits narrow, not displaced from base, apex; inner carinae glabrous, together forming convex discal area, sloped
gradually into paramedian grooves; medial margin of outer carina with narrow pollinose strip; basal impression small,
oblique; outer carina broad, its width near middle about 0.5 of width on inner carina at same level; outer carina slightly
curved, of nearly even width, anterior end truncate; marginal groove impressed, complete, clearly visible in dorsal view;
submarginal groove represented by impression in basal 0.12 of pronotum; pronotal setae absent.

Elytra with sides parallel, base scarcely narrowed; sutural stria deep, entire, obsoletely punctate; Interval II less
depressed than Interval I; parasutural stria impressed, impunctate, base straight; Interval III slightly convex, apex
(subapical tubercle) not swollen; intratubercular stria complete,impressed, impunctate; marginal stria fine, complete,
impressed; apical tubercles slightly swollen, contiguous at suture, with minute pore ventral to them in midline; marginal
stria with three to six setae below apical tubercle; elytral setae otherwise absent.

Metasternum without median sulcus; abdominal Sterna III— V with prominent transverse sulci; in male. III, IV
continuous across midline, V narrowly interrupted at midline; in female all sulci narrowly interrupted medially (Fig. 32);
both sexes with enlarged lateral pit on Sternum IV; Sternum VI with transverse sulci at base, curved submarginal sulcus;
abdominal sulci narrow; middle, hind tibiae rather slender, with inner spur shorter than outer one; anterior femur of male
tuberculate ventrally, with small ventral tooth; all trochanters of male rounded distally; both pairs of calcars small, acutely
pointed.

The broad pronotum, broad outer carinae with anterior end truncate and the flat head and
small eyes are distinctive of this species. The most similar member of the myopicus group is R.
frater of Borneo. The latter species is much longer and more slender, has the outer carina of the
pronotum narrower, and lacks the temporal seta. R. mishmicus , of the mishmicus group, is also
rather similar in appearance but lacks a tuft of minor setae on antennal Segment IV, lacks the

Quaest. Ent., 1985, 21 (1)

Bell and Bell

transverse sulci on the abdominal sterna, and has the spurs of each middle and hind tibia equal.

R. patruus is the only other member of the species group from the Malay peninsula. It
differs from the present species in having the outer carina very narrow, the head narrow and
elongate, and in having only one tibial spur and an apicolateral process.

Range. — R. myopicus is known only from the Malay Peninsula. In addition to the

lectotype, we have seen the following specimens: one male, labelled: “Malaya, G. Batu, Brinchang, 6500',
VI- 19-62, coll. E. S. Ross & D. Cavagnaro” (CAS); two females, labelled: “Malaya, Pahang, Cameron Highlands, Mt.
Brinchang, coll. L. W. Quate” (BPBM). One is dated, 1-4-1959, 1980 m., the other 5-1-1959 at 1900 m. “in dead wood”.

Rhyzodiastes ( Temoana ) vadiceps new species
(Figs. 28, 33)

Type Material. — HOLOTYPE male, labelled: “Mjoberg Coll., W. W. Funge Bequest” (CAS). No locality is
given. Borneo is a likely if unproven provenance, Firstly because the closest relatives of the species, R. frater, R.
bifossulatus, and R. denticauda are all from Borneo, and secondly, because Mjoberg is known to have collected in Borneo.
The type specimens of Omoglymmius fraudulentus Bell and Bell and Rhyzodiastes denticauda , described herein, both
have labels identical to that on the type of R. vadiceps, in addition to labels for specific localities in Borneo. If not from
Borneo, this species might be from one of the Greater Sunda Islands or possibly from the Malay Peninsula.

Description. — Length 8.5 mm. Antennal stylet acute, longer than in other members of group; antennal Segment
XI slightly compressed; slightly longer than wide; outer antennal segments strongly thickened, oblate sphaeroidal; tufts of
minor setae present on Segments IV-X; basal setae entirely absent; antennal pollinosity restricted to Segments I, II; head
1.5 longer than wide antennal lobe glabrous; median lobe triangular, tip acute, opposite anterior 0.33 of eye; frontal
grooves shallow, glabrous; postantennal pit small; lateral margin of frontal groove sloped gradually to temporal lobe;
medial margin of temporal lobe long, oblique, sinuate near occipital angle; temporal lobe 2.5 longer than wide; margin
lined with short pollinosity near occipital angle; orbital groove represented by very minute pollinosity medial to eye,
invisible except under high magnification; small preorbital pit present; eye short, about 0.5 of length of temporal lobe,
broader than in R. myopicus ; temporal setae absent; genal lobe prominent, rectangular, nearly glabrous, separated from
temporal lobe by deep, pollinose notch.

Pronotum elongate, length/greatest width 1.60; basal margin transverse; lateral margins strongly curved; base, apex
strongly narrowed; anterior angles indistinct; median groove fine, linear; median pits large, oval, not displaced from base,
apex; inner carinae glabrous, together forming convex disc, sloped gradually to paramedian grooves; medial margin of
outer carina with only a trace of pollinosity; basal impressions small; outer carina tapered anteriorly, widest at middle;
marginal groove entirely absent; submarginal groove absent; pronotal setae absent.

Elytra with sides parallel near middle; base strongly narrowed; sutural stria deep, its apex effaced, obsoletely punctate;
parasutural stria complete, impressed, more so apically, faintly punctate, base bent medially nearly to base of sutural stria;
Interval III convex, its apex elevated, forming distinct preapical tubercle; intratubercular stria fine, complete; apical
tubercles swollen, contiguous at midline, meeting medially above slit-like pore; elytral setae entirely absent.

Metasternum without median sulcus; midline of abdomen elevated, slightly cariniform, separating dilated transverse
sulci; Sternum VI with transverse sulci, also with dilated submarginal sulcus; shallow lateral pit present on Sternum IV in
male (Fig. 33); tibiae moderately thick; spurs equal; male with ventral surface of all femora with many minute tubercles;
front, middle trochanters of male rounded at apex hind trochanter acutely pointed; middle, hind calcars large, acutely
pointed, proximal margin strongly angulate; female unknown.

This species is closest to R. frater , from which it differs most conspicuously in the absence of
the marginal grooves of the pronotum, in the very shallow frontal grooves with indistinct lateral
margins, and in the great reduction of the pollinosity of the postorbital region and of the genal
tubercles.

Rhyzodiastes ( Temoana ) frater (Grouvelle 1903) NEW COMBINATION

(Figs. 34, 40, 42)

Clinidium frater Grouvelle 1903: 135-136.

Rhyzodiastes frater (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Born. Occ., Riv. Sambay, pres Ngabang, J. B. Ledru 1887”
(MNHN). This locality is in the northwestern part of Indonesian Borneo.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Description. — Length 5.9-8.0 mm. Antennal stylet minute, conical; antennal Segment XI compressed, as wide as
long; outer antennal segments very short, thick, cylindrical, disc-like; tufts of minor setae present on Segments IV-X; basal
setae present, though sparse on Segments III-X; Segments I-III with subapical pollinose rings; pollinosity of outer
segments restricted to areas close to bases of setae; Segment I with prominent swellings around base of two setae of
anterior aspect.

Head 1.5 times longer than wide; antennal lobe glabrous; median lobe hastate, short, narrow, tip acute, just posterior
to anterior margin of eye; frontal grooves glabrous, moderately deep, margins distinct; tentorial, postantennal pits large;
medial margin of temporal lobe long, oblique; temporal lobe 3.0 longer than wide; posterior margin and posterior 0.33 of
medial margin fringed with long pilosity; orbital groove very fine, shallow, pollinose, in a few specimens interrupted
posterior to eye; eye narrow, crescentic, 0.67 as long as temporal lobe; temporal setae absent; genal lobe, lower surface of
temporal lobe long pilose, partly concealing notch between them.

Pronotum elongate; length/greatest width 1.54; widest behind middle; basal margin slightly oblique on either side of
midline, where obtusely angulate; lateral margins curved; apex strongly narrowed; median groove fine, linear; median pits
large, oval, not displaced from base, apex; inner carina glabrous, together forming convex discal area, sloped gradually to
paramedian groove; medial margin of outer carina with narrow strip of pollinosity; basal impressions small, oblique; outer
carina widest posterior to middle, tapered anteriorly; lateral part of outer carina shallowly concave; marginal groove fine,
linear, ventrad to concavity of outer carina; submarginal groove, pronotal setae absent.

Elytra elongate, lateral margins parallel near middle, obliquely narrowed to humerus; swollen apical tubercles form
large but ill-defined cauda; sutural stria nearly complete, impressed, minutely punctate, apex effaced; parasutural stria
complete, impressed, base bent medially nearly to base of sutural stria; Interval III convex, apex elevated, forming distinct
preapical tubercle; intratubercular stria fine, complete; marginal stria effaced near humerus, otherwise narrow, complete
to suture; apical tubercles swollen, contiguous at midline, meeting medially above round pore (Fig. 42); elytral setae
entirely absent.

Metasternum with fine trace of median sulcus in posterior 0.5; mid-line of abdomen elevated, cariniform, separating
dilated transverse sulci; transverse sutures of abdomen deeply impressed; Sternum VI in both sexes with transverse sulci,
also broad, deep subapical depression occupying 0.67 of sternum, bounded posteriolaterally by impressed marginal groove;
in both sexes, Sternum IV with rather small lateral pit; tibiae thick; middle and hind tibiae each with one spur, also with
acute, medially-curved apicolateral process (Fig. 40); ventral surface of femur in male smooth, not tuberculate; front, hind
trochanters of male acutely pointed; middle, hind calcars large, acutely pointed, proximal margin angulate.

This species is closest to R. patruus of the Malay Peninsula, which it resembles in having
only one spur and an apicolateral process on the middle and hind tibiae. R. frater differs from
the latter species in lacking a temporal seta, in having basal antennal setae, in having the
intratubercular stria impressed, and in lacking tubercles on the femora of the male.

It differs from the two remaining Bornean species, R. denticauda and R. bifossulatus in not
having the apical tubercle separated from the outer intervals by a notch, and in having the
preapical tubercle less elevated and less tooth-like.

Range. — Northwestern Borneo, including both the Indonesian Borneo and Sarawak. In

addition to the holotype we have studied a series of 13 males, three females labelled: “Mt. Matang,
W. Sarawak, G. E. Bryant, Bryant Colin., 1919-147” with various dates from XII- 1 9 1 3 to 11-1914 (BMNH).

Rhyzodiastes ( Temoana ) patruus new species
(Figs. 35,41,43)

Type Material. — HOLOTYPE male, labelled: “Malaisie Johor, Sedili Kechil, 15-VIII-72. T. Jaccoud” (GVA).

Description. — Length 5.3 mm. Antennal stylet minute, conical; Segment XI slightly compressed, as wide as long;
outer antennal segments very short, thick, cylindrical, disc-like; tufts of minor setae present on Segments IV-X; basal setae
entirely absent; Segments I-X with complete subapical pollinose rings; Segment I with carina on dorsal surface.

Head 1.5 longer than wide; antennal lobe glabrous; median lobe hastate, short, narrow, tip acute, just posterior to
anterior margin of eye; frontal grooves glabrous, moderately deep, margins distinct; tentorial, postantennal pits large;
medial margin of temporal lobe long, oblique; temporal lobe 3.0 longer than wide, posterior margin, posterior 0.33 of
medial margin fringed with pollinosity; orbital groove very fine, pollinose, complete; eye narrow, crescentic, 0.67 as long as
temporal lobe; lateral margin of temporal lobe posterior to eye more oblique than in R. frater. temporal seta present; genal
lobe, lower surface of temporal lobe long, pilose, partly concealing notch between them.

Pronotum elongate; length/greatest width 1.58; widest behind middle; basal margin slightly oblique on either side of
midline, but not angulate at midline; lateral margins curved; apex strongly narrowed; median groove fine, linear; median
pits large, oval, not displaced from base, apex; inner carinae glabrous, together forming convex discal area, sloped
gradually to paramedian groove; medial margin of outer carina with narrow strip of pollinosity; basal impression small;

Quaest. Ent., 1985, 21 (1)

Bell and Bell

outer carina widest posterior to middle, tapered anteriorly; lateral part of outer carina shallowly concave; marginal groove
fine, linear, ventrad to concavity of outer carina; submarginal groove of outer carina absent.

Elytra elongate; lateral margins parallel near middle, obliquely narrowed to humerus; apical tubercles less swollen
than in R. frater, sutural stria complete, impressed, minutely punctate; parasutural stria complete, impressed, base bent
medially nearly to base of sutural stria; Interval III convex, apex elevated, forming distinct preapical tubercle (Fig. 43);
intratubercular stria impressed, pollinose near apex, otherwise effaced, marked only by elevated medial border of Interval
IV; apical tubercles contiguous at midline, meeting above minute pore; setae of elytral striae entirely absent.

Metasternum without median sulcus; midline of abdomen less distinctly elevated than in R. frater, transverse sutures
of abdomen deeply impressed; Sternum VI in male with transverse sulci, narrow subapical depression; tibiae thickened;
middle, hind tibiae each with one spur, also with acute, medially curved apicolateral process; ventral surfaces of all femora
of male tuberculate, anterior femur extensively so, middle femur with a few tubercles, hind one more densely tuberculate;
trochanters of front, hind leg acutely pointed; calcars acutely pointed; medial surface of hind tibia concave between calcar
and basal angle, latter prominent, truncate; medial surface with two minute tubercles on either side of middle of length
(Fig. 41); female unknown.

This species is closest to R. frater which it resembles in the presence of one tibial spur and a
conspicuous apicolateral process on the middle and hind tibia. It differs from the latter in the
presence of a temporal seta, absence of basal antennal setae, unimpressed intratubercular stria,
and in having tuberculate ventral surfaces on the femora of the male.

Rhyzodiastes ( Temoana ) bifossulatus (Grouvelle 1903) NEW COMBINATION

(Figs. 36, 44)

Clinidium bifossulatum Grouvelle 1903: 139-140.

Rhyzodiastes bifossulatus (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Borneo, Oberthuer Coll.” (MNHN).

Description. — Length 6. 7-8.0 mm. Antennal stylet conical, short; antennal Segment XI compressed, as wide as
long; outer antennal segments very short, thick, cylindrical, disc-like; basal setae absent; Segments I-X each with subapical
pollinose ring; latter interrupted ventrally on Segments IV-X by tuft of minor setae; Segment I with swellings around bases
of two prominent setae on anterior aspect.

Head as wide as long; antennal lobe glabrous; median lobe very short, its tip acute, opposite anterior margin of eye;
frontal grooves glabrous, moderately deep; frontal space very wide; tentorial, postantennal pits very large; antennal groove
oblique, slightly dilated; temporal lobes 2.5 longer than wide; closest together opposite middle of eye, posterior 0.67
oblique, divergent; posterior margin of temporal lobe fringed with long pilosity; orbital groove complete, fine, pollinose,
reaching posterior margin of temporal lobe; eye narrow, crescentic, 0.67 as long as temporal lobe; one temporal seta, in
orbital groove posterior to eye; posterior margin of temporal lobe, postorbit long, pilose, concealing cleft between temporal,
genal lobes; latter smaller, or shortly pilose than in R. denticauda.

Pronotum elongate; length/greatest width 1.57; widest behind middle; basal margin curved; lateral margins curved;
apex strongly narrowed; base moderately so; median groove fine, linear; anterior median pit oval, elongate; posterior
median pit round, separated from base by 0.2 of length of pronotum; inner carinae glabrous, together forming convex
discal area, sloped gradually to paramedian groove; medial margin of outer carina with narrow strip of pollinosity; basal
impressions small, deep, oblique; outer carina narrow, curved, bounded laterally by shallow broad concavity; ventral
margin of latter with trace of minute pollinosity; posterior end of outer carina with tuft of pilosity; submarginal groove,
pronotal setae absent.

Elytra elongate, lateral margins parallel near middle, obliquely narrowed to humerus; humerus with small lobe; apical
lobes form prominent, narrow cauda; sutural stria impressed, faintly punctate, apex joining parasutural; parasutural stria
complete, impressed, base bent medially to base of sutural stria; Interval III narrow, cariniform for most of length, base
forming prominent, fringed medial angle; elevated preapical tubercle prominent; preapical tubercles closer together than in
related species, separated by 0.5 of combined width of sutural intervals (Fig. 44); in lateral view, preapical tubercle with
posterior margin emarginate; intratubercular stria shallow, broad; becoming obsolete ventrad to preapical tubercle;
marginal stria impressed, connected by deep, narrow, glabrous impression to preapical impression, isolating apical tubercle
from remainder of Interval III; portion of marginal stria beyond impression bent ventrally, passing across lower surface of
apical tubercle to suture; apical tubercles swollen, contiguous at midline, meeting medially above slit-like pore; apex of
marginal stria with one seta.

Metasternum with Fine median sulcus in posterior 0.5; mid-line of abdomen scarcely carinate; abdominal sterna with
deep, wide transverse sulci, scarcely interrupted in midline, interruption very narrow, pollinose; abdominal sutures deeply
impressed; Sternum VI with transverse sulci, also deep marginal groove, delimiting central tubercle; small lateral pit on
Sternum IV in both sexes; tibiae thick, coarsely punctate, each puncture with prominent seta; middle, hind tibiae each with
two spurs; those of hind tibia equal; those of middle tibia very unequal, posterior spur minute, anterior one large, curved
anteriorly; no apicolateral process; male with ventral surface of anterior, posterior femora with many small tubercles; male

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

with anterior, posterior trochanters pointed; middle calcar acute, hind calcar smaller than middle one, triangular; proximal
margins of calcars not angulate.

This species and R. denticauda are characterized by having the preapical tubercle elevated
and tooth-like, separated by a notch from the apical tubercle. In this species, the apical
tubercles are contiguous at the suture, while in R. denticauda , they are separated.

Range. — Borneo, possibly restricted to the northeastern part. We have seen the following

specimens with specific locality data, both from Sabah (The former British North Borneo): one
male, labelled: “Sandakan, B. N. Borneo, Baker” (NMNH); one female labelled: “British North Borneo, Tawau, Quoin
Hill, Cocoa Res. Sta., 30-VI-1962, Y. Hirashima, coll.” (BPBM).

Rhyzodiastes ( Temoana ) denticauda new species
(Figs. 37, 45)

Type Material. — HOLOTYPE female, labelled: “Mt. Murud, Borneo, Mjoberg Coll., W. W. Funge Bequest”
(CAS). This locality is in eastern Sarawak. PARATYPE one female, labelled: “SARAWAK: Claudetown, 25, vii, 1932,
primitive white sand forest; Oxford Univ. Exp., B. M. Hobby and A. W. Moore, B.M. -1933-254” (BMNH). This locality
is now called Marudi and is in eastern Sarawak, not far from Mt. Murud.

Description. — Length 6. 8-7.0 mm. Antennal stylet conical, short; antennal Segment XI compressed, as wide as
long; outer antennal segments very short, thick, cylindrical, disc-like; basal setae absent; Segments I-X each with subapical
pollinose ring; latter interrupted ventrally on Segments IV-X by tuft of minor setae; Segment I swollen near bases of two
prominent setae on anterior aspect.

Head 1.5 longer than wide; antennal lobe glabrous; median lobe short, hastate, tip acute, even with anterior margin of
eye; frontal grooves glabrous, moderately deep; tentorial, postantennal pits very large; antennal groove transverse, very
fine, its lateral 0.5 effaced; temporal lobe three times longer than wide, closest together opposite middle eye; posterior 0.67
oblique, divergent; posterior margin, posterior 0.33 of medial margin of temporal lobe fringed with pilosity; orbital groove
very fine, complete, pollinose, reaching posterior margin of temporal lobe; eye relatively short, less than 0.5 of length of
•temporal lobe, narrow, crescentic; lateral margin of temporal lobe oblique posterior to eye; temporal setae absent; postorbit
long, pilose, concealing notch between temporal, genal lobes; dorsal surface with conspicuous, reticulate microsculpture.

Pronotum very elongate; length/greatest width 1.67; widest behind middle; basal margin curved; lateral margins
curved, convergent anteriorly; apex strongly narrowed; base moderately narrowed; median groove fine, linear: median pits
oval, elongate; posterior median pit separated from base of pronotum by 0.2 of length of pronotum; inner carinae glabrous,
together forming convex discal area; sloped gradually to paramedian grooves; medial margin of outer carina with narrow
strip of pollinosity; basal impression small, transverse, oval; outer carina narrow, curved, bounded laterally by shallow
broad concavity; ventral margin of latter with trace of minute pollinosity; posterior end of outer carina with tuft of
pollinosity; submarginal groove, pronotal setae absent.

Elytra elongate, lateral margins parallel near middle, obliquely narrowed to humerus; latter with prominent lobe
bounded posteriorly by pilose notch; apical lobes form prominent, narrow cauda; sutural stria scarcely impressed, faintly
punctate; parasutural stria shallowly impressed, complete; base bent medially to base of sutural stria; Interval III narrow,
cariniform for most of its length, base forming prominent, fringed medial angle; apex of Interval III forming prominent,
elevated preapical tubercle; preapical tubercles separated by combined width of sutural intervals; in lateral view, preapical
tubercle with posterior margin emarginate; intratubercular stria shallow, broad, becoming obsolete ventrad to preapical
tubercle; marginal stria broad, shallowly impressed, connected by deep, semicircular notch to preapical impression,
isolating apical tubercle from remainder of Interval IV; portion of marginal stria posterior to notch bent ventrally, passing
across lower surface of apical tubercle; apical tubercles swollen, subtruncate medially, separated by about 0.5 of distance
between preapical tubercles (Fig. 45); elytron entirely without setae.

Metasternum with fine median sulcus in posterior 0.5; midline of abdomen elevated, cariniform separating broad, deep
transverse sulci; abdominal sutures deeply impressed; Sternum VI with transverse sulci at base, apex deeply impressed,
impression bounded posteriorly by pollinose submarginal groove; female with deep lateral pit on Sternum IV; tibiae thick;
middle, hind tibiae each with two small, equal spurs; apicolateral process absent. Male unknown.

This species is easily recognized by the separated apical tubercles of the elytra. These,
together with the subapical tubercles, form four tooth-like elevations bounding the deep
preapical impression.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

THE GESTROI GROUP

This group consists of three species, two from Sumatra and one from the Nicobar Islands.
They resemble the members of the myopicum group in most respects, but have the median
groove of the pronotum narrowly dilated, with its sides pollinose and its floor glabrous. In
contrast to the fairmairei group, the median groove is narrower than the median pits. The
elytral setae are more extensive than in the myopicum group, with at least one seta in the
intratubercular stria.

Phylogeny. — R. bonsae and R. propinquus are clearly closely related. They share the
following characters: outer carina of pronotum concave dorsad to marginal groove; parasutural
stria with many setae; median groove shallow, its margins glabrous posterior to posterior
median pit; antennae short, thick, outer segments short, cylindrical, disc-like; Segment XI as
wide as long, compressed, stylet short, conical; preapical tubercle strongly elevated; antennal
lobe glabrous; temporal lobes divergent posteriorly.

The third species, R. gestroi, is more distantly related, and has the following contrasting
characters: carina of pronotum convex; parasutural stria without setae; median groove basad to
posteriormedian pit deep, its margins pollinose; antennae longer, more slender, outer segments
oblate sphaeroidal; Segment XI longer than wide, not compressed; stylet longer, acute;
preapical tubercle scarcely elevated; antennal lobe largely pollinose; temporal lobes strongly
convergent posteriorly.

Rhyzodiastes ( Temoana ) gestroi (Grouvelle 1903) NEW COMBINATION

(Fig. 38)

Clinidium gestroi Grouvelle 1903: 136-137.

Rhyzodiastes gestroi (Grouvelle) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) male, labelled: “Luglio, Gunong Singalang, Beccari, 1878”
(MNHN). PARALECTOTYPES one specimen, sex not recorded, labelled: “Si Rambe, Modigliana, Sep. 1892”
(MNHN); one male, same label as lectotype (GEN); three specimens (GEN), labelled: “Syntype. Si Rambe, Modigliana,
XI 1-90- 1 1 1-91” are not R. gestroi but R. indigens. Both localities are in Sumatra.

Description. — Length 6.2-7.0 mm. Antennal stylet acuminate; antennal Segment XI longer than broad, scarcely
compressed; few basal setae on Segment X, absent from more proximal segments; antennae longer, more slender, than in
related species, outer segments oblate spheroids; Segments I-X each with subapical pollinose ring; head slightly longer
than wide; antennal lobe largely pollinose, with a few irregular glabrous areas; median lobe short, rather narrow,
shield-shaped, its tip obtuse, opposite anterior 0.33 of eye; frontal grooves rather broad, deep, pollinose; medial margins of
temporal lobe oblique, closest together posteriorly, where medial angles are narrowly separated, nearly closing frontal
space posteriorly; temporal lobe 2.5 longer than wide; temporal lobe evenly rounded posteriorly, with conspicuous pollinose
border; orbital groove complete, narrow; eye narrowly crescentic, 0.5 as long as temporal lobe; one temporal seta, in orbital
groove posterior to eye; postorbit pilose.

Pronotum moderately elongate; length/greatest width 1.44; ovate, basal margin strongly curved; apex strongly
narrowed; base moderately narrowed; lateral margins curved; median groove narrowly dilated, about 0.5 times as wide as
anterior median pit; posterior median pit separated from base by 0.2 of length of pronotum; median groove basad to
posterior median pit deep, margins pollinose; inner carina highest next to median groove, sloped gradually laterally to
paramedian groove; medial margin of outer carina with narrow strip of pollinosity; outer carina convex, relatively broad,
marginal groove linear, entire, pollinose, pronotal setae absent.

Elytra relatively short, broad, margins parallel near middle; base strongly, obliquely narrowed to humeri; apex broadly
rounded, cauda not distinct; stria pollinose; sutural, parasutural scarp-like, with lateral margin much higher than medial
margin; intratubercular, marginal striae impressed; base of parasutural stria bent medially; Interval III subcarinate, base
forming prominent medial angle; apex forming slightly elevated preapical tubercle; apical tubercles inflated, contiguous,
no pore beneath them in midline; intratubercular stria with one seta just anterior to preapical tubercle; apex of marginal
stria with four setae.

Metasternum without median sulcus; transverse sulci of abdominal sterna broadly interrupted at midline, shallow,
barely impressed, each sulcus with pit at medial end; female with deep lateral pit in Sternum IV; submarginal sulcus of

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Sternum VI well separated from transverse sulci; tibiae moderately slender, middle, hind tibiae with two spurs, these equal
in female, posterior spur smaller in male; anterior femur of male with minute ventral tooth near apex; male with all
trochanters pointed; calcars narrowly triangular, acute.

This species is easily recognized by the form of the temporal lobes, which nearly meet
posteriorly.

Range. — Known only from the west coast of Sumatra. In addition to the type material, we

have seen three specimens with the following label: “Gunung Singgaiang, Sumatra’s Westkust, 1800 m.
VII- 1 925, leg. E. Jacobson”. Of these specimens, there is one male (NMNH) and one male, one female (AMS).

Rhyzodiastes ( Temoana ) propinquus new species
(Fig. 39)

Type Material. — HOLOTYPE female, labelled: “Nicobars, Rhyzodiastes propinquus Grouv.” (MNHN). The
latter is an unpublished species name which we are happy to adopt.

Description. — Length 6.8 mm. Antennal stylet minute, conical; antennal Segment XI as broad as long, somewhat
compressed; few basal setae on Segment X, absent from more proximal segments; antennae very short, thick; outer
segments short, disc-like cylinders; tufts of minor setae present on Segments IV-X; Segments I-X each with subapical
pollinose ring; head slightly longer than wide; antennal lobe glabrous; median lobe short, hastate, its tip acute, opposite
anterior 0.33 of eye; frontal grooves deep, glabrous; medial margins of temporal lobes curved, closest together opposite
posterior part of eyes; temporal lobes appear to diverge posteriorly, because of broad posteriomedial glabrous area on each
temporal lobe; temporal lobe about 2.5 longer than wide; posterior margin of temporal lobe pilose; orbital groove complete;
eye narrowly crescentic, approximately 0.5 as long as temporal lobe; possibly small temporal seta, in posterior part of
orbital groove; postorbit pilose.

Pronotum elongate; length/greatest width 1.50; widest posterior to middle, ovate; basal margin curved, apex strongly
narrowed; base moderately narrowed; lateral margins curved; median groove narrowly dilated, about 0.5 as wide as
anterior median pit; posteriomedian pit separated from base by 0.2 of length of protonum; median groove based to
posteriomedian pit shallow, margins glabrous; inner carina highest next to median groove, sloped gradually laterally to
paramedian groove; medial margin of outer carina with narrow strip of pollinosity; outer carina in dorsal view appearing
narrow, because lateral 0.67 of outer carina is concave; marginal groove linear, entire, pollinose; pronotal setae absent.

Elytra moderately elongate; margins parallel near middle; base slightly narrowed to humeri; apex evenly rounded, not
forming cauda; striae pollinose; sutural, parasutural striae scarp-like, with lateral margin higher than medial margin;
intratubercular, marginal striae impressed, base of parasutural bent medially; base of Interval III forming prominent,
pilose angle; Interval III laterad to basal angle glabrous; apex of Interval III forming rounded, elevated preapical tubercle;
apical tubercles scarcely inflated, contiguous; round pore ventral to apical tubercles in midline; parasutural stria with six
setae forming complete row; one seta at base of Interval III; intratubercular stria with one seta at base, one seta opposite
anterior end of preapical tubercle; three or four setae in apex of marginal stria.

Metasternum with fine median sulcus; transverse sulci of abdominal sterna broadly interrupted at midline; deep,
pollinose; female with deep lateral pit in Sternum IV; SternUm VI with marginal groove, posterior 0.33 impressed; tibiae
moderately slender; middle, hind tibiae each with two equal spurs; male unknown.

This species is most similar to R. bonsae from which it can be distinguished by the apparent
divergence of the temporal lobes posteriorly, the glabrous base of Interval IV, and the
contiguous, scarcely inflated apical tubercles.

Rhyzodiastes ( Temoana ) bonsae new species
(Fig. 46)

Type Material. — HOLOTYPE female, labelled: “Sumatra, Mt. Tenggamoes, Lampongs”. (MNHN) The
locality is now spelled “Gunung Tanggamus”, and is near the southern tip of Sumatra.

Description. — Length 7.8 mm. Antennal stylet minute, conical; antennal Segment XI as broad as long, somewhat
compressed; few basal setae on Segment X, basal setae absent from more proximal segments; antennae very short, thick;
outer segments short, disc-like cylinders; Segments I-X each with subapical pollinose ring; head slightly longer than wide;
antennal lobe glabrous; median lobe short, hastate, its tip acute, opposite anterior 0.33 of eye; frontal grooves deep,
glabrous; medial margins of temporal lobes closest together opposite eyes, nearly parallel, very slight divergent posteriorly;
medial and posterior margins of temporal lobe pilose fringe of even width; glabrous area of each temporal lobe about 2.5
longer than wide; orbital groove complete; one small, inconspicuous temporal seta in posterior part of orbital groove;
postorbit with conspicuous, rather long golden pilosity.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Pronotum elongate; length/greatest width 1.54; ovate, widest near middle; lateral margins more strongly curved than
in R. propinquus , base curved; base rather strongly narrowed; apex very strongly narrowed; median groove narrowly
dilated, about 0.5 as wide as anteriomedian pit; posterior median pit separated from base by 0.2 of length of pronotum;
median groove basad to posterior median pit shallow, margins glabrous; inner carina highest next to median groove, sloped
gradually laterally to paramedian groove; medial margin of outer carina with narrow strip of pollinosity; outer carina in
dorsal view appearing narrow, because lateral 0.67 of outer carina is concave; marginal groove linear, pollinose, entire;
pronotal setae absent.

Elytra moderately elongate; margins parallel near middle; base slightly narrowed to humeri; apical tubercles
protruding, forming broad but distinct cauda; sutural stria with very narrow, inconspicuous line of pollinosity; remaining
striae with broader, more conspicuous pollinose lines; sutural, parasutural striae scarp-like, with lateral margin higher than
medial margin; intratubercular, marginal striae very shallow, scarcely impressed; base of parasutural stria bent medially;
base of Interval III forming prominent pilose medial angle; latter connected to humerus by band of pollinosity crossing
base of Interval III; apex of Interval III forming low preapical tubercles, these more rounded, further apart than R.
propinquus ; apical tubercles prominent, inflated, separately rounded medially, nearly touching at one point, medial
surfaces pollinose; large rounded pore ventrad to them in midline; parasutural stria with row of about 10 setae, base of this
row follows medially bent portion of parasutural stria to base of sutural stria, while most posterior setae of this row is
displaced slightly laterad to stria, arising from medial surface of preapical tubercle; one seta at base of Interval III laterad
to parasutural stria; one or two setae on preapical tubercle; one or two setae in apex of intratubercular stria; several setae
near apex of marginal stria.

Metasternum with faint trace of median sulcus in posterior 0.5; abdominal sterna with transverse sulci well defined but
glabrous, broadly interrupted in midline; small pit at medial end of each sulcus; female with deep, round lateral pit on
Sternum IV; Sternum VI with transverse sulci, narrow marginal groove, disc not impressed; tibiae moderately slender;
middle hind tibiae each with two equal spurs; male unknown.

Among species with the median groove narrowly dilated, this species may be recognized by
the nearly parallel medial margins of the temporal lobes and by the inflated, separately
rounded apical tubercles.

It is a pleasure to name this species for Madame Andree Bons, of the National Museum of
Natural History in Paris, in gratitude for the aid that she has given to us and to many other
coleopterists over the years.

THE FAIRMAIREI GROUP

There are four species in this group, all from the mainland of Southeast Asia. Among the
species with a tuft of minor setae on Segment IV, they are easily recognized by the greatly
dilated median groove of the pronotum.

Phytogeny. — The relationships within the group are relatively obvious. R.fairmairei differs
from all the rest in the great enlargement of the posteriomedian pit of the pronotum, and in the
absence of temporal and elytral setae. It is probably only distantly related to the three
remaining species, which have the median groove of even width, not dilated by the enlargement
of the posteriomedian pit; and which have one or more temporal setae, and, in most specimens,
elytral setae. Among these three species, the two Vietnamese species appear to be very closely
related, with R. spissicornis of Malaya being more distant. The latter species has a median
sulcus on the metasternum, equal tibial spurs, setae in the sutural stria, in the fourth interval,
and on the apical tubercle, and the head elongate, with margins oblique posterior to the eyes. In
the two Vietnamese species, the median sulcus is absent, the tibial spurs are unequal, and the
elytral setae are limited to the parasutural stria and the apex of the marginal stria, or else are
absent.

Rhyzodiastes ( Temoana ) fairmairei Grouvelle 1895b NEW COMBINATION

(Fig. 47)

Rhyzodiastes fairmairei Grouvelle 1895b: 762-763.
Clinidium fairmairei (Grouvelle) Grouvelle 1903.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Rhyzodiastes fairmairei (Grouvelle 1895b) Bell and Bell 1978.

(Incorrect citation: Grouvelle originally described the species in Rhyzodiastes.)

Type Material. — HOLOTYPE female, labelled: “Carin Cheba, 900-1100 m., L. Fea, V-XII-88” (MNHN). In
the original description, the locality is given as “Montagnes des Carin, district des Carin Cheba”. It is in eastern Burma.

Description. — Length 7.0 mm. (our measurement; Grouvelle gives the length as 6.5 mm). Antennal stylet acute,
rather long; antennal Segment XI somewhat compressed, as wide as long; outer antennal segments very short, thick,
cylindrical, disc-like; basal setae absent; Segments I-X each with subapical pollinose ring; latter interrupted ventrally on
Segments IV-X by tuft of minor setae; head 1.25 longer than wide; antennal lobe glabrous; median lobe short, hastate, tip
acute, extending slightly posterior level of anterior margin of eye; frontal grooves rather broad, deep, pollinose; antennal
groove oblique, deep, pollinose; temporal lobe three times longer than wide; medial margins curved, closest together
opposite posterior margin of eye; posterior margin rounded, broadly fringed with pilosity, latter extended obliquely
anteriorly along medial margin; orbital groove complete, pollinose; eye crescentic, rather short, about 0.33 as long as
temporal lobe; temporal seta absent.

Pronotum elongate, length/greatest width 1.51; oval, widest at middle, lateral margins curved; base, apex both
narrowed; base strongly curved; median groove very broad, resembling keyhole, posterior portion (posteriomedian pit)
round, about 0.33 of width of pronotum; anterior portion with parallel margins, about 0.2 of width of pronotum; inner
carinae broad, sloped laterally; medial margin of outer carina with narrow strip of pollinosity; in dorsal view, outer carina
appears narrow, curved; outer carina with lateral surface concave, bounded ventrally by poorly defined marginal groove;
submarginal groove, pronotal setae absent.

Elytra elongate, rather narrow; lateral margins parallel near middle; humeri abruptly, obliquely narrowed; sutural,
parasutural stria finely punctate, impressed, scarp-like, with lateral margin of each much higher than medial margin, base
of parasutural stria bent medially; intercalary, marginal striae rather coarsely punctate, scarcely impressed except at apex;
subapical tubercle slightly elevated; apical tubercles inflated, contiguous at midline; elytral setae entirely absent.

Metasternum without median sulcus; transverse sulci of abdomen broadly interrupted at midline; sulci largely
glabrous, each with median, lateral pit; lateral pit of Sternum IV enlarged in female; submarginal sulcus of Sternum VI
widely separated from transverse sulcus at base, latter scarcely impressed, appearing as pair of pits on either side; middle
and hind tibia each with spur equal. Male unknown.

The broad, median groove of this species resembles a keyhole, and is unique within the
genus.

Rhyzodiastes ( Temoana ) spissicornis Fairmaire 1895 NEW COMBINATION

(Fig. 48)

Rhyzodiastes spissicornis Fairmaire 1895: 11-12.

Clinidium spissicorne (Fairmaire) Grouvelle 1903.

Rhyzodiastes spissicornis (Fairmaire) Bell and Bell 1978.

(Incorrect citation; Fairmaire originally described the species in Rhyzodiastes.)

Type Material. — LECTOTYPE male, labelled: “Puolo Pinang, Raffray, Ty., voisin de parumcostatum de Fairm
de Madagascar” (MNHN). PARALECTOTYPE female, labelled: “Singapore” (MNHN).

Description. — Length 5. 5-7.0 mm. Antennal stylet small, conical; antennal Segment XI compressed, as broad as
long; basal setae absent; antennae very short, thick; outer segments very short cylinders, 2.5 wider than long; tufts of minor
setae present on Segments IV-X; Segments I-X each with subapical pollinose ring; head elongate, length/width 1.5;
antennal lobe glabrous; medial lobe very short, shield-shaped, its tip obtusely pointed, at level with anterior margin of eye;
frontal grooves deep, glabrous; medial margins of temporal lobe curved, closest together opposite middle of eye; temporal
lobes divergent posteriorly; temporal lobe three times longer than wide; posterior half of medial margin, posterior margin
of temporal lobe fringed with pilosity; orbital groove complete, sinuate; eye crescentic, short, 0.5 as long as temporal lobe;
one small temporal seta, halfway between posterior end of eye, occipital angle; postorbits pilose.

Pronotum elongate; length/greatest width 1.55; widest posterior to middle, ovate; basal margin curved; base
moderately narrowed; apex strongly so; lateral margins curved; median groove dilated, about 10 times longer than broad,
gradually narrowed anteriorly; anterior median pit separated from apex of pronotum by about 0.1 of pronotal length;
posterior median pit separated from base by about 0.33 of length of pronotum; median groove very shallowly impressed in
front of anterior median pit and behind posterior median pit; inner carina highest next to median groove, sloped gradually
laterally to paramedian groove; medial margin of outer carina with narrow strip of pollinosity; outer carina in dorsal view
appearing narrow, because lateral 0.67 of outer carina is concave; marginal groove represented only by inconspicuous line
of minute pollinosity at ventral margin of concavity; pronotal setae absent.

Elytra elongate; margins parallel near middle; narrowed near humeri; latter prominent, tooth-like in dorsal aspect;
apex evenly rounded, not forming cauda; all striae scarp-like, with lateral margin higher than medial one; sutural,
parasutural, marginal impressed, pollinose; intratubercular with base, apex impressed, pollinose, middle not pollinose,
scarcely impressed; base of parasutural stria bent medially; base of Interval III forming prominent medial angle, latter

Quaest. Ent., 1985, 21 (1)

Bell and Bell

fringed with pilosity; base of Interval III with prominent lateral swelling just posterior to humeral angle; apex of Interval
III forming narrow, elevated preapical tubercle; apical tubercles slightly inflated, contiguous; sutural, parasutural stria
and Interval IV each with complete row of many setae; posthumeral elevation, apical tubercle with setae; apex of marginal
stria with several setae.

Metasternum with median sulcus; transverse sulci of abdomen broad, deeply impressed, each with narrow transverse
line of pollinosity; transverse grooves well separated at midline; those of Sternum VI slightly oblique, well separated from
submarginal groove; lateral pit of Sternum IV enlarged in female; tibiae thick; middle, hind tibiae each with two equal
spurs; ventral surface of anterior femur of male with many minute tubercles, but without ventral tooth; male with front,
hind trochanters pointed; calcars acute, triangular.

This species can be recognized by the dilated median groove in combination with the great
development of elytral setae. The elongate head, with relatively short eyes, also separates it
from the two species from Viet Nam.

Range. — Malay Peninsula. In addition to the type material we have seen the following

Specimens: one female, labelled: “Perka” (BMNH); one female, labelled: “Penang” (BMNH); one female, labelled:
“Malaya, Kuala Lumpur, 90 m. VI-7- 1962, coll. E. S. Ross and D. Q. Cavagnaro” (CAS), one male, labelled: “P. Penang,
Raffray” (GEN), one male, labelled: “P. Penang, 600-800 M., Loria e Fea” (GEN), also one male without locality label
(MNHN).

Rhyzodiastes ( Temoana ) alveus new species
(Figs. 49, 52)

Type Material. — HOLOTYPE male, labelled: “Hoa Binh, Tonkin, de Cooman, B.M. 1929-299” (BMNH).
PARATYPES two males, same label as holotype (BMNH); one male, three females, same label as holotype except that
acquisition number reads “B,M, 1925-251” (BMNH).

Description. — Length 5.0-6. 8 mm. Antennal stylet small, conical; antennal Segment XI not compressed, longer
than wide, basal setae absent; antennae moderately short, thick; outer segments oblate spheroids, less than two times wider
than long; tufts of minor setae present on Segments IV-X; pollinosity in some specimens present on all antennal segments,
in other specimens limited to Segments I-III; head slightly longer than wide; median lobe short, shield-shaped, its tip
acute, opposite anterior margin of eye; frontal grooves deep, glabrous; medial margins of temporal lobes curved, closest
together opposite middle of eye; posterior 0.5 of medial margin, lateral margin posterior to eye, with long pilosity; glabrous
area of temporal lobe three times longer than wide, tapered to point posteriorly, its medial margin concave opposite
posterior 0.5 of medial margin; orbital groove complete, margin with lateral pilosity posterior to eye; eye narrow,
crescentic, larger than R. spissicornis, 0.67 as long as temporal lobe; one small temporal seta opposite posterior margin of
eye; postorbit pilose.

Pronotum moderately elongate; length/greatest width 1.44; widest at middle; base, apex equally narrowed; lateral
margins moderately curved; base moderately curved; median groove dilated, its deep portion Five times longer than wide,
margins nearly parallel; posterior median pit separated from base by about 0.25 of length of protonum; median groove
shallowly impressed in front of anterior median pit, behind posterior median pit; inner carina highest next to median
groove, sloped gradually to paramedian groove; medial margin of outer carina with narrow strip of pollinosity; Fine line of
pollinosity in concavity of outer carina, distant from lateral margin and notopleural suture (Fig. 52); outer carina, in dorsal
view, appearing narrow because lateral 0.67 of outer carina is concave; pronotal setae absent.

Elytra moderately elongate; margins parallel near middle, narrowed slightly near humeri; apex evenly rounded, not
forming cauda; all striae scarp-like, with lateral margin much higher than medial one; all striae impressed, with very Fine
line of minute pollinosity; marginal stria interrupted posteriorly, apical portion detached, on ventral surface of apical
tubercle; base of parasutural stria bent medially; base of Interval III forming prominent pilose medial angle; base of
Interval III without lateral swelling; apex of Interval III forming narrow, elevated preapical tubercle; in posterior view,
posterior margin of preapical tubercle emarginate, apex overhanging base; apical tubercles slightly inflated, contiguous;
apex of marginal stria with several setae; elytron otherwise without setae.

Metasternum without median sulcus; transverse sulci of abdominal sterna narrow, not interrupted at midline, with pair
of dilated pits on either side of midline; those of Sternum VI connected laterally to base of marginal groove; both sexes
with lateral pit in Sternum IV, that of female larger than that of male; tibiae relatively slender; spurs of middle and hind
tibiae unequal, medial one about 0.5 as long as lateral one; male with ventral tooth on anterior femur; male with hind
trochanter pointed; calcars blunt, middle one narrow, hind one triangular, its proximal margins slightly angulate.

The short head, lack of a median metasternal sulcus, and greatly reduced elytral setae
separate this species from R. spissicornis. The absence of setae from the parasutural stria and
the broader median groove separate it from R.fossatus.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Range. — Northern Viet Nam, west and north of Hanoi. In addition to the type series, we

have seen the following specimens: two females, labelled: “N. Viet Nam, northwest of Tam Dao, Shou-Zuong,
1-2-1962, 200, 300 m., Kabakov” (LEN); one female, same data, 300 m., 20-2-1962 (LEN); one female, same data,
31-1-1962 (LEN); one male, labelled: “North Viet Nam, hills 50 km. NW of Thai-Nguyen, 19-12-1962, 400 m. Kabakov”
(LEN); one male, same data, except 9-3-1963, 300 m. Kabakov (LEN); one female, same data, except 8-II-1963, Kabakov
(LEN).

Variation. — The type series, from southwest of the Son Koi (Red River), have subapical
pollinose rings on all antennal segments, and have a pollinose area on the lateral surface of the
hind, and in most specimens, of the middle tibia. Specimens from northeast of the Song Koi
(Thai-Nguyen, Tam Dao) have pollinose rings only on antennal segments I and II. and lack
pollinosity on the lateral surfaces of the middle and hind tibiae, although there is a pollinose
line on the posterior face of the tibia. The latter is also present in the specimens from Hoa Binh.
Further collecting may demonstrate that the northeastern populations represent a separate
species or subspecies.

Rhyzodiastes ( Temoana ) fossatus new species
(Figs. 50, 53, 54)

Type Material. — HOLOTYPE male, labelled: “N. VIET NAM, hills s.w. Kui Chau, 300 m. 14-1-1963,
Kabakov” (LEN). PARATYPES five males, five females, same locality, several dates from 12-1-1963 to 15-11-1963
(LEN). The locality is in the north part of the former Annam, about 200 Km. south of Hanoi.

Description. — Length 5.6-7.0 mm. Antennal stylet small, conical; antennal Segment XI slightly compressed, as
wide as long; basal setae absent; antennae moderately thick, short; outer segments oblate spheroids, less than two times
wider than long; tufts of minor setae present on Segments IV-X; Segments I-X with subapical pollinose rings, in some
specimens interrupted near some of the apical setae; head slightly longer than wide; median lobe short, shield-shaped, tip
acute, opposite anterior margin of eye; frontal grooves deep, glabrous; medial margins of temporal lobes curved closest
together opposite middle of eye; posterior 0.5 of medial margin, lateral margin posterior to eye, with long pilosity; glabrous
area of temporal lobe three times longer than wide, tapered to point posteriorly, its medial margin oblique, straight or
nearly so opposite posterior 0.5 of medial margin; orbital groove complete, merging with lateral pilosity posteriorly; eye
narrow, crescentic, 0.67 as long as temporal lobe; one small temporal seta opposite posterior margin of eye; postorbit pilose.

Pronotum moderately elongate; length/greatest width 1.42; widest at middle; base, apex equally narrowed; lateral
margins moderately curved; base moderately curved, median groove dilated, but narrower than in R. alveus , deep portion
six times longer than wide, margins nearly parallel; floor of groove minutely pollinose with narrow glabrous median line;
posterior median pit separated from base by about 0.20 of length of pronotum; median groove shallowly impressed in front
of anterior pit, behind posterior median pit; inner carina highest next to median groove, sloped gradually to paramedian
groove; medial margin of outer carina with narrow strip of pollinosity; line of pollinosity along lateral margin of pronotum
just dorsad to notopleural suture (Fig. 53); outer carina, in dorsal view, appearing narrow because lateral 0.67 of outer
carina is concave; pronotal setae absent.

Elytra moderately elongate; margins parallel near middle, narrowed slightly near humeri; apex evenly rounded, not
forming cauda; all striae scarplike, with lateral margin much higher than medial margin; all striae impressed, with line of
pollinosity; marginal stria interrupted posteriorly, apical portion detached, on ventral surface of apical tubercle; base of
parasutural stria bent medially; base of Interval III forming prominent pilose medial angle; base of Interval III without
lateral swelling; apex of Interval III forming narrow, elevated preapical tubercle; in posterior view, posterior margin of
preapical tubercle emarginate, apex overhanging base; apical tubercles slightly inflated, contiguous; parasutural stria with
five to eight setae, in a few specimens forming complete row, in most specimens with a gap near middle; apex of marginal
stria with several setae (Fig. 54).

Metasternum without median sulcus; transverse sulci of abdominal sterna narrow, not interrupted at midline, with pair
of dilated pits on either side of midline; those of Sternum VI connected laterally to base of marginal groove of Sternum VI;
both sexes with lateral pit in Sternum IV, that of female larger than that of male; tibiae relatively slender; spurs of middle,
hind tibiae unequal, medial one about 0.5 as long as lateral one; male with ventral tooth on anterior femur; male with hind
trochanter pointed; calcars as in R. alveus ; lateral surface of tibia in both sexes with extensive pollinose area containing
glabrous tubercles.

This species is close to R. alveus , from which it differs most conspicuously in the presence of
setae in the parasutural stria and in having a narrower, minutely pollinose median groove on
the pronotum.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

SUBGENUS RHYZOSTRIX NEW SUBGENUS

Type species. — Rhyzodes maderiensis Chevrolat 1873a.

Description. — Antennal stylet present, though minute in some species; tufts of minor setae present on Segments
IV-X; clypeal setae present; compound eye relatively broad, oval; posterior margin of temporal lobe broadly pilose; gena
with curved band of pollinosity ventrad to eye; temporal seta absent; inner carina of pronotum sloped gradually to
paramedian groove; pollinosity limited to narrow line on medial slope of outer carina; base of pronotum with pollinose
border; paramedian grooves straight to slightly curved; outer carina not greatly enlarged or broadened at middle; pronotal
setae absent; intercalary stria absent; elytral striae coarsely punctate.

The large, oval eyes are diagnostic of this subgenus. Otherwise, it is superficially similar to
Temoana. The very coarsely punctate elytral striae will separate it from all except R. (T.)
sulcicollis. The appearance is quite different from Rhyzodiastes sensu stricto, the other
subgenus found in South America. The latter genus has narrow, costate outer carinae, smaller,
more crescentic eyes, and broadly pollinose paramedian and marginal grooves. However, the
two South American subgenera have some characters in common, such as pollinosity of the
gena in a curved, c-shaped band, elytral striae coarsely punctate, and posterior margin of
temporal lobe with very broad band of pollinosity. Perhaps they are related to one another.

Clinidium quadristriatum (Chevrolat) was used by Vulcano and Pereira (1975b) as the
name for a species of this genus. Their species is probably distinct from any known to us, but, as
indicated below, there is doubt as to whether C. quadristriatum is the correct name for it. They
illustrate another species under the name C. integrum Grouvelle, but this is not the species
described by Grouvelle (a Clinidium s. str.), but is probably yet another undescribed
Rhyzostrix.

Rhyzostrix is found in South America, in the Amazon Basin and Guiana and south along
the coast to Rio de Janeiro. It is largely allopatric to Rhyzodiastes sensu stricto.

Phytogeny. — Of the species known to us, R. menieri and R. maderiensis appear to be
closely related. If the convex, setose sutural interval is regarded as a synapomorphy, R. nitidus
is closer to the two preceding than to R. davidsoni. If the flat non-setose sutural interval of the
latter species is an apomorphy, R. davidsoni may be merely the most specialized species, and
not the most phylogenetically distinct one. Current data are insufficient to choose between
these possible phylogenies or to place R. quadristriatus in the phylogeny.

KEY TO SPECIES

1 Elytral Intervals I-III undulating, irregular, invaded by enlarged strial

punctures R. quadristriatus (Chev), p. 49

I' Elytral Intervals I-III not undulating, irregular 2

2 (1') Sutural interval flat, without setae; antenna without basal setae; hind

calcar cultrate R. davidsoni new species, p. 49

2' Sutural interval convex, with two to four setae near apex; basal setae of

antenna present; hind calcar straight 3

3 (2') Punctures of sutural, parasutural striae very coarse, nearly as broad as

Interval II; Stria III with pilosity limited to punctures; tip of preapical

tubercle slightly dentate, its posterior margin emarginate; Sternum VI of
female not impressed R. nitidus new species, p. 52

3 Punctures of sutural, parasutural striae smaller, less than 0.5 of width of

Interval II; Stria III with continual pollinosity; tip of preapical tubercle not
dentate, rounded posteriorly; Sternum VI of female impressed, with

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

tubercle at midline 4

4 (30 Sternum VI of female with pollinose pit posterior to small tubercle;

Sternum IV without lateral pits in the female

R. menieri new species, p. 52

4' Sternum VI of female with pair of pollinose areas or crescent area posterior
to tubercle; Sternum IV of female with lateral pits evident, though small

R. maderiensis (Chevrolat), p. 53

Rhyzodiastes ( Rhyzostrix ) quadristriatus (Chevrolat 1873a) NEW COMBINATION

Rhyzodes quadristriatus Chevrolat 1873a: 211.

Clinidium quadristriatum (Chevrolat) Grouvelle 1903.

Rhyzodiastes quadristriatus (Chevrolat) Bell and Bell 1978.

Type Material. — HOLOTYPE (sex not stated) according to the original description “Cayenna ex museo
Banoni”. We have not been able to locate the type, which was not studied by Vulcano and Pereira (1975b). “Cayenne”
refers to French Guiana.

Description. — Vulcano and Pereira (1975b) assigned a specimen from Brazil (Para, Taperinha perto de
Santarem, 1-10, VII. 1927 Zerny leg.) to this species. We have not studied this specimen. As indicated in our key, it differs
from all species seen by us in having Intervals l-III undulating and irregular, invaded by enlarged punctures of sutural and
parasutural striae. However, there is doubt as to whether the Chevrolat name really applies to this specimen. The original
description does not mention the undulating, irregular intervals. Grouvelle (1903) did cite undulating, irregular intervals
as characteristic of this species, and of R. maderiensis as well, but did not state that he had studied the type of R.
quadristriatus. Unless the type can be located, R. quadristriatus should probably be regarded as a nomen dubium, and the
specimen attributed to it by Vulcano and Pereira should be given a new name.

Rhyzodiastes ( Rhyzostrix ) davidsoni new species
(Figs. 51,55,62, 64)

Type Material. — HOLOTYPE male, labelled: “Brazil, Amazonas, 1 km. W. Taruma Falls, 100 m., 11-1-1981,
coll. R. Davidson” (CMP). PARATYPES three males, three females, same data as holotype (CMP); one male, labelled:
“Manaus, Amazonas, Brasil, VIII- 1962, coll. K. Lenko” (MZSP).

Description. — Length 5. 9-6. 8 mm. Antennal stylet minute; basal setae of antenna absent; head longer than wide;
frontal grooves very narrow, shallow; median lobe longer than in related species, its tip even with middle of eye; gena with
horizontal pollinose line just below eye, but without curved ventral continuation.

Pronotum short for subgenus, length/greatest width 1.53; suboval, with apex more truncate, hind angles more distinct
than in other members of subgenus; widest just posterior to middle; lateral margins constricted just anterior to middle,
width anterior to constriction almost equal to greatest width; marginal groove strongly abbreviated posteriorly, ending just
posterior to middle of pronotum.

Elytra elongate, lateral margins parallel through most of length; humeri narrowed; sutural stria, fine with about 10
moderately fine punctures; parasutural striae impressed, wider than others, with about 10 coarse punctures: intercalary,
marginal striae impressed, rather finely punctate; sutural interval completely flat (Fig. 62); second interval convex,
subcarinate; third interval elevated above parasutural stria, medial margin broadly pollinose; third intervals strongly
convergent anteriorly; apex of third interval forming elevated preapical tubercle, latter with posterior margin strongly
emarginate; preapical tubercles dentate, nearly contiguous in midline; apical tubercle with one or two setae; apex of
marginal stria with several setae; sutural interval without setae.

Metasternum with complete, deep, median sulcus; abdominal sterna each with narrow, coarsely punctate transverse
sulcus; Sulci III-IV complete, V, VI complete or narrowly interrupted in midline; submarginal sulcus of Sternum VI of
female with expanded pit at either side (Fig. 55), male without such expanded pit; middle, hind femora of male angulate
beneath; hind trochanter pointed in male; middle calcar very narrow, straight, acute; hind calcar elevated above tibial
spurs, strongly cultrate (Fig. 64).

The flat sutural interval, entirely without setae, differentiates this species from the rest of the subgenus. The strongly
dentate preapical tubercles, short pronotum with distinct hind angles and truncate apex, and the curved, hooklike hind
calcars, are also diagnostic.

Range. — In addition to the type series we have seen three males, three females, labelled:

“Brasil, Amazonas, BR. 174, Km. 18, 5-XII-1979, Elias Brasil” (INPA).

Quaest. Ent., 1985,21 (1)

Bell and Bell

Plate 6. Figs. 59-65. Genus Rhyzodiastes , new Subgenus Rhyzostrix. Figs. 59-62, Right elytron, posterior aspect; Fig. 59,
R. (R.) menieri new species; Fig. 60, R. (R.) nitidus new species; Fig. 61, R. (R.) maderiensis (Chevrolat); Fig. 62, R. (R.)
davidsoni new species; Figs. 63-64, Hind tibia, apex, male; Fig. 63, R. (R.) maderiensis (Chevrolat); Fig. 64, R. (R.)
davidsoni new species; Figs. 65-70, Head and pronotum, dorsal aspect; Fig. 65, R. (R.) maderiensis (Chevrolat). Figs.
66-74. Subgenus Rhyzodiastes sensu stricto. Fig. 66, R. (s. str.) pentacyclus new species; Fig. 67, R. (s. str.) liratus
(Newman); Fig. 68, R. (s. str.) parumcostatus (Fairmaire); Fig. 69, R. (s. str.) suturalis new species; Fig. 70, R. (s. str.)
costatus (Chevrolat); Figs. 71-73, Left elytron, dorsal aspect; Fig. 71, R. (s. str.) liratus (Newman); Fig. 72, R. (s. str.)
suturalis new species; Fig. 73, R. (s. str.) costatus (Chevrolat); Fig. 74, Antennal Segments IX-XI, R. (s. str.) pentacyclus
new species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent., 1985,21 (1)

Bell and Bell

Variation. — One of the females from Taruma Falls has a tubercle in the middle of Sternum
VI of the abdomen, while the other two females lack any trace of a tubercle. This might prove
to be a specific character; however, the tuberculate and one of the nontuberculate females were
taken in copula with apparently identical males. It is possible that the females are morphs of a
polymorphic population, analagous to Clinidium veneficum Lewis.

In addition to type material, we provisionally assign to this species a male, labelled:
“Taracua, Rio Uaupes, Amazonas, Brasil, VIII- 1964 Pereira and Machado” (MZSP). It
conforms to the description of R. davidsoni in most respects, but has a low second interval, only
slightly more convex than the sutural interval. The hind trochanter is less distinctly pointed
than in the type series. This locality is far to the west of Manaus, and is near the Colombian
border. This form might be a distinct, though closely related species, a subspecies, or the
differences might be clinal. A decision must await collections in the intervening area.

Rhyzodiastes ( Rhyzostrix ) nitidus new species
(Figs. 56, 60)

Type Material — HOLOTYPE male, labelled: “Santarem, Brazil, Acct. No. 2966” (CMP). PARATYPES one
male, one female, same data as holotype (CMP); two males, one female, labelled: “Rio de Jan., Brazil, Acct. No. 2966”
(CMP); one male, one female, labelled: “Amaz., Para” (MNHN). The female of this pair is labelled: “ Clinidium nitidum
Grouv.”, an unpublished name.

Description. — Length 6.0-7. 2 mm. Antennal stylet small, narrowly conical; basal setae present on Segment
VII-X; head longer than wide; frontal grooves moderately wide, shallow; median lobe very short, its tip opposite anterior
margin of eye; gena with curved pollinose line.

Pronotum oval, elongate; length/greatest width 1.59; widest near middle; base strongly curved; apex moderately
curved; hind angles indistinct; marginal groove over 0.6 as long as pronotal margin, separated from base, apex by 0.2 of
length of margin.

Elytra elongate, lateral margins parallel through most of length; humeri narrowed; sutural stria impressed, coarsely
punctured, punctures nearly as wide as Interval II; parasutural stria deeply impressed, coarsely punctured, both striae with
about 8 elongate punctures; intratubercular, marginal striae broad, impressed, with medial margins sloped gradually from
intervals, coarsely punctate, though less coarse than punctures of sutural, parasutural striae; sutural intervals together
convex (Fig. 60); Interval II convex, lower than I or III; Interval III elevated above parasutural stria, medial margin
broadly pollinose; bases of third intervals weakly convergent; apex of third interval thickened, forming elevated preapical
tubercle, latter with posterior margin emarginate; preapical tubercles weakly dentate, separated from one another by width
of one sutural interval; sutural interval with two to four setae in apical 0.5; apical tubercle with one seta; apex of marginal
stria with several setae.

Metasternum with complete median sulcus; abdominal Sterna III- VI each with narrow, coarsely punctate transverse
sulcus, narrowly interrupted at midline; female with deep, large, round lateral pit on Sternum IV (Fig. 56); male without
lateral pit; Sternum IV alike in both sexes, not impressed; middle, hind calcars similar, narrow, straight, acute.

The very coarse punctures of the sutural and parasutural striae and the separated pilose
punctures of Stria III are distinctive of this species. The dentate preapical tubercle will separate
it from R. menieri and R. maderiensis, while the convex and setose sutural interval separates it
from R. davidsoni.

Range. — Coastal lands of Brazil, from the lower Amazon south to Rio de Janeiro.

Rhyzodiastes ( Rhyzostrix ) menieri new species
(Figs. 57, 59)

Type Material. — HOLOTYPE female, labelled: “GUYANE, Haut-Carsevenne, F. Geay, 1898” (MNHN).

Description. — Length 7.1 mm. Antennal stylet minute, conical; basal setae present on Segments VII-X; head
slightly longer than wide; frontal grooves shallow, moderately narrow; median lobe with tip opposite anterior 0.25 of eye;
gena with curved pollinose line.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Pronotum oval, elongate, length/greatest width 1.57, widest just posterior to middle; margin, marginal groove,
paramedian groove slightly constricted near middle; base strongly curved; apex moderately curved; hind angles indistinct;
marginal groove of pronotum nearly complete, separated from basal pollinosity by 0.1 of length of pronotum.

Elytra elongate, lateral margins parallel through most of length; humeri narrowed; sutural stria deeply impressed, with
about 10 punctures, latter less than 0.25 as wide as Interval II; parasutural stria impressed, with 10 punctures;
intratubercular, marginal striae broad, impressed, with medial margins sloped gradually from intervals, rather finely
punctate; sutural intervals together, convex (Fig. 59); Interval II convex, lower than I or III; Interval III elevated above
parasutural stria, medial margin broadly pollinose; bases of third intervals weakly convergent; apex of third interval
thickened, forming elevated preapical tubercle, latter with posterior margin rounded, not dentate; preapical tubercles
separated by combined width of both sutural intervals; sutural interval with two setae near apex; apical tubercle with one
seta; apex of marginal stria with several setae.

Metasternum with complete shallow median sulcus; abdominal Sterna 1 1 1- VI, each with coarsely punctate transverse
sulcus, narrowly interrupted at midline; female without lateral pit on Sternum IV (Fig. 57); Sternum VI in female
impressed in apical 0.33; impression bounded anteriorly in midline by small tubercle; small median pollinose pit posterior
to tubercle; male unknown.

This species is close to R. maderiensis , but the female differs in having a smaller tubercle
with one median pollinose pit posterior to it on Sternum VI and in lacking the lateral pit on
Sternum IV.

It is a pleasure to name this species for Dr. Jean-Jacques Menier of the Museum National
d’Histoire Naturelle in appreciation of his aid in our study of Rhysodini.

Rhyzodiastes ( Rhyzostrix ) maderiensis (Chevrolat 1873a) NEW COMBINATION

(Figs. 58,61,63,65)

Rhyzodes maderiensis Chevrolat 1873a: 21 1-212.

Clinidium maderiensis (Chevrolat) Grouvelle 1903.

Rhyzodiastes maderiensis (Chevrolat) Bell and Bell 1978.

Type Material. — HOLOTYPE (sex not specified), according to the original description, labelled “Madereo”,
and collected by Lethierryo. The type locality refers to the Rio Madeira, a major tributary of the Amazon River. We have
not studied the type specimen but Vulcano and Pereira (1965b) have seen the type from the Vienna Museum collection.
The figure and description agree with specimens seen by us, and on which the description below is based.

Description. — Length 6.0-7.0 mm (according to Vulcano and Pereira, five to eight mm). Antennal stylet minute,
conical; basal seta present on Segments VI-X or VII-X; head longer than wide; frontal grooves shallow, moderately
narrow; median lobe with tip opposite anterior 0.25 of eye; gena with curved pollinose line.

Pronotum oval, elongate; length/greatest width averaging 1.60, ranging from 1.55-1.65, widest just posterior to
middle, margin slightly constricted at middle, marginal, paramedian grooves slightly sinuate opposite constriction; base
strongly curved; apex moderately curved; hind angles indistinct; marginal groove of pronotum less complete than in R.
menieri, separated from basal pollinosity by 0.2 or more of length of pronotum.

Elytra elongate, lateral margins parallel through most of length; humeri narrowed; sutural stria deeply impressed, with
about 10 punctures, latter less than 0.25 as wide as Interval II; parasutural stria impressed; with about 10 punctures like
those of sutural stria; intratubercular, marginal stria broadly impressed, with medial margin sloped gradually from
intervals, rather Finely punctate; sutural intervals together convex (Fig. 61); Interval II convex, lower than I or III; Interval
III elevated above parasutural stria, medial margin broadly pollinose; bases of third intervals weakly convergent; apex of
third interval thickened, forming elevated preapical tubercle, latter with posterior margin rounded, not dentate; preapical
tubercles separated by combined width of both sutural intervals; sutural interval with three to five setae in apical 0.67;
apical tubercle with one seta or without; apex of marginal stria with several setae.

Metasternum with median sulcus incomplete, anterior part effaced; abdominal Sterna III-VI narrow, coarsely
punctate; all sulci narrowly interrupted in midline in female; sulci of Sterna III, IV not interrupted in male; Sternum VI
not impressed in male; Sternum VI of female with apical 0.33 deeply impressed, impression bounded anteriorly in midline
by tubercle; pair of median pollinose pits posterior to tubercle (Fig. 58); male with femora obtusely angulate ventrally;
both pairs of calcars acute, triangular, straight; hind calcar larger, more broad based than middle one (Fig. 63).

R. maderiensis is close to R. menieri but differs in the female having a larger tubercle on
Sternum VI with paired pollinose pits or a crescent shaped pit and in having a lateral pit on
Sternum IV.

Range. — We have seen the following specimens all from Manaus, Brasil: three males, four
females, Manaus, 1 km. W. Taruma Falls, 100 m., 1 1-1-1981, coll. R. Davidson, (CMP); two males, three females, VIII,
1962, coll. K. Lenko (MZSP); one male, one female, 26- VIII- 1 962, coll. W. L. Brown (MZSP). Manaus is about 125

Quaest. Ent., 1985, 21 (1)

Bell and Bell

kilometers west of the mouth of the Rio Madeira. The latter is listed as the type locality, but there is no information as to
where on the river it was taken.

SUBGENUS RHYZODIASTES SENSU STRICTO

Type species. — Rhyzodes parumcostatus Fairmaire 1868.

Description. — Antennal stylet compressed, broad, obliquely truncate, resembling chisel blade; tufts of minor setae
begin on Segment IV, V, or VI; clypeal setae present; eye crescentic, narrow in most species, broad in one species; gena
with curved band of pollinosity; pronotum elongate; inner carina with lateral margin sharply defined; paramedian groove
broad, pollinose, at least 0.5 as wide as outer carina; marginal groove broad, sharply defined, pollinose, visible in dorsal
view; pronotal setae absent; elytron with intercalary stria absent; elytral intervals, especially Interval III costate (least so in
R. pentacyclus ); elytral setae absent; metasternum with median sulcus.

This subgenus is easily recognized by the broad, pollinose paramedian grooves, narrow,
sharply defined inner carinae, and broad, chisel-like antennal stylet. It is found in southern and
eastern Brazil, in the coastal mountains and the Mato Grosso, and reaches northern Argentina.
It apparently does not penetrate the Amazon Basin.

Phytogeny. — R. pentacyclus is the most distinctive species and probably represents the
sister group to the remaining species. The elytral intervals are not costate, while among the
remaining species at least Interval III is strongly costate. In this character state, R. pentacyclus
is obviously the least modified member of the subgenus. The absence of tufts of minor setae
from antennal Segments IV and V is probably also plesiomorphic, if it is accepted that the
general tendency in the subtribe has been for the number of tufted segments to increase. On the
other hand, the circular form of the tuft, with a raised rim, is probably an apomorphy for R.
pentacyclus.

R. suturalis resembles R. pentacyclus in having a deeply impressed sutural stria and
relatively limited pollinosity on the ventral surface. However, it resembles the remaining
species in having the minor setae in transverse, unrimmed tufts. It is perhaps the sister species
of the remaining species. It has tufts on antennal Segments V-X. The remaining species, R.
liratus, R. costatus , and R. parumcostatus are closely related, with a very narrow sutural
interval, sutural stria scarcely impressed, and ventral surface strongly pollinose. R.
parumcostatus has tufts on Segments IV-X, in contrast to the two remaining species, which
have them on Segments V-X.

KEY TO SPECIES

1 Minor setae in round, rimmed tufts on Segments VI-X

R. pentacyclus new species, p. 55

T Minor setae in transverse, oval tufts on Segments V-X or IV-X 2

2 (T) Minor setae on Segments V-X 3

2' Minor setae on Segments IV-X R. parumcostatus (Fairmaire), p. 55

3 (2) Sutural stria deeply impressed, coarsely punctate; lateral margin of

pronotum sinuate anterior to hind angle R. suturalis new species, p. 59
3' Sutural stria very shallow, impunctate or finely, shallowly punctate; lateral

margin or pronotum not sinuate anterior to hind angle 4

4 (3') Sutural stria pollinose, impunctate or with shallow punctures; medial

margin of parasutural stria pollinose; hind calcar slightly to strongly

convex dorsally R. liratus (Newman), p. 56

4' Sutural stria not impressed, represented by shallow, pollinose punctures,

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

not pollinose between punctures; medial margin of parasutural stria
glabrous; hind calcar narrow, triangular, dorsal margin straight
R. costatus (Chevrolat), p. 58

Rhyzodiastes ( sensu stricto) pentacyclus new species
(Figs. 66, 74)

Type Material. — HOLOTYPE male, labelled: “BRASILIA, Alto da Serra, Stanzel-Lachnit, CNHM-1955.
Karl Brancik Colin., ex Eduard Knirsch” (AMNH). PARATYPES one male, labelled:. “A. Serra, 1921” (collector’s name
illegible) (MZSP); two females, labelled: “Est. Biol. Boraceia, Salesopolis, Sao Paulo, BRAZIL, 17-10-1960, 12-V-1961,
K. Lenko col.” (MZSP); one male, same data as preceding but dated 17-10-1960, (MZSP); four specimens with same data
as preceding but dated as follows: one male, one female 16-1 9-VIII- 1 966, Biasi, Costa & Silva (MZSP); one male, V-966,
E. Rabalo (MZSP), one female, 21-22-III-1973, J. Vanin & M. Jorge, “sob a casco de tronco caido” (MZSP); two males,
two females, labelled “Paranapiacaba, S. P. Brasil, 30-IX-1974, EXP. MUS. ZOOL, col tronco caido” (MZSP); one male,
without locality label (MZSP).

Description. — Length 6.7-8. 9 mm. Each tuft of minor setae in flat, circular space, surrounded by raised rim,
present on Segments VI-X (Fig. 74); basal setae present on Segments VII-X; head short, length/greatest width 1.1;
median lobe long, triangular, tip opposite posterior margin of eye, obtusely pointed; glabrous part of temporal lobe oblique,
length/greatest width 4.0, glabrous area separated posteriorly from lateral margin of head by broad pollinose space, latter
wider than glabrous area; eye crescentic, rather narrow; eye separated from posteriolateral angle of head by 0.3 of length
of eye, medial margin of eye straight.

Pronotum short for subgenus, length/greatest width about 1.48; widest posterior to middle, base moderately narrowed;
apex very strongly narrowed; lateral margins curved, base oblique on either side of midline; apex truncate; median groove
narrow between median pits; anterior median pit broad; posterior median pit narrower, separated from base by 0.30 of
length of pronotum; median groove posterior to it broad, containing secondary, shallower pit at base; paramedian grooves
relatively narrow, sinuate; marginal groove dilated, about 0.5 as wide as outer carina at middle; inner carina sinuate,
broadest posterior to middle, where three times as wide as paramedian groove; outer carina of nearly even width, widest
near middle, where about 0.5 as wide as greatest width of inner carina; narrow marginal carina visible in dorsal view;
prosternum with tubercle posterior to coxa.

Elytra elongate, moderately narrow; sutural stria impressed, with about 12 very coarse punctures; parasutural stria
impressed, with 12-14 coarse punctures, anteriorly equal to sutural stria, posteriorly becoming slightly more dilated;
intratubercular stria impressed, very coarsely punctate, abruptly narrowed opposite preapical tubercle; marginal stria
impressed, coarsely punctate; punctures of all striae each about as wide as elytral interval; intervals glabrous, convex;
sutural interval only slightly less convex than Interval II; latter tapered posteriorly; Interval III with apex forming
preapical tubercle, latter less prominent than in other members of subgenus; apical tubercle scarcely inflated; metasternum
glabrous, with median sulcus; transverse sulci of abdominal Sterna V, VI narrowly interrupted at midline, those of III, IV
continuous in female, narrowly interrupted in male; male with small lateral pit on Sternum IV, female with large one;
anterior femur of male with many minute round tubercles belqw; middle, hind tibiae with traces of tubercles; middle calcar
acute, straight, very slender; hind calcar cultrate, its ventral margin raised well above bases of spurs.

The circular tufts of minor setae, with raised rims, are distinctive of this species, as is the
absence of tufts from Segments IV and V. The inner pronotal carinae are broader than in other
species, and the elytral intervals are nearly equal and not costate.

Variation. — The series from Salesopolis differ from the remaining specimens in having the
paramedian groove closed or nearly closed anteriorly by a junction of the inner and outer
carinae. The outer carinae of the pronotum also averages narrower than in the remaining
localities, Alto da Serra and Paranapiacaba.

Rhyzodiastes (sensu stricto ) parumcostatus (Fairmaire 1868) NEW COMBINATION

(Fig. 68)

Rhyzodes parum-costatus Fairmaire 1868: 782.

Clinidium parumcostatum (Fairmaire) Dajoz 1975.

Rhyzodiastes parumcostatus (Fairmaire) Bell and Bell 1978.

This species was synonymized with Clinidium costatum (Chevrolat) by Arrow (1942), and
so regarded by Hincks (1950). Dajoz (1975) recognized it as a distinct species and listed
differences between it and C. costatum.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Type Material. — HOLOTYPE male, labelled: “Madag., Rhysodes parumcostatum Fairmaire” (MNHN). This
must be a mislabelling, as the species is represented by numerous specimens from Brazil and one from northern Argentina,
and has never been collected in Madagascar.

Description. — Length 5.6-7.7 mm. Each tuft of minor setae is oval, transverse depression, not rimmed; tufts
present on Segments IV-X; basal setae present on Segments VII-X; head relatively elongate, length/greatest width about
1.23; median lobe glabrous medially, pollinose laterally, tip acute, opposite to posterior portion of eye; glabrous part of
temporal lobe slightly curved, about 4.5 longer than wide; posteriolateral angle of head nearly rectangular, widely
spearated from glabrous part of temporal lobe; eye crescentic in lateral view; medial margin of eye slightly curved; gena
with curved band of pollinosity, and diffuse pollinose area ventroposterior to it; in some specimens gena entirely pollinose.

Pronotum elongate; length/greatest width about 1.60; widest near middle; base, apex only slightly narrowed; lateral
margins nearly straight, parallel, very slightly undulating; apex truncate; base strongly curved; median groove moderately
broad between median pits; anterior median pit much broader than median groove; posterior median pit inconspicuous,
small, separated from base by 0.3 of length of pronotum; secondary, inconspicuous posteriomedian pit at base of pronotum;
median groove entirely pollinose; paramedian, marginal grooves broad, deep, pollinose; pronotal carinae largely pollinose,
but each with narrow glabrous line; those of inner carinae strongly undulating; those of outer carinae nearly straight,
complete (most specimens) or undulating, abbreviated posteriorly (southern specimens); marginal groove with very narrow
glabrous line; prosternum without tubercle posterior to coxa.

Elytra elongate, narrow, convex; sutural stria not impressed, scarcely evident, with about 1 2 punctures, coarse in most
specimens, in some specimens scarcely evident; parasutural stria deeply impressed, with about 12 coarse punctures;
intratubercular stria impressed, coarsely punctate; marginal stria broad, shallow, scarcely impressed except near apex;
parasutural stria glabrous between punctures; other striae pollinose; sutural interval flat, represented by very narrow
glabrous line; Intervals II, III subcostate, largely pollinose, but with glabrous line; that of II complete; that of III complete
in some specimens, limited to anterior 0.25 and preapical tubercle in others; Interval IV slightly convex, with glabrous line
near humerus, latter incomplete in some specimens; preapical tubercles slightly inflated, rounded posteriorly; apical
tubercles scarcely inflated; metasternum with median sulcus; metasternum largely pollinose, but with glabrous area on
either side of sulcus anterior to hind coxae; transverse sulci of abdominal sterna not interrupted in midline in most
specimens, in a few (both sexes) narrowly interrupted on V or VI, VI; abdominal sterna extensively pollinose, both in
transverse sulci, and along posterior margin of each sternum; median longitudinal pollinose area connecting transverse
sulcus with posterior margin on Sterna III, IV; male with lateral pit scarcely evident on Sternum IV; that of female deep;
anterior femur of male with many minute tubercles on ventral surface; middle calcar acute, straight, very slender; hind
calcar triangular, acute, moderately narrow, slightly cultrate, proximal margin convex, distal margin concave.

This species is the only member of the subgenus in which the tufts of minor setae begin on
antennal Segment IV.

Range. — Southeastern Brazil and northern Argentina. We have studied the following

specimens: ARGENTINA: one male, labelled: “Misiones, Dep. Concep., Sta. Maria X-1948, M. J. Viana” (MZSP).
This is one of the specimens which Viana (1951) recorded as C. costatus Chevrolat. Viana listed two females and one
additional male, and also one female from Santiago del Estero. We have not located these specimens, which were in
Viana’s personal collection; 19 specimens, labelled: “Rep. Arg., Misiones” without date or collector (MNHN). BRAZIL:
SANTA CATARINA: three females, labelled: “Corupa (Hansa Humboldt), Nov. 1945, Dec. 1944, A. Mailer coll., Frank
Johnson, donor” (AMNH); one male, labelled: “Hansa, Sta. Catarina, VIII, 1910, Leuderw.” (MZSP); three males,
labelled: “Sainte Catherine, Deyrolle 1847” (MNHN); two males, Santa Cath. (BMNH); BRAZIL, SAO PAULO, one
female, labelled: “Caioba, 25-50, 48-40 (latitude, longitude), 10 m., F. Plaumann IV-1965” (MZSP); one female, labelled:
“Cantareira, S.P. 20-11-1958, K. Lenko” (MZSP); two females, labelled “Ilha de Vitoria, S. Paulo 16-27 III, 1964, Exp.
Dep. Zool.” (MZSP); five males, ten females, labelled: “Ilha dos Buzios, S. Paulo, 16-X-4-XI 1963, Exp. Dep. Zool.”
(MZSP); one female, labelled: “Brasil, Cn Fairm” (GEN).

There are five additional specimens (BMNH) without precise locality data.

Variation. — The specimens from Argentina, and some of those from Santa Catarina differ
from more northern specimens in having the pollinosity more extensive, with the glabrous lines
of the outer carinae abbreviated posteriorly and those of Interval III obsolete except at the base
and on the preapical tubercle. These might represent an additional taxon, but more specimens
are required to confirm it.

Rhyzodiastes {sensu stricto) liratus (Newman 1838) NEW COMBINATION

(Figs. 67,71)

Rhysodes liratus Newman 1838: 665-666.
Clinidium liratum (Newman) Fairmaire 1873b.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Rhyzodiastes liratus (Newman) Bell and Bell 1978.

Grouvelle (1903) synonymized this species with Clinidium costatum (Chevrolat), while Bell and Bell (1978)
resurrected it.

Type Material. — LECTOTYPE (here designated) female, labelled: “ Rhysodes liratus Newm., Rio” (BMNH).
According to the original description, it was collected by Charles Darwin. PARALECTOTYPES. According to Newman,
there were five specimens in the type series. We have not located any in addition to the lectotype, and do not know whether
any of the paralectotypes are still preserved. Newman indicated that they were in Darwin’s personal collection.

Description. — Length 6.2-7. 5 mm. Each tuft of minor setae in oval transverse depression, latter not rimmed; tufts
present on Segments V-X; head elongate, length/greatest width about 1.38; median lobe narrow, elongate, extending
posterior to middle of eye, glabrous part of temporal lobe very narrow, elongate; pollinosity of orbital groove as broad as
glabrous part of temporal lobe; posteriolateral portion of head completely pilose; eye broad, nearly semicircular in lateral
view; eye separated from posteriolateral angle of head by 0.5 of length of eye; medial margin of eye curved.

Pronotum extremely elongate; length/greatest width about 1.83; base only slightly narrowed; apex moderately
narrowed; lateral margins nearly straight in some specimens, slightly emarginate anterior to middle in others; base
strongly curved; median groove glabrous medially, lateral slopes broadly pollinose; anterior median pit large; paramedian
grooves broad, deep, largely pollinose, but with narrow glabrous area in bottom; marginal groove broad, pollinose; inner
carina narrow, curved around median pits, in most specimens not broadened posteriorly, in some specimens distinctly
broadened posteriorly; outer carina narrow, abbreviated posteriorly; margin with very narrow pollinose line; propleuron
pollinose; prosternum pollinose except for part of intercoxal process; prosternum without postcoxal tubercle.

Elytra elongate, narrow, convex; sutural stria impunctate or faintly punctate, scarcely impressed, separated from
suture by very narrow flat glabrous interval (Fig. 71), parasutural stria very deeply impressed, with 12-15 deep coarse
punctures; intratubercular stria impressed, punctate, entirely pollinose; marginal stria shallowly impressed, punctate, but
punctures obscured by thick pollinosity; lateral margin of Interval III pollinose; Intervals II, III, IV forming narrow
glabrous carinae; preapical tubercles inflated, tapered posteriorly; apical tubercles inflated; metasternum with median
sulcus; metasternum pollinose except for posterior margin, lateral borders of median suclus; anterior part of abdomen
largely pollinose; transverse sulci narrowly interrupted in midline; transverse sulci each with row of coarse punctures;
female with deep lateral pit on Sternum IV; anterior femur of male with many minute tubercles on ventral surface; middle
calcar acute, triangular; hind calcar with dorsal margin convex, strongly so in most specimens, only slightly so in a few
specimens.

This species is most likely to be confused with R. costatus, which also has the tufts of minor
setae beginning on Segment V, and has the sutural stria scarcely impressed. The latter species
has the sutural stria punctate, with the pollinosity interrupted between the punctures. The
lateral margin of Interval II is glabrous. Also, the hind calcar is triangular, with the dorsal
margin not or scarcely convex, and the inner carinae of the pronotum are more broadened
posteriorly.

Range. — Southeastern Brazil, north to Bahia State and south to Sao Paulo State. All

localities are near to the Atlantic Coast. We have seen specimens from the following localities:
BAHIA, two females, labelled: “Bahia Lewis” (BMNH), one-female, two males, labelled: “Retiro, Bahia” (BMNH); two
males, labelled: “Una, Bahia, Oct. 27, Friedrich” (BSL); GUANABARA (former Federal District), two females, no
further data (MZSP); one female, labelled: “Corcovado 14-12-1945, Wygodzinsky” (MZSP); one male, labelled:
“Corcovado, Guanabara, 700 m., Nov. 1-7, 1963, Wygodzinsky” (AMNH); one male, labelled: “Rio de J., Wygodzinsky”
(BSL); RIO DE JANEIRO (STATE), one female, labelled: “Angra, E. do Rio, Pisseral. X-935, L. Tr, et Lopes”
(MZSP), one female, labelled: “Floresta de Dijuca, 1 7-VII- 1 960, R. Schubartel” (MZSP); SAO PAULO: one female,
labelled: “Santos, 17-23 2-99” (MNHB); one male, labelled: “Santos, 7-1 1-93” (MNHN); two males, one female, labelled:
“Sao Paulo, J. Metz” (CNHM); STATE UNCERTAIN: one female, labelled: “Mendes, 4-IX-33, Eidmans” (BSL); one
female, labelled: “P. N. do Itaiaia, 1.1958, L. C. Alvaranca” (MZSP). In addition, we have seen several specimens labelled
simply “Brazil”, including two members of the type series for R. costatus (Chevrolat) (NMW), labelled “ costatum ,
Brasilia, Chevrolat”.

Variation. — This species shows considerable variation in many characters, including the
shape of the pronotum, the length of the marginal carina, the distinctness of the posterior
median pit, and the convexity of the dorsal margin of the hind calcar. The sutural stria in most
specimens is narrowly pollinose and impunctate, but in a few specimens there are indistinct
punctures. The variation appears on the basis of very limited material to be geographical. The
specimens from Bahia State have the posterior median pit virtually absent, the outer carina as
long as the inner one, and the hind calcar less convex than in specimens from other areas. The
hind calcar varies considerably within this population, in some specimens being scarcely more

Quaest. Ent., 1985,21 (1)

Bell and Bell

convex than in R. costatus. The specimens from Rio de Janeiro have the posterior median pit
distinct, the outer carina as in the Bahia specimens, and the hind calcar strongly convex. The
specimens from Sao Paulo State have the posterior median pit distinct, the hind calcar strongly
convex, and the outer carina of the pronotum abbreviated posteriorly. The available specimens
are too few to be certain whether these differences represent subspecies or not. This species is in
need of more detailed study.

Rhyzodiastes ( sensu stricto) costatus (Chevrolat 1829)

(Figs. 70, 73)

Rhysodes costatus Chevrolat 1829, t. 18, f. 12, in Guerin-Meneville 1829-1844.

Rhyzodes costatus (Chevrolat) Chevrolat 1844 (altered spelling of generic name).

Clinidium costatum (Chevrolat) Lewis 1888.

Rhyzodiastes costatus (Chevrolat) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) female, labelled: “costatus, Guerin, Brasilia, Chevrolat”, with
red “typus” label (NMW). PARALECTOTYPES: The type series is a mixture of three species. One male and one female
are R. liratus. One male and one female are R. parumcostatus. All are labelled like the lectotype, and all are in NMW.
We have restricted the name to the lectotype, the one species in the series which has not been described elsewhere.

Description. — Length 6.9-7. 3 mm. Each tuft of minor setae in oval transverse impression, not rimmed; tufts
present on Segments V-X; head moderately elongate; length/greatest width about 1.29; median lobe narrow, elongate,
extending posterior to middle of eye; glabrous portion of temporal lobe narrow, elongate; pollinosity of orbital groove as
broad as glabrous portion of temporal lobe; posteriolateral portion of head completely pilose; eye broad, nearly
semicircular in lateral view; eye separated from posteriolateral angle of head by 0.5 of length of eye; medial margin of eye
curved.

Pronotum elongate; length/greatest width about 1.77; base only slightly narrowed; apex moderately narrowed; lateral
margins slightly curved; base strongly curved; median groove broad, pollinose; paramedian grooves broad, deep, largely
pollinose, but with narrow glabrous area in bottom; marginal groove broad, pollinose; inner carinae narrow, curved around
anterior median pits, broader posteriorly, where distinctly broader than paramedian groove; outer carina narrow, of even
width; marginal carina slightly narrower than outer carina; propleuron pollinose; prosternum pollinose except for part of
intercoxal process; postcoxal tubercle absent.

Elytra elongate, narrow, convex; sutural stria not impressed, represented by row of shallow pollinose punctures,
pollinosity absent between punctures except in some specimens, where present in posterior 0.25 of stria (Fig. 73);
parasutural stria impressed, punctured, medial margin glabrous, lateral margin pollinose; intratubercular stria deeply
impressed, pollinose; marginal stria shallowly impressed, punctures obscured by thick pollinosity; Interval II only
moderately convex; Intervals III, IV forming narrow, glabrous carinae; preapical tubercles scarcely inflated posteriorly;
apical tubercles slightly inflated; metasternum with median sulcus; metasternum very finely pollinose or microsculptured;
abdominal sterna dull, very finely pollinose or microsculptured; transverse sulci narrowly interrupted in midline, each with
pit at medial end, otherwise impunctate or obscurely punctate; female with deep lateral pit on Sternum IV; anterior femur
of male with many minute tubercles on ventral surface; middle calcar acute, triangular; hind calcar narrow, acute,
distinctly proximad to tibial spurs.

This species differs from R. liratus in having the sutural stria represented by a row of
isolated punctures, these not connected by pollinosity except near elytral apex, in lacking
pollinosity on the medial margin of the parasutural stria (in other words, on the lateral margin
of the second interval), and in having the hind calcar narrowly triangular with the dorsal
margin straight.

Range. — Southern Brazil, except for one female, labelled: “Rio Jano., FRY” (BMNH), from more
inland localities than R. liratus. We have seen one male and three females, labelled “Matto Grosso, de Castelnau, 12-47”
(MNHN), and one male, labelled: “Vicosa, M. G., 23-7-57, coll. J. Becker” (MZSP). “M.G.” indicated Minas Gerais

State. The characters of this species are approached by some of the variants of R. liratus , and it
is possible that the two are only subspecifically distinct. However, the presence of both forms at
Rio de Janeiro makes this doubtful. Like R. liratus , this form needs more study.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Rhyzodiastes ( sensu stricto) suturalis new species
(Figs. 69, 72)

Type Material. — HOLOTYPE female, labelled: “Espiritu Santo, Sooretama, Linhares, X. 962, Pereira, Alv.
Martins” (MZSP).

Description. — Length 7.4 mm. Each tuft of minor setae in oval transverse depression, not rimmed; tufts present
on Segments V-X; head elongate, length 1.33 times greatest width; median lobe narrow, elongate, extending posterior to
middle of eye, glabrous part of median lobe narrow, only slightly broader than pollinosity of orbital groove; posteriolateral
portion of head completely pilose; eye broad, nearly semicircular in lateral view; medial margin of eye curved; eye
separated from posterior angle of head by 0.5 of length of eye.

Pronotum elongate; length/greatest width 1.71; base, apex slightly narrowed; lateral margin with long, shallow
sinuation anterior to hind angle; base strongly curved; median groove broad, bottom glabrous, margins pollinose;
paramedian grooves broad, deep, pollinose; marginal groove moderately broad, pollinose, abbreviated anterior to sinuation
of lateral margin; inner carinae narrow, convex, curved around anterior median pit; outer carina narrow, convex, complete;
marginal carina complete; base of pronotum narrowly margined with pollinosity; propleuron largely glabrous, but very
finely pollinose or microsculptured near notopleural suture; prosternum glabrous; postcoxal tubercle absent.

Elytra moderately narrow, elongate, but broader than in R. liratus, humeri strongly narrowed; sutural stria deeply
impressed, with about 11 coarse punctures, bottom glabrous, margins pollinose; parasutural stria deeply impressed,
becoming broader posteriorly, with about 12 coarse punctures; parasutural stria glabrous, lateral margin pollinose, medial
margin nearly glabrous, with trace of pollinosity posteriorly; intratubercular stria impressed, broad, coarsely punctate,
punctures obscured by pollinosity; marginal stria narrow, impressed, pollinose; sutural interval broad, convex, wider than
second interval, glabrous; second interval convex, posterior 0.5 subcarinate; third interval narrow convex, fourth interval
nearly flat; preapical tubercle strongly inflated, apex tapered; apical tubercle inflated; metasternum with median sulcus,
largely glabrous; abdominal sterna largely glabrous; transverse sulci narrowly interrupted in midline, with enlarged pits at
medial ends, coarsely punctate, narrowly pollinose; female with round lateral pit in Sternum IV; male unknown.

This species resembles R. liratus and R. costatus in having the minor setae in oval tufts on
Segments V-X. It differs from them in having the sutural stria very strongly impressed and
coarsely punctate and in the shape of the pronotum. The deep sutural stria gives it the
appearance of R. pentacyclus. The latter species, however, has the minor setae in circular,
rimmed tufts on Segments VI-X.

GENUS CLINIDIUM KIRBY 1835

Description. — Part I: 62

KEY TO SUBGENERA (slightly revised from Part I: 62)

1 Cleaning organ of anterior tibia entirely proximad to basal articulation of

anterior tarsus 2

V Cleaning organ more distad, basal articulation of tarsus opposite its

midpoint 4

2 (1) Tufts of minor setae present on Antennal Segments VII-X; pronotum

widest near middle; angular seta present; marginal setae absent;
intercalary stria ending blindly posteriorly, except in C. halffteri and some
C. guatemalenum Mexiclinidium Bell and Bell, p. 60

2' Tufts of minor setae present on antennal Segments VI-X; pronotum widest
distinctly behind middle; either both angular, marginal setae present, or
else both absent; intercalary stria not ending blindly posteriorly 3

3 (2') Parasutural stria complete anteriorly, reaching base of elytron; pronotum

without setae Protainoa Bell and Bell, p. 69

3' Parasutural stria restricted to posterior 0.5 to 0.25 of elytron; pronotum

Quaest. Ent., 1985, 21(1)

Bell and Bell

with marginal, angular setae Tainoa Bell and Bell, p. 70

4 (T) Marginal stria clearly sixth from suture; all striae well developed; inner
elytral intervals carinate, marginal groove of pronotum double or single

Arctoclinidium Bell, p. 75

4' Striation more reduced; marginal stria fourth or fifth from suture;
supramarginal stria absent; inner elytral intervals not carinate or scarcely
so; marginal groove of pronotum single Clinidium sensu stricto , p. 93

SUBGENUS MEXICLINIDIUM BELL AND BELL 1978

Type species. — Clinidium mexicanum Chevrolat 1873a.

Description. — Antenna with tufts of minor setae on Segments VII-X; antennal stylet small; 1 temporal seta
present; eye narrowly crescentic (narrower than in Arctoclinidium)-, orbital groove pollinose, complete, reaching posterior
margin of temporal lobe; pronotum with lateral margins curved, marginal groove double or single; pronotum with angular
setae (except for newtoni), but without marginal setae; sternopleural grooves absent; elytral striae complete; marginal stria
fifth or sixth from suture; supramarginal stria impressed in most specimens, represented by row of punctures in some
specimens, absent in C. championi-, intercalary stria ending blindly anterior to preapical tubercle, except in C. halffteri and
some specimens of C. guatemalenum-, intervals of elytra elevated, costate in most species; elytral setae more numerous than
in Arctoclinidium-, metasternum without median sulcus; female with enlarged lateral pit in Sternum IV; female without
elytral cauda; anterior femur of male with ventral tooth; anterior tibia of male with proximal tooth present or absent; base
of anterior tarsus entirely distad to cleaning organ; calcars small, hind calcar smaller than middle calcar.

The deep elytral striae and carinate intervals make most members of this subgenus
superficially similar to Arctoclinidium. The position of the cleaning organ and the more
numerous elytral setae separate it from the latter subgenus. Most Mexiclinidium differ from
Arctoclinidium in the anastomosis of Intervals III and IV posterior to the end of the intercalary
stria. In C. halffteri and some specimens of C. guatemalenum , the intercalary stria is complete
and the intervals do not anastomose.

Mexiclinidium is known from central and southern Mexico and from Guatemala.

Phylogeny. — The nine species can be grouped as follows:

I. mexicanum group

C. mexicanum
C. balli
C. triplehorni

II. blomi group

C. blomi
C. iviei

III. guatemalenum group

C. guatemalenum
C. newtoni

IV. championi group

C. championi
C. halffteri

The mexicanum group contains three very similar species which differ mainly in secondary
sexual characters. The group occupies a compact area on the Mexican Plateau and its eastern
margin. The outer marginal groove of the pronotum is shallow or absent, the transverse sulci of
the abdomen are deep and widely separated, elytral setae are few, the male first trochanter is
toothed, and Sternum VI has many round punctures.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

The blomi group has two species, one on either side of the gap in the mountain chain at
Tehuantepec. These have the outer marginal groove deep but hidden in dorsal view, the
transverse sulci deep and widely separated, the male first trochanter toothed. Elytral setae are
many, and Sternum VI either has elongate punctures (iviei) or else two pairs of impressions
{blomi).

The guatemalenum group has two species in the highlands of Chiapas and Guatemala. They
have a pair of precoxal setae. The outer marginal groove is deep and is visible in dorsal view.
The median lobe of the head is shorter than in the two preceding groups. The transverse sulci of
the abdomen are deep and are only narrowly separated medially. Elytral setae are few. The
male first trochanter is toothed ( newtoni ) or rounded {guatemalenum). Sternum VI is coarsely
punctate.

The championi group has two species, one in the Quiche Mountains of Guatemala, the other
from a relatively low elevation near the Gulf of Mexico. These species have Sternum VI with a
narrow submarginal groove and a pair of pits near the base. The first trochanter of the male is
rounded, and the transverse sulci of the abdomen are relatively shallow. Otherwise the two
species are quite dissimilar, and perhaps are not closely related. C. championi has the median
lobe of the head elongate, the outer marginal groove of the pronotum deep and visible in dorsal
view (as in the guatemalenum group), and the elytral setae few. C. halffteri has the median
lobe short and truncate, the outer marginal groove absent, and the elytral setae numerous.

KEY TO SPECIES

1 ' Outer marginal groove of pronotum visible in dorsal view 2

V Outer marginal groove of pronotum not visible in dorsal view 4

2 Transverse sulci of abdomen present, pilose; Sternum VI coarsely

punctured, submarginal sulcus absent 3

2' Transverse sulci of abdomen absent, represented by isolated punctures;

Sternum VI with crescent-shaped submarginal sulcus
C. championi new species, p. 62

3 Apex of pronotum narrowed, evenly curved; postorbital, suborbital tubercle
present; medial ends of transverse sulci without enlarged pits

C. newtoni new species, p. 63

3' Apex of pronotum truncate; postorbital, suborbital tubercle absent; medial
ends of transverse sulci with enlarged pits

C. guatemalenum Sharp, p. 63

4 Intercalary stria ending blindly posteriorly; Intervals III, IV anastomosing

posteriorly; median lobe of head elongate, tip acute, opposite or behind
posterior region of eye 5

4' Intercalary stria complete; Intervals III, IV not anastomosing; median lobe
of head short, tip truncate, opposite middle of eye
C. halffteri new species, p. 66

5 Sternum VI with a pair of median pilose, oval impressions

C. blomi Bell, p. 66

5' Sternum VI with scattered, round or elongate punctures 6

6 Punctures of Sternum VI elongate, coalesced; Sternum VI impressed

C. iviei new species, p. 69

Quaest. Ent., 1985,21 (1)

62 Bell and Bell

6' Punctures of Sternum VI large, round; Sternum VI unimpressed 7

7 Calcars present, males 8

T Calcars absent, females 10

8 Proximal tooth of anterior tibia present 9

8' Proximal tooth of anterior tibia absent

C. triplehorni new species, p. 68

9 Proximal tooth of anterior tibia large; femoral tooth large, almost carinate

C. mexicanum Chevrolat, p. 67

9' Proximal tooth of anterior tibia small, oblique; femoral tooth small, oblique
C. balli new species, p. 68

10 Lateral pit of Sternum IV glabrous 11

10' Lateral pit of Sternum IV pollinose C. triplehorni new species, p. 68

1 1 Basal impressions of pronotum relatively large, 0.25 of length of pronotum;

supramarginal stria impressed or represented by coarse punctures

C. mexicanum Chevrolat, p. 67

11' Basal impressions small, less than 0.20 of length of pronotum;

supramarginal stria not impressed, represented by fine punctures

C. balli new species, p. 68

Clinidium ( Mexiclinidium ) championi new species
(Figs. 75, 98)

Type Material. — HOLOTYPE male, labelled: “Quiche Mountains, 7-9000 ft.. Champion” (BMNH). This
locality is in Guatemala near Totonicapan.

Description. — Length 6.0 mm. Head as broad as long; median lobe long, tip opposite posterior margin of eye;
medial margin of temporal lobe nearly straight.

Pronotum relatively short; length/greatest width 1.30; lateral margin moderately curved; base slightly narrowed, apex
moderately narrowed; basal impression relatively large, length 0.33 of length of pronotum; basal impression closed
posteriorly; inner, outer marginal grooves equally deep, outer marginal groove conspicuous in dorsal view; marginal carina
curved, narrow; prosternum without precoxal seta on each side.

Striae impressed, coarsely punctured, narrowly pollinose; intervals convex, but not distinctly costate; supramarginal
stria absent; sutural stria with one seta near apex; intercalary stria with two setae in apical 0.33; intratubercular stria
without setae; marginal stria with five setae near apex; transverse sulci broadly interrupted in midline, scarcely impressed,
not pollinose, each represented by row of punctures; male with small pollinose lateral pit on Sternum IV; Sternum VI with
small round pit near each anteriolateral angle, curved submarginal groove (Fig. 98); male with small obtuse ventral tooth
on anterior femur; male with anterior trochanter rounded; anterior tibia of male without proximal tooth; middle calcar
triangular, acute, base relatively broad; hind calcar small, triangular, apex acute, distal margin raised well above level of
spurs; female unknown.

This species differs from all others in the subgenus except C. halffteri in having the striae
shallower and the inner intervals not truly costate. The reduction of the transverse sulci to rows
of punctures is also distinctive. The absence of precoxal setae and impunctate sixth sternum
easily separate it from the sympatric C. guatemalenum. C. halffteri differs in lacking the outer
marginal groove and in having the supramarginal stria impressed.

We have named this species for the collector, George Champion, who collected fine series of
Rhysodidae in Central America.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Clinidium ( Mexiclinidium ) newtoni new species
(Figs. 77, 86)

Type Material. — HOLOTYPE male, labelled: “MEXICO, Chiapas, 8 mi. N. Pueblo Nuevo S., 6000', cl. for.
26-27 VIII-73, N. 541 A. Newton” (BSRI).

Description. — Length 7.0 mm. Head slightly longer than broad; median lobe long, ended just anterior to posterior
margin of compound eye; frontal grooves relatively shallow, convergent posteriorly; medial margins of temporal lobes
oblique, convergent posteriorly; small postorbital, suborbital tubercles present.

Pronotum relatively short; length/greatest width 1.40; lateral margins curved; apex strongly narrowed; base
moderately narrowed; basal impression large, 0.3 of length of pronotum; basal impression open posteriorly; inner, outer
marginal grooves equally deep, outer marginal groove conspicuous in dorsal view; marginal carina curved, narrow;
prosternum with precoxal setae; angular seta apparently absent.

All striae, including supramarginal deeply impressed; sutural stria with one or two setae near apex; intercalary stria
without setae; intratubercular stria with two setae near apex; transverse sulci of abdominal sterna narrowly interrupted in
midline, medial ends of sulci not enlarged; Sternum VI with a few very coarse punctures; anterior femur of male with very
prominent ventral tooth, latter truncate with apex in form of oblique ridge; anterior trochanter of male with obtuse ventral
tooth; male anterior tibia without proximal tooth, posterior face of anterior tibia with conical tooth opposite middle of
cleaning organ (Fig. 86); middle calcar small, acute; hind calcar larger, dorsal margin emarginate near base (calcar thus
slightly falcate); female unknown.

The distinct postorbital and suborbital tubercles of this species are unique within the
subgenus. The presence of precoxal setae and the narrow interruption of the transverse sulci of
the sterna are points of similarity to C. guatemalenum , but the latter species has enlarged pits
at the medial ends of the transverse sulci, and the pronotum is much less narrowed and rounded
anteriorly. C. blomi, which is probably sympatric with C. newtoni, differs in having the
transverse sulci broadly interrupted, the precoxal setae absent, the pronotum much less
narrowed anteriorly, and in numerous secondary characters of the male.

The species is named in honor of the collector, Alfred F. Newton, Jr.

Clinidium ( Mexiclinidium ) guatemalenum Sharp 1899
(Figs. 76, 101)

Clinidium guatemalenum Sharp 1899: 489.

Clinidium ( Arctoclinidium ) guatemalenum (Sharp) Bell 1970.

Clinidium ( Mexiclinidium ) guatemalenum (Sharp) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) male, labelled: “GUATEMALA, San Geronimo, Vera Paz
Prov., coll. Champion” (BMNH). PARALECTOTYPES one male, three females, same data as lectotype (BMNH).

Description. — Length 7. 0-7. 7 mm. Head longer than broad; median lobe short, tip opposite middle of eye; medial
margin of temporal lobe curved.

Pronotum relatively elongate, length/greatest width 1.48; lateral margins curved; apex strongly narrowed, basal
margin moderately narrowed; basal impression relatively large, 0.4 of length of pronotum; basal impression closed
posteriorly, closed or open laterally; inner, outer marginal grooves equally deep, outer marginal groove conspicuous in
dorsal view; marginal carina curved, narrow; prosternum with precoxal seta on each side.

All striae, including supramarginal deeply impressed; sutural stria with one or two setae near apex; intercalary stria
with two or three setae in posterior 0.5; intratubercular stria with two or three setae in posterior 0.3; marginal stria with
three to five setae near apex; transverse sulci of abdominal sterna very narrowly interrupted in midline, medial end of each
sulcus with enlarged pit; Sternum VI coarsely punctate (Fig. 101); lateral pit of Sternum IV in female glabrous; anterior
femur of male, with acute, narrow, ventral tooth with one seta; male with anterior trochanter rounded; male anterior tibia
with large proximal tooth; middle calcar triangular, acute, base relatively broad; distal margin not elevated above spurs;
hind calcar small, triangular, apex acute, distal margin elevated above level of spurs; female without ventral tooth on
anterior femur.

In the form of the pronotum, this species is closest to C. blomi , but the coarsely punctate
Sternum VI separates it from the latter species, and the closely approximate medial pits on the
transverse sulci are unique within the subgenus. The presence of precoxal setae is shared only
with C. newtoni. The latter species differs in male secondary sexual characters and in the

Quaest. Ent., 1985,21 (1)

Bell and Bell

Plate 7. Figs. 75-86. Genus Clinidium, Subgenus Mexiclinidium. Figs. 75-83, Head and pronotum, dorsal aspect; Fig. 75,
C. (M.) championi new species; Fig. 76, C. (M.) guatemalenum Sharp; Fig. 77, C. (M.) newtoni new species; Fig. 78, C.
(M.) balli new species; Fig. 79, C. (M.) triplehorni new species; Fig. 80, C. (M.) blomi Bell; Fig. 81, C. (M.) mexicanum
Chevrolat; Fig. 82, C. (M.) iviei new species; Fig. 83, C. (M.) halffteri new species; Figs. 84-86, Anterior leg, male
(excluding tarsus); Fig. 84, C. (M.) iviei new species; Fig. 85, C. (M.) balli new species; Fig. 86, C. (M.) newtoni new
species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent., 1985, 21 (1)

Bell and Bell

absence of pits at the medial ends of transverse sulci and in the presence of postorbital and
suborbital tubercles.

Range. — Guatemala. In addition to the type series we have studied the following specimens:
two males, two females, labelled: “Chichivac, Chimaltenango Prov., 8600', Aug. 19, 1926, J. R. Sievin’’ (CAS); two
females, labelled: “El Quiche, 7.3 km from Chichicastenango, 14° 54' N, 91° 07' W, 2400 m., May 28, 1973, T. L. & L. J.
Erwin” (NMNH); one male, one female, labelled: “Quiche Mts., 8500-10500 ft., Totonicapam, coll. Champion”
(BMNH).

In the type series, the parasutural stria is interrupted for a short distance just anterior to the
apex of the intercalary stria, so that a narrow bridge connects Intervals II and III.

In the specimens from other localities, this is not true. The specimens from Chichivac have a
shallow median impression on the metasternum, though it does not form a discrete sulcus.
There is no trace of this impression in the type series. It is not certain whether these variations
are individual differences or represent geographic variation.

Clinidium ( Mexiclinidium ) blomi Bell 1970
(Fig. 80)

Clinidium ( Arctoclinidium ) blomi Bell 1970: 309.

Clinidium ( Mexiclinidium ) blomi (Bell) Bell and Bell 1978.

Type Material. — HOLOTYPE MCZ 31747, male, labelled: “Rancho Nuevo 8 1 /2 miles SE of San Cristobal de
las Casas, Chiapas, Mexico, coll. R. T. Bell, D. H. Van Horn, July 23, 1956” (MCZ). PARATYPES, three females
collected with the type (UVM); four males, eight females, collected at same locality. Sept. 1, 1967, by Ball, Erwin, and
Leech (ALB).

Description. — Length 6. 1-7.5 mm. Head nearly as broad as long; median lobe long, tip posterior to hind margin of
eye; medial margin of temporal lobe curved.

Pronotum relatively short; length/greatest width about 1.40; lateral margins curved; apex strongly narrowed; base
moderately narrowed; basal impression about 0.3 of length of pronotum; basal impression closed posteriorly, laterally;
inner, outer marginal grooves equally deep; outer groove placed more laterally than in C. guatemalenum, scarcely visible
in dorsal view; marginal carina curved, conspicuous; prosternum without precoxal setae.

All striae, including supramarginal, deeply impressed; sutural stria with two to four setae, most anterior of them
anterior to middle of stria in most specimens; intercalary stria with complete row of five setae, most anterior of them near
to elytral base; intratubercular stria with two or three setae near apex; marginal stria with five or six setae in apical 0.5;
transverse sulci of abdominal sterna broadly interrupted in midline, without medial pits; Sternum VI not punctate, but
with two pairs of oblique impressions; lateral pits of Sternum IV of female pollinose; anterior femur of male with large,
broad ventral tooth with one seta; anterior trochanter of male dentate; anterior tibia of male with large proximal tooth;
middle calcar acute, base narrow, not elevated above spurs; hind calcar small, triangular, apex acute; distal margin
elevated above level of spurs; female with anterior femur not dentate.

This species resembles C. guatemalenum in having inner and outer marginal grooves of the
pronotum equally developed, separated by a narrow marginal carina. In this species, however,
the marginal carina is directed more laterally than in C. guatemalenum so that the outer
groove is almost hidden in dorsal view. Unique to C. blomi are the great development of the
elytral setae and the absence of coarse punctures on Sternum VI.

Range. — High Plateau of Chiapas, southeastern Mexico. In addition to the type material

we have seen one specimen labelled: “Mexico: 5 mi. w. of San Cristobal, 7500', V-23-1961, J. M. Campbell”
(BSRI).

Clinidium ( Mexiclinidium ) halffteri new species
(Figs. 83, 99)

Type Material. — HOLOTYPE male, labelled: “MEXICO, Ver., Amates, 29-V-1964, Catemaco, Halffter,
Reyes” (MZSP). PARATYPES two males, same label as holotytpe (MZSP). The type locality is in southern Vera Cruz
State, near the Gulf of Mexico, at a low elevation.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Description. — Length 6. 1-6.5 mm. Head slightly longer than broad; median lobe short, tip subtruncate, opposite
middle of eye; frontal grooves very narrow; frontal space very small; medial margins of temporal lobes convergent to
narrowly separated medial angles, latter posterior to hind margin of eye; posteriomedial margin oblique.

Pronotum rather elongate; length/greatest width 1.58; lateral margins weakly curved; apex less narrowed than base;
basal impressions very small, 0.12 of length of pronotum, closed posteriorly; outer marginal groove entirely absent;
prosternum without precoxal setae.

Elytra much broader than pronotum; margins parallel for most of length; humeri strongly, obliquely narrowed; elytral
intervals convex, inner ones not carinate; intercalary stria complete; Intervals III, IV not anastomosing posteriorly;
intratubercular stria impressed at base, apex; middle portion not impressed, represented by row of coarse punctures;
supramarginal stria not impressed; represented by row of punctures very close to those of intratubercular stria in middle
0.33; base, apex absent; marginal stria with base, apex impressed, middle 0.33 represented by row of punctures; sutural
stria with two setae near apex; parasutural stria with four or five setae in apical 0.5; intercalary stria with complete row of
five or six setae; intratubercular stria with one seta near apex; marginal stria with four or five setae in apical 0.33;
transverse sulci of abdomen narrow, each with row of coarse punctures, broadly interrupted in midline; without enlarged
punctures at medial end; Sternum VI of abdomen with small pit near each anteriolateral angle, long submarginal groove,
its end angled medially (Fig. 99); anterior femur of male with large, sharp ventral tooth, without setae; anterior trochanter
of male rounded; anterior tibia of male without proximal tooth; middle calcar slender, rather long, apex obtuse; hind calcar
triangular, apex obtuse, only slightly elevated above level of spurs; female unknown.

This species is distinguished from all other members of the subgenus in having the median
lobe short and truncate, and the temporal lobe with distinct medial angles. The pronotum is
also distinctive, with the base slightly more narrowed than the apex and the outer marginal
groove entirely absent. It is the only member of the subgenus to have setae in the parasutural
stria.

The species is named for the collector, Dr. Gonzalo Halffter, a skilled specialist in
Scarabaeidae and insect behavior.

Clinidium ( Mexiclinidium ) mexicanum Chevrolat 1873a
(Figs. 81, 100)

Clinidium mexicanum Chevrolat 1873a: 214.

Clinidium ( Arctoclinidium ) mexicanum (Chevrolat) Bell 1970.

Clinidium ( Mexiclinidium ) mexicanum (Chevrolat) Bell and Bell 1978.

Type Material. — According to the original description, there is a type series of seven specimens, collected in
May, 1855 by Aguste Salle, under pine bark at Jacale, at the Park of Orizaba. We have studied one male and one female
of this series, labelled: “Jacale, Mexico, Salle coll.” (BMNH). Both are labelled as cotypes. We have not located the other
five specimens. The pair which we have studied fit the concept of C. mexicanum of previous authors in all respects except
one; both specimens are virtually without the outer marginal groove of the pronotum. This is probably an individual
variation, as we have seen a few specimens from other localities which have the groove strongly reduced. Nevertheless,
there might be two taxa in the type series, so we feel it improper to designate a lectotype until we have seen the rest of the
series.

Description. — Length 6. 0-8. 5 mm. Head slightly longer than broad; median lobe long, tip posterior to hind
margin of eye; medial margins of temporal lobes nearly straight, nearly parallel.

Pronotum relatively elongate; length/greatest width about 1.47; lateral margins curved, apex strongly narrowed; base
moderately narrowed; basal impressions small, less than 0.25 of length of pronotum, open posteriorly; outer marginal
groove completely hidden in dorsal view, shallower than inner marginal groove, fine but complete in most specimens,
effaced anteriorly in a few specimens, nearly absent in a few specimens; prosternum without precoxal setae.

Supramarginal stria shallow in most specimens, in some specimens not impressed, represented by row of coarse
punctures; remaining striae impressed; sutural stria with two or three setae near apex; intercalary stria with two or three
setae posterior to middle; intratubercular stria with two or three setae posterior to junction with supramarginal; marginal
stria with five or six setae near apex; transverse sulci of abdomen broadly interrupted in midline, without pits at medial
ends; Sternum VI of abdomen coarsely punctate (Fig. 100); lateral pit of Sternum IV in female glabrous; anterior femur of
male with large, sharp ventral tooth with several setae; anterior trochanter of male dentate; anterior tibia of male with
large proximal tooth; middle calcar acute, base moderately broad, elevated above level of spurs; hind calcar small, less
acute than in C. blomi, base broad, elevated well above level of spurs.

This species is most easily separated from C. triplehorni and C. balli by the secondary
sexual characters of the male. It is also larger than the two related species, and has the

Quaest. Ent., 1985, 21 (1)

Bell and Bell

supramarginal stria better developed.

Range. — Mountains of the southern end of the Mexican Plateau, from Jalisco to Vera Cruz
State, from 5000 to 12,000 feet elevation. Bell (1970) gives a list of localities. We have seen

Specimens from the following additional localities. MEXICO STATE: Temescaltepec, long series of both
sexes, coll. Hinton, Usinger (BMNH) (this locality is near to Tejupilco des Hidalgo); four males, four females 18 km. SW
of Toluca, meadow, 3400 m., April 22, 1977, coll. J. S. Ashe, H. E. Frania, D. Shpeley (ALB); MORELOS: one female, 7
mi. s. of Tres Cumbres, VII-7, 1975, coll. Triplehorn (OSU); PUEBLA, one male, one female, 50.8 km. se of Azuabilla,
2480 m., oak pine forest, logs, ground, 78B-36a, Dec. 24-25, 1978, G.E., K. E. Ball (ALB); six males, nine females, 37.5
km. se of Azuabilla, 2500 m., wet oak-pine, 78B-37, G.E., K.E. Ball (ALB); one female, 7.6 km. e. of Santa Maria del
Monte, wet pine-oak forest, 2480 m., VII-9-1975, G. E. Ball, H. E. Frania (ALB). These new records do not significantly
extend the range of the species.

Variation. — The most significant variation is in the develoment of the outer marginal
groove of the pronotum. In a large majority of specimens, it is complete. In a few, the anterior
part is effaced, and in the two cotypes studied by us, it is entirely effaced, as in C. balli and C.
triplehorni. It appears to us that this is an individual aberration, without taxonomic
importance, though its presence in both the cotypes is surprising.

Clinidium ( Mexiclinidium ) balli new species
(Figs. 78, 85, 97)

Type Material. — HOLOTYPE male, labelled: “G. Ball Colin., MEXICO: Hgo, 16 mi. N. Zimapan 8000', at
night, V-27-1974, C. & L. O’Brien & Marshall” (NMNH). PARATYPES two males, one female, same data as type
(NMNH); three males, two females, labelled: “MEX:S. Luis Potosi, 14 mi. W. Xilitla, 4800'; VI-29-73, A. Newton”
(MCZ); one male labelled: “MEX, Hidalgo, 4 mi. S.W. Chapalhuacan, 3500', VII-5-1976, A. Newton” (MCZ).

Description. — Length 5. 0-6.0 mm. Head slightly longer than broad; median lobe not quite so long as in C.
mexicanum, tip even with posterior margin of eye, blunter than in C. mexicanunr, medial margins straight, parallel.

Pronotum elongate; length/greatest width 1.54; lateral margins curved; apex slightly more narrowed than base; basal
impression very small, about 0.15 of length of pronotum, open posteriorly; outer marginal groove almost absent, less than
0.1 of length of pronotum, ventrad to hind angle; prosternum without precoxal setae.

Supramarginal stria not impressed, represented by row of fine punctures; marginal stria impressed at apex,
represented at middle by row of fine punctures; both marginal, supramarginal striae with anterior 0.33 entirely effaced;
sutural stria with one seta near apex; intercalary stria with one seta near apex; intratubercular stria without setae;
marginal stria with three or four setae near apex; transverse sulci of abdomen broadly interrupted at midline, without pits
at medial ends; Sternum VI of abdomen coarsely punctate; lateral pit of Sternum IV in female glabrous (Fig. 97); anterior
femur of male with small, oblique ventral tooth, without setae; anterior trochanter of male dentate; anterior tibia of male
with small, obtuse proximal tooth; middle calcar acute, base narrow, not raised above level of spurs; hind calcar small, base
broad, scarcely raised above level of spurs.

This species is close to C. mexicanum , but has the supramarginal and marginal striae
effaced anteriorly. The males is easily recognized by the small proximal tooth on the anterior
tibia and the oblique tooth on the anterior femur (Fig. 85). C. triplehorni has the proximal
tooth entirely lacking.

We take pleasure in naming this species for our longtime friend, Dr. George Ball, who has
made long series of Mexican Clinidium available to us.

Clinidium ( Mexiclinidium ) triplehorni new species
(Figs. 79, 96)

Type Material. — HOLOTYPE male, labelled: “MEXICO, Hgo. 7 mi. ne. of Jacala, VI-23-1975, C. A., W. E.,
B. W. Triplehorn” (OSU). According to Dr. Triplehorn (in litt.), the elevation of the type locality is about 3200'. Despite
the similar spelling, this locality is not the same as Jacale, the type locality for C. mexicanum. PARATYPES four males,
two females, same label as holotype (OSU).

Description. — Length 6. 0-6. 7 mm. Head slightly longer than broad; median lobe long, tip posterior to hind
margin of eye, blunter than in C. mexicanum ; medial margins of temporal lobes straight, nearly parallel.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Pronotum elongate, length/greatest width 1.54; lateral margins curved; apex strongly narrowed; base moderately
narrowed; basal impression small, about 0.2 of length of pronotum, open posteriorly; outer marginal groove reduced to 0.2
of length of pronotum, ventrad to hind angle; prosternum without precoxal setae.

Supramarginal stria not impressed, represented by row of very fine punctures; marginal stria impressed near apex,
base, otherwise represented by row of fine punctures; sutural stria without setae or with one near apex; intercalary stria
without setae or with one or two near apex; intratubercular with one seta near apex; marginal stria with three or four near
apex; transverse sulci of abdomen broadly interrupted in midline, without pits at medial ends; Sternum VI of abdomen
coarsely punctate; lateral pit of Sternum IV in female pollinose (Fig. 96); femur of male with very small obtuse ventral
tooth, without setae; anterior trochanter of male dentate; anterior tibia of male without proximal tooth; middle calcar
acute, base broad, not raised above level of spurs; hind calcar small, acute, base narrow, raised well above level of spurs.

This species resembles C. mexicanum, but differs strongly in secondary sexual characters,
entirely lacking the proximal tooth on the anterior tibia in the male. The female, unlike C.
mexicanum and C. balli , has the lateral pit of Sternum IV pollinose.

We dedicate this species to the collector, Dr. Charles Triplehorn, in appreciation of his
making the type series and other Mexican Rhysodini available for study.

Clinidium ( Mexi clinidium) iviei new species
(Figs. 82, 84, 102)

Type Material — HOLOTYPE male, labelled: “MEX: Oaxaca, 2 mi. S. Cerro Pelon, 03 Jul 1982, 8-9000 ft. M.
A. Ivie Coll., ex rotten pine” (NMNH). PARATYPES two males, eight females, same data as holotype (NMNH); one
male, one female each (UCD, UVM, MAI). All forementioned paratypes with same data as holotype; one male, two
females (UVM); one male, one female (OSU); one male, one female (R. S. Miller Colin.); one male, one female sent to
Thomas Atkinson at the Colegio de Post-graduados, Chapingo, Mexico. All forementioned paratypes with same locality
data as holotype but labelled “July 2, 1982, (R. S. Miller Coll.)”; one female, labelled “Mexico, Oaxaca, 28 mi. N. Ixtlan
de J., 10,000', VII-23-29, 1971, A. Newton, under pine bark (MCZ).

Description. — Length 6. 2-8.0 mm. Head slightly longer than broad; median lobe long, tip posterior to hind
margin of eye; medial margins of temporal lobes slightly curved.

Pronotum relatively elongate, length/greatest width 1.54; lateral margins curved; apex less narrowed than in C.
mexicanum , anterior angles more distinct; base moderately narrowed; basal impressions larger than in C. mexicanum ,
about 0.3 of length of pronotum; basal impression closed posteriorly, open laterally; outer marginal groove completely
hidden in dorsal view, as deep as inner marginal groove; prosternum without precoxal setae.

All striae impressed, pollinose, including supramarginal, marginal; sutural stria with two or three setae near apex;
intercalary stria without setae; intratubercular stria with complete row of five or six setae; marginal stria with five or six
setae near apex; transverse sulci of abdomen broadly interrupted in midline, without pits at medial ends; Sternum VI of
abdomen shallowly impressed near apex in both sexes, in profile forming distinct angle; Sternum VI coarsely punctate,
most punctures elongate, some reaching margin of sternum (Fig. 102); lateral pit of Sternum IV of female, large glabrous;
anterior femur of male with large, broad, rather obtuse ventral tooth; surface proximal to tooth tuberculate; tooth without
setae; anterior trochanter of male dentate; anterior tibia of male with large proximal tooth (Fig. 84); middle calcar acute,
scarcely raised above level of spurs; hind calcar small, acute, elevated well above spurs; proximal margin of hind calcar
slightly emarginate.

This species is unique in having elongate, slightly confluent punctures on abdominal
Sternum VI. The pronotum is not as narrowed anteriorly as in C. mexicanum. The shape of the
pronotum, the well developed outer marginal groove which is hidden in dorsal view, and the
numerous elytral setae, all suggest C. blomi. The latter species, however, has four large
impressions on Sternum VI, rather than elongate punctures.

The species is named for the collector, Michael A. Ivie, to whom we are grateful for the
opportunity to study the type series.

SUBGENUS PROTAINOA BELL AND BELL 1978

Type species. — Clinidium ( Protainoa ) extrarium Bell and Bell 1978.

Description. — Antennal stylet slender, acuminate, long; tufts of minor setae present on Segments VI-X; one or
two temporal setae present; orbital groove abbreviated posteriorly; marginal groove of pronotum single; angular seta.

Quaest. Ent., 1985, 21 (1)

Bell and Bell

marginal setae absent; sternopleural groove absent; elytral striation reduced; marginal stria fifth complete stria from
suture; supramarginal stria represented by sparse row of fine punctures below intratubercular stria; sutural, parasutural,
intercalary, intratubercular striae complete; setae present in sutural, intercalary, intratubercular, marginal striae;
metasternum with broad median concavity, which is continued posteriorly to abdominal Sternum III; female with lateral
pit on abdominal Sternum IV; male with anterior trochanter dentate; all femora of male with ventral surface tuberculate;
tubercles most numerous on anterior femur; anterior femur of male with inconspicuous ventral tooth; male without
proximal tooth on anterior tibia; base of anterior tarsus distad to cleaning organ; calcars small; middle one longer but more
slender, more acute than hind one; latter triangular, its apex acute, slightly proximad of level of spurs; hind tibia of male
with medial apical tooth, resembling third spur.

Clinidium ( Protainoa ) extrarium Bell and Bell 1978
(Fig. 87)

Clinidium ( Protainoa ) extrarium Bell and Bell 1978: 63-64.

Type Material. — HOLOTYPE male, labelled: “Am. Bor. Rhysodes”, “N. Amerika” (BSL) PARATYPE
female, same data (BSL).

Description. — Length 6. 1-6.3 mm. Antenna with basal setae present on Segments VII-X; head slightly longer
than wide; median lobe short, tip acute, opposite anterior margin of eye; medial margins of temporal lobes slightly curved;
posterior margin of temporal lobe long, pilose; temporal lobe with two prominent, isolated punctures, probably both setose
(but setae probably broken off in type series); holotype with only one puncture on right side.

Pronotum elongate; length/greatest width 1.67; widest at basal 0.33; strongly tapered anteriorly; basal impressions
deep, oval, closed posteriorly; length of basal impression 0.16 of length of pronotum; hind angle with prominent tooth,
preceded by pollinose pit.

Sutural, parasutural, intercalary stria impressed, coarsely punctate, entire; intratubercular stria impressed near apex,
remainder not impressed, represented by row of fine punctures; marginal stria impressed, coarsely punctate near apex,
effaced in middle 0.33, represented by row of fine punctures near humerus; sutural stria with two setae near apex;
intercalary stria with complete row of four setae; intratubercular stria with two setae near apex; marginal stria with several
setae near apex; metasternum with large transverse curved pilose area near anterior margin; transverse sulci of Sterna
III- V prominent, rather narrowly interrupted at midline, medial, lateral ends each with prominent pits; in male, Sterna
III-V also each with median pit; female with median pit on Sternum III but not on other sterna; Sternum VI with
transverse sulci near base, curved submarginal sulci near apex; latter very narrowly interrupted at midline.

This isolated species resembles its nearest relatives in subgenus Tainoa in the shape of the
pronotum, but differs strongly in having much more complete elytral striation, denticulate hind
angles, and in lacking all pronotal setae.

The country of origin is unknown. We think it likely to be the tropical lowlands of Mexico or
northern Central America. It seems likely that the endemic subgenus Tainoa of the Greater
Antilles is derived from an ancestor much like C. extrarium.

SUBGENUS TAINOA BELL AND BELL 1978

Type species. — Clinidium darlingtoni Bell 1970.

Description. — Antennal stylet acuminate, long; tufts of minor setae present on antennal Segments VI-X; basal
setae present on Segments VI-X; two or three temporal setae; orbital groove present, abbreviated at posterior margin of
eye; marginal groove of pronotum single; angular, one or more marginal setae present; sternopleural groove absent; elytral
striation strongly reduced; parasutural stria effaced anteriorly, reduced to remnant in posterior part of elytron;
supramarginal not impressed, represented by row of punctures, incomplete; metasternum neither sulcate not impressed;
female with lateral pit in Sternum IV: Sternum VI with two pairs of impressions, both oblique, posterior pair divergent
posteriorly; in some specimens with additional pair of round anteriomedial pits; anterior femur with cleaning organ very
large, entirely proximad to base of anterior tarsus; anterior trochanter of male dentate; anterior femur of male without
ventral tooth.

Phytogeny. — The two Cuban species, C. curvicosta and C. chevrolati are obviously closely
related, differing mainly in secondary sexual characters. C. xenopodium, of Hispaniola, and C
darlingtoni, of Jamaica, are rather distantly related to one another, but share enough
characters, including a strongly abbreviated parasutural stria, to suggest that they are
descended from a common ancestor different from that which led to the Cuban species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

KEY TO SPECIES

1 Parasutural stria relatively long, anterior and near middle of elytron 2

V Parasutural stria very short, arising at or behind apical 0.33 of elytron 5

2 (1) Males, calcars present 3

2 ' Females, calcars absent 4

3 (2) Hind calcar with dorsal margin slightly sinuate, largest seta of hind calcar

smaller, scarcely longer than width of calcar

C. curvicosta Chevrolat, p. 7 1

3' Hind calcar with dorsal margin strongly angulate, largest seta more than
twice as long as width of calcar C. chevrolati Reitter, p. 74

4 (20 Sternum VI sloped gradually posteriorly, in profile view angulate

C. curvicosta Chevrolat, p. 7 1

4' Sternum VI strongly impressed posteriorly, anterior margin of impression
forming median tubercle; in lateral view, tubercle forming sharp angle
C. chevrolati Reitter, p. 74

5 (10 Parasutural stria not attached to intercalary anteriorly; intercalary ending

blindly posteriorly C. darlingtoni Bell, p. 74

5' Parasutural stria attached to intercalary stria both anteriorly and

posteriorly, isolating small remnant of Interval III

C. xenopodium Bell, p. 75

Clinidium ( Tainoa ) curvicosta Chevrolat 1873a
(Figs. 88, 93, 94)

Clinidium curvicosta Chevrolat 1873a: 215.

Clinidium ( Tainoa ) curvicosta (Chevrolat) Bell and Bell 1978.

Type Material. — Not seen by us. According to the original description, collected in Cuba by F. Poey. Vulcano
and Pereira (1975b) illustrated the elytron of a specimen in the Museum of Natural History in Vienna, which is labelled as
the type. As previously noted (Part 1:64), this specimen does not correspond to the original description, and probably is
labelled incorrectly. The original description could apply either to the present species, or to the one subsequently described
as C. chevrolati. Until an authentic type is located, it seems best to continue to use the Chevrolat name for the present
species, as was the practice of Bell (1970) and previous authors.

Description. — Length 4. 3-6. 2 mm (according to Chevrolat, the type measured 8 mm). Basal setae present on
antennal Segments VII-X; median lobe of head elongate, tip acute, opposite posterior margin of eye; medial margins of
temporal lobe nearly parallel opposite frontal space; medioposterior margin of temporal lobe nearly evenly rounded,
completely fringed with pilosity; temporal lobe with 3 setae in most specimens, two opposite eye, one near occiput; either
anterior or posterior of those near eye absent in some specimens.

Pronotum elongate, length/greatest width 1.74; basal impression about 0.15 of total length of pronotum; in most
specimens, basal impression pointed, suggesting rudimentary distal striole, four or five marginal setae.

Sutural stria impressed, entire, punctate; parasutural impressed, punctate, base near middle of elytron, not connected
to neighboring striae, apex connected to intercalary stria; intercalary stria entire, impressed, punctate; intratubercular
impressed near apex, otherwise not impressed, represented by row of coarse punctures; supramarginal effaced at base,
apex, middle 0.33 represented by row of punctures; marginal stria coarsely punctate, shallowly impressed; sutural stria
with four to six setae in nearly complete row, though absent from basal 0.25; intercalary stria with complete row of seven
or eight setae; intratubercular stria with three to five setae in apical 0.25; marginal stria with 10-12 setae forming nearly
complete row, though absent from basal 0.25 (Fig. 93); Sternum VI with reflected margin, anteriomedial pits present in
most specimens, absent in 1; anteriolateral pits elongate; posterior pits convergent posteriorly, connected by fine
submarginal groove (Fig. 94); anterior tibia of male with proximal tooth small, opposite proximal end of cleaning organ;
calcars very small; hind calcar triangular, dorsal margin nearly straight, largest seta scarcely longer than width of calcar;
lateral pit of Sternum IV of female laterad to sulcus; Sternum VI in female similar to that of male, only slightly concave in
lateral view; neither impressed nor tuberculate, female with tip of elytra evenly rounded in posterior view.

Quaest. Ent., 1985,21 (1)

Bell and Bell

Plate 8. Fig. 87. Genus Clinidium, Subgenus Protainoa, Head and pronotum, dorsal aspect, C. (P.) extrarium Bell and
Bell. Figs. 88-95. Genus Clinidium, Subgenus Tainoa ; Figs. 88-91, Head and pronotum, dorsal aspect; Fig. 88, C. (T.)
curvicosta Chevrolat; Fig. 89, C. (T.) chevrolati Reitter; Fig. 90, C. (T.) darlingtoni Bell; Fig. 91, C. (T.) xenopodium Bell;
Figs. 92-93, Left elytron, dorsal aspect; Fig. 92, C. ( T .) xenopodium Bell; Fig. 93, C. (T.) curvicosta Chevrolat; Figs.
94-95, Sterna III— VI, right half, female; Fig. 94, C. (T.) curvicosta Chevrolat; Fig. 95, C. (T.) chevrolati Reitter. Figs.
96-102. Genus Clinidium, Subgenus Mexiclinidium. Fig. 96-97, Sterna IV-VI, right half; Fig. 96, C. (M.) triplehorni
new species, female; Fig. 97, C. (M.) balli new species, female; Fig. 98, Metasternum, abdomen, right half, male C. (M.)
championi new species; Figs. 99, 101, Sterna V-VI; Fig. 99, C. (M.) halffteri new species; Fig. 101, C. [ M .) guatemalenum
Sharp; Figs. 100, 102, Sternum VI; Fig. 100, C. (M.) mexicanum Chevrolat; Fig. 102, C. (M.) iviei new species (bisected,
showing range of variation).

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent., 1985,21 (1)

Bell and Bell

The relatively long parasutural stria, 0.5 the length of the elytron, distinguishes this species
from all others except C. chevrolati. From the latter, the male can be distinguished by the
triangular hind calcar with a relatively short major seta, and the female by the unmodified
Sternum VI.

Range. — Cuba, central and eastern part of the Sierra Maestra. Bell (1970) lists localities.

Clinidium ( Tainoa ) chevrolati Reitter 1880
(Figs. 89, 95)

Clinidium chevrolati Reitter 1880: 30-31.

Clinidium turquinense Bell 1970.

Clinidium ( Tainoa ) chevrolati (Reitter) Bell and Bell 1978.

Type Material. — For C. chevrolati HOLOTYPE female, labelled: “Neu Granada, Chevr., type Cl., granatense,
chevrolati Reitter” (NMW). It is not clear why this specimen should be labelled as a type of C. granatense. It does not
match the description of the latter species, which is represented by another type, belonging to Subgenus Clinidium s. str.
(NMW). The type specimen of C. chevrolati undoubtedly bears an incorrect locality label, as it appears identical to C.
turquinense of Cuba. For C. turquinense HOLOTYPE male, labelled: “Pico Turquino, 3000-5000 ft., June, 1936, coll. P.
J. Darlington” (MCZ 31752). PARATYPE one female, with head, prothorax missing, same data as type (MCZ).

Description. — Length 6. 8-7.0 mm. Very similar to C. curvicosta except for the following points: body more
robust; pronotum less elongate, length/greatest width 1.60; five or six marginal setae; sutural stria with only two to four
setae limited to portion behind middle; hind calcar of male very large, strongly angulate on proximal margin, with very
long, curved seta; female with Sternum VI deeply impressed in posterior 0.5, impression preceded by median tubercle (Fig.
95); female with tips of elytra separately angulate in posterior view.

Range. — Pico Turquino is the western Sierra Maestra of Cuba, and outside the known
range of C. curvicosta.

Clinidium ( Tainoa ) darlingtoni Bell 1970
(Fig. 90)

Clinidium (s. str.) darlingtoni Bell 1970: 317-318.

Clinidium ( Tainoa ) darlingtoni (Bell) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Whitfield Hall, St. Thomas parish, JAMAICA, Jan. 9, 1967,
coll. R. T. Bell, J. R. Bell, B. B. Chiolino” (MCZ 31751). PARATYPES ten males. Five females, same data as holotype
(MCZ, UVM); two males, same locality, coll. P. J. Darlington, Aug. 13-20, 1934 (MCZ).

Description. — Length 4.9-6. 7 mm. Basal setae present on antennal Segments VIII-X; median lobe elongate, tip
acute, opposite posterior margin of eye; medial margins of temporal lobes parallel, rather close together; medioposterior
margin of temporal lobe nearly evenly rounded, fringed with pilosity; temporal lobe with three setae (preorbital,
postorbital, occipital).

Pronotum relatively short; length/greatest width 1.58; basal impression very small, about 0.10 of length of pronotum;
three or four marginal setae, one near angular seta, others anterior to middle of pronotum.

Sutural stria shallowly impressed, very coarsely punctate; parasutural stria deeply impressed, coarsely punctate, base
far posterior to middle of elytron, not connected to neighbouring striae; apex attached to intratubercular stria; intercalary
stria very deeply impressed, coarsely punctured, ending posteriorly just anterior to base of parasutural stria; interval
laterad to intercalary stria elevated, forming medial-facing scarp; intratubercular stria impressed near apex, otherwise
represented only by row of very minute punctures; supramarginal stria absent; marginal stria with basal 0.33 entirely
effaced, middle 0.33 represented by row of minute punctures; apical 0.33 impressed; sutural stria with three or four setae
near apex; intercalary stria with six to eight setae forming complete row; intratubercular stria with three or four setae near
apex; marginal stria with seven or eight setae in apical 0.25.

Sternum VI with reflected margin; anteriomedial pits absent; anteriolateral pits elongate; posterior pits convergent
posteriorly; anterior tibia of male with proximal tooth small, opposite proximal end of cleaning organ; calcars small, hind
one scarcely larger than middle one; hind calcar angulate to proximal margin, with small, proximally directed seta; lateral
pit of Sternum IV of female large, triangular, with short trace of transverse sulcus medial to it; Sternum VI of female
slightly concave in lateral view, but not distinctly impressed.

This species is easily recognized by the very short parasutural stria which ends blindly
anteriorly, close to the blind posterior end of the intercalary stria.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Range. — Jamaica from sea level to 4500'. Not known from west of Runaway Bay and Mt.
Diable. Bell (1970) gives a list of localities. In addition, we have studied a series of eight
specimens labelled: “JAMAICA, St. Andrew Parish, Hardwar Gap, 4000', J. Peck, 16, XII,
1972” (BSRI).

Clinidium ( Tainoa ) xenopodium Bell 1970
(Figs. 91, 92)

Clinidium ( sensu stricto) xenopodium Bell 1970: 316.

Clinidium ( Tainoa ) xenopodium (Bell) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Loma Vieja, near Santa Constanza, Dominican Republic, 6000
ft., August 1938, coll. P. J. Darlington” (MCZ 31750). PARATYPES two males, one female, labelled: “La Cavite,
Dominican Republic, March 5, 1917, coll. R. H. Beck” (MCZ, UVM).

Description. — Length 5. 8-6. 5 mm. Basal setae present on antennal Segments VII-X; median lobe short, tip acute,
opposite middle of eye; medial margins of temporal lobes divergent posteriorly; occipital angle glabrous, prominent,
interrupting fringe of pollinosity on margin of temporal lobe; temporal lobe with three setae, preorbital, postorbital,
occipital.

Pronotum moderately elongate, length/greatest width 1.65; basal impression oval, about 0.25 of length of pronotum;
one marginal seta near apex of pronotum, also one angular seta.

Sutural stria impressed, finely punctate; parasutural stria impressed, short, joined at both ends to intercalary stria,
isolating small oval area; intercalary stria entire, impressed, finely punctate; intratubercular stria impressed near apex,
otherwise represented by row of minute punctures; supramarginal stria effaced at base, apex, represented by row of fine
punctures in middle 0.33 of elytron; marginal stria deeply impressed at apex, remainder shallow impressed; sutural stria
with two or three setae near apex, or else these setae on Interval I, medial to sutural stria; intercalary stria with four setae
near apex; intratubercular stria with two setae near apex; marginal stria with seven or eight setae in posterior 0.5 (Fig. 92).

Sternum VI of abdomen without raised rim; anteriomedial pits absent; anteriolateral pits elongate; posterior pits
parallel or divergent posteriorly; anterior tibia of male with proximal tooth large, distinctly proximad to cleaning organ;
calcars very large; middle calcar narrowly triangular, acute, 0.33 as long as tibia; hind calcar broadly triangular, apex
acuminate, calcar more than 0.5 as long as tibia; female with lateral pit of Sternum IV very large, rounded medially,
without trace of transverse sulcus medial to it; Sternum VI of female not impressed posteriorly.

The short parasutural stria, connected both anteriorly and posteriorly to the intercalary
stria, is diagnostic of this species.

SUBGENUS ARCTOCLINIDIUM BELL 1970

Type species. — Clinidium sculptile (Newman)

Description. — Antennal stylet small; tufts of minor setae present on Antennal Segments VI-X (C. veneficum) or
VII-X (all other species); temporal seta one or absent; eye crescentic; orbital groove complete, joined posteriorly to
marginal pollinosity of temporal lobe; pronotum with lateral margins curved, base, apex truncate; marginal groove double
or single; pronotum with angular seta present (C. marginicolle) or absent (all other species); marginal setae absent;
sternopleural groove present or absent; elytral striation complete; marginal stria sixth from suture; inner intervals of
elytron convex or costate; elytral setae very few, at most one in apex of parasutural or sutural stria, several in apex of
marginal stria, one on apical tubercle; metasternum with or without median sulcus; female with enlarged lateral pit on
Sternum III or IV or both; base of anterior tarsus opposite cleaning organ.

The more distal position of the cleaning organ and the relatively few elytral setae separate
this subgenus from Mexiclinidium. North American species have the inner intervals costate,
and look similar to the larger Mexiclinidium , from which they can be distinguished by the
presence of the sternopleural groove and the complete intercalary stria. European and western
Asian species have the intervals not costate, and lack the sternopleural grooves.

This subgenus is Holarctic. It has five species in the eastern U.S.A., and one each on the
Pacific Coast of North America, Japan, the Caucasus, and Southern Europe.

Phytogeny. — Our concept of the interrelationship of the nine species are as illustrated in
Diagram 2. The subgenus is derived from two ancestral populations, Species 2 and Species 3,

Quaest. Ent., 1985,21 (1)

canalic u latum

76 Bell and Bell

Phylogenetic Diagram 2. Reconstructed Phylogeny of species of Clinidum Subgenus Arctoclinidium

rosenbergi

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

both descended from the common ancestor, Species 1. From Species 2 have arisen the two
European species, C. canaliculatum and C. marginicolle. They are characterized by the
following features: elytral intervals not carinate, though convex; sternopleural groove absent;
precoxal setae present; parasutural stria incomplete at apex, so Intervals II, III fuse posteriorly;
supramarginal row of punctures, not impressed; marginal stria not impressed except at extreme
apex; one seta near apex of either sutural or parasutural stria; abdominal sterna in both sexes
glabrous medially.

Species 3 was the ancestor of the six North American and one Japanese species. Among
these species, the inner elytral intervals are carinate, the sternopleural groove is present;
precoxal setae are absent; parasutural stria is complete, so Intervals II, III do not fuse; the
supramarginal stria is impressed; marginal is completely impressed; setae are absent from
apices of inner elytral striae, and the abdominal sterna of the male have a pattern of pollinose
areas.

Synapomorphies of Species 2 and its descendants include loss of sternopleural groove; loss of
apex of parasutural stria; possible synapomorphies are the reduction of marginal and
supramarginal striae. Synapomorphies of Species 3 and its descendants are the development of
ventral pollinosity in the male, the highly costate elytral intervals, and possibly the loss of setae
from the apex of the sutural or parasutural striae.

The interpretation of the costate intervals is debatable. Noncostate, slightly convex intervals
are found in most Rhysodini with functional hind wings, and probably represent the primitive
character state for the tribe. Costate intervals have arisen several times, and can be regarded as
advanced. However, it is quite possible that the costae can be reduced secondarily. Within
Clinidium , both Mexiclinidium and Arctoclinidium have costate and noncostate species, and in
the more highly modified members of subgenera Tainoa and Clinidium sensu stricto, the
intervals are not costate. Thus, Species 2 may have lost its costae secondarily. The absence of
the specialized pollinose areas on the abdomen of the male, however, suggests that Species 2
and its descendants are a separate phyletic line from the costate species.

Species 2 probably had the intercalary stria unmodified. The status of other characters is
less definite. C. marginicolle is the only member of the subgenus to have an angular seta. It
could be argued that this seta has been lost separately in C. canaliculatum , and in Species 3.
However, in Rhysodini generally the greatest number of tactile setae are found in highly
modified species with cryptic habits and strongly reduced eyes. This might imply that a
proliferation of tactile setae has happened independently, and that the angular seta has
appeared de novo in C. marginicolle. There are similar possibilities in relation to the presence
of a temporal seta in both species descended from Species 2, and in C. valentinei alone among
those descended from Species 3. Either C. valentinei retains the temporal seta, which was lost
independently in C. veneficum , C. calcaratum and Species 5, or else the temporal seta was
developed independently in C. valentinei and in Species 2.

Species 3 probably gave rise to Species 4 and 5. In Species 5, the metasternum developed a
longitudinal sulcus. The four species descended from it are much alike, and all are found in
eastern North America. The existence of Species 4 is less strongly indicated than that of
Species 5. There are no clearly derived characters in common among C. veneficum , C.
valentinei , and C. calcaratum , though a possible synapomorphy is the fact that the pollinose
area of Sternum II is narrowed anteriorly, while in the remaining species, it is as wide or wider
anteriorly than posteriorly. Otherwise, the three species without a sulcus are more widely
divergent from one another than are those with a sulcus, and are widely distributed, with one

Quaest. Ent., 1985,21 (1)

Bell and Bell

species each in Japan, the North American Pacific Coast, and the Appalachians.

We hypothesize the descent of C. valentinei and C. veneficum from Species 6. Common
features include a ventral tooth on the anterior femur of the male, a relatively broad marginal
carina on the pronotum, and a distinct cauda on the elytron in the female. In C. calcaratum, in
contrast, the ventral tooth is lacking, the marginal carina is linear, and the cauda is small but
distinct in both sexes. In all of these characters, there is uncertainty about which character
state is plesiomorphic, and alternative arguments could be made to support other phyletic
arrangements. Thus, C. calcaratum and C. valentinei both have the anterior trochanter of the
male pointed, while it is rounded in C. veneficum. C. calcaratum and C. veneficum lack a
temporal seta and a ventral tooth on the hind femur, while they have the lateral pits of Sternum
IV enlarged. In C. valentinei a temporal seta and a tooth on the hind femur of the male are
present, and the lateral pits of Sternum III are enlarged.

Both the posterior and anterior trochanters of the male are pointed in C. calcaratum , a
feature it shares with C. canaliculatum and C. marginicolle. This suggests that pointed
trochanters were a feature of the common ancestor of the subgenus, Species 1 . C. valentinei is
unique in having the anterior trochanter pointed, but the posterior one rounded, an
intermediate condition, between the ancestral character state, and the condition seen in C.
veneficum and all four descendants of Species 5, which have both anterior and posterior
trochanters rounded.

C. valentinei has the lateral pit on Sternum III enlarged, while C. calcaratum and C.
veneficum have that of Sternum IV enlarged. The latter character state is probably the
primitive one. Sternum IV has enlarged pits in C. canaliculatum and C. marginicolle , and in
the overwhelming majority of Rhysodini in other subtribes. It seems likely that a shift to
Sternum III is a specialization that has occurred in C. valentinei independently of the species
descended from Species 5. Of the latter, the shift is only partial in Species 8 and its
descendants, C. rosenbergi and C. sculptile which have both III and IV enlarged, but is
complete in the remaining species, C. apertum and C. baldufi, which resemble C. valentinei in
having only the pits of Sternum III enlarged.

Species 5, with the metasternum sulcate, gave rise to two descendants, Species 7 and 8.
Species 7 had enlarged lateral pits only on Sternum III in the female, and had the prosternum
glabrous in the male. It gave rise to C. apertum and C. baldufi. Species 8 developed a pollinose
area on the prosternum of the male, a feature not seen elsewhere in the genus, while the lateral
pits of the female were enlarged on both Sterna III and IV. It gave rise to C. rosenbergi and C.
sculptile.

KEY TO SPECIES

1 Parasutural stria abbreviated near apex, Intervals II, III fused posteriorly;

elytral intervals merely convex 2

1' Parasutural stria complete, Intervals II, III not fused posteriorly; elytral

intervals costate 3

2 (1) Angular seta absent from pronotum; intercalary stria only slightly broader,
deeper than parasutural; anterior femur of male without ventral tooth . .

C. canaliculatum (Costa), p. 83

2' Angular seta present; intercalary stria twice as broad and deep as
parasutural stria, strongly dilated in apical 0.33; anterior femur of male

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

with ventral tooth C. marginicolle Reitter, p. 83

3 (10 Metasternum without median sulcus

3 Metasternum with median sulcus

4 (3) Males, calcars present

4' Females, calcars absent

5 (4) Anterior femur without ventral tooth; hind trochanter pointed

C. calcaratum Leconte, p. 84

5' Anterior femur with ventral tooth; hind trochanter rounded

6 (5') Posterior femur with ventral tooth; anterior trochanter pointed

C. valentinei Bell, p. 85

6' Posterior femur without ventral tooth; anterior trochanter rounded

C. veneficum Lewis, p. 88

7 (40 Lateral pit of abdominal Sternum III enlarged

C. valentinei Bell, p. 85

7' Lateral pit of Sternum IV enlarged

8 (70 Marginal carina of pronotum linear; outer marginal groove not visible in

dorsal view; basal impression of pronotum open posteriorly

C. calcaratum Leconte, p. 84

8' Marginal carina relatively broad; outer marginal groove visible in dorsal
view; basal impression closed posteriorly C. veneficum Lewis, p. 88

9 (30 Males, calcars present

9' Females, calcars absent

10 Prosternum glabrous

10' Prosternum with median pollinose area

11 (10) Abdominal Sternum V with pollinosity interrupted by narrow median

carina; anterior tibia without proximal tooth C. baldufi Bell, p. 89

1 V Abdominal Sternum V without median carina; anterior tibia with proximal
tooth C. apertum Reitter, p. 90

12 (100 Pollinose area present between transverse sulci on Sternum V; hind calcar

large C. sculptile (Newman), p. 92

12' Sternum V glabrous between transverse sulci; hind calcar very small

C. rosenbergi Bell, p. 91

13 (90 Lateral pit in Sternum III of abdomen

13' Lateral pits present in Sterna III, IV

14 (13) Sternum VI with posterior 0.5 deeply impressed; cauda of elytra

prominent, trapezoidal C. baldufi Bell, p. 89

14' Sternum VI with posterior 0.5 not impressed; cauda small, rounded

C. apertum Reitter, p. 90

15 (130 Lateral pits of Sterna III, IV equally large; cauda very small, rounded;

Sternum VI impressed in some specimens, not impressed in others

C. rosenbergi Bell, p. 91

15' Lateral pits of Sternum IV smaller than those of Sternum III; cauda
prominent, rounded; Sternum VI with posterior 0.5 deeply impressed
C. sculptile (Newman), p. 92

. 4

. 5

. 7

. 6

. 8

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Plate 9. Figs. 103-1 11. Genus Clinidium, Subgenus Arctoclinidium. Figs. 103-109, Head and pronotum, dorsal aspect;
Fig. 103, C. (A.) rosenbergi Bell; Fig. 104, C. (A.) baldufi Bell; Fig. 105, C. (A.) calcaratum Leconte; Fig. 106, C. (A.)
canaliculatum (Costa); Fig. 107, C. (A.) marginicolle Reitter; Fig. 108, C. (A.) valentinei Bell; Fig. 109, C. (A.) veneficum
Lewis; Fig. 1 10, Elytra, dorsal aspect (showing strial variation), C. (A.) veneficum Lewis; Fig. Ill, Sternum VI, C. (A.)
veneficum Lewis.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Plate 10. Figs. 112-123. Genus Clinidium, Subgenus Arctoclinidium Metasternum, abdomen, halved. Figs. 112, 1 18, C.
(A.) rosenbergi Bell; Figs. 1 13, 1 19, C. (A.) baldufi Bell; Figs. 1 14, 120, C. (A.) apertum apertum Reitter; Figs. 115, 121,
C. (A.) sculptile (Newman); Figs. 116, 122, C. (A.) valentinei Bell; Figs. 117, 123, C. (A.) calcaratum Leconte.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Clinidium ( Arctoclinidium ) canaliculatum (Costa 1839)
(Fig. 106)

Ips canaliculatus Costa 1839: 104.

Rhysodes trisulcatus Germar 1840: 441.

Rhysodes sulcipennis Mulsant 1853: 6.

Clinidium trisulcatum (Germar) Chevrolat 1873a.

Clinidium canaliculatum (Costa) Grouvelle 1903.

Clinidium ( Arctoclinidium ) canaliculatum (Costa) Bell and Bell 1978.

Type Material. — We have not located type material for any of the three nominate species of Clinidium
described from southern Europe. As there appears to be only one species of Clinidium in Europe, we follow Grouvelle
(1903) in regarding R. trisulcatus Germar and R. sulcipennis Mulsant as being pure synonyms.

Description. — Length 6.0-7. 5 mm. Antennal Segments VII-X with tufts of minor setae; basal setae sparse,
limited to Segments IX-X; one temporal seta present; pronotum rather short, length/greatest width about 1.47; greatest
width of pronotum slightly behind middle; pronotum strongly narrowed anteriorly, only slightly narrowed posteriorly;
angular seta absent; basal impressions open posteriorly; outer marginal groove not visible in dorsal view, in most specimens
abbreviated anteriorly; precoxal setae present on prosternum; sternopleural groove absent.

Elytral striae punctate; sutural stria complete, shallowly impressed; parasutural impressed, abbreviated near apex;
intercalary stria much deeper than other striae; portion between 0.55 and 0.70 of length deeper but scarcely wider than
remainder; intratubercular stria shallowly impressed; supramarginal stria not impressed, represented by line of very fine
punctures near middle of elytron; base, apex effaced; marginal stria effaced at base, middle portion not impressed,
represented by line of rather coarse punctures; apex impressed; Intervals I-III broad, nearly flat, of nearly uniform width;
Interval IV not flattened laterad to deepened part of intercalary stria; elytron with one seta in apex of sutural stria, absent
or one in apex of intercalary stria; two to four near apex of marginal stria.

Metasternum not sulcate; transverse sulci of abdominal sterna widely separated in midline, without median pollinosity
in either sex; Sternum VI coarsely punctate; female with lateral pits on abdominal Sternum IV; Sternum VI of female
without impression; male with anterior, posterior trochanters pointed; male without ventral tooth on anterior femur,
without proximal tooth on anterior tibia; middle calcar acute, narrow; hind calcar large, somewhat cultrate, dorsal margin
curved, ventral margin shallowly emarginate; male with prosternum, abdominal sterna glabrous.

This species is similar to C. marginicolle, from which it differs in the absence of the angular seta, in the wider, flatter
elytral intervals, and in the smaller, narrower subapical enlargement of the intercalary stria. The males can also be
distinguished by the different shape of the hind calcar, and by the absence of the ventral tooth on the anterior femur in C.
canaliculatum. Some specimens of C. canaliculatum resemble C. marginicolle in having a complete outer marginal groove
on the pronotum, although it is not visible except in lateral view.

Range. — Southern Italy (Sicily, Calabria), Greece. We have studied specimens with

Specific locality labels: GREECE: Taygetos, Morea (CNHM, UVM). ITALY: Aspromonte, Calabria (CNHM);
Sta. Eufemia, Calabria (CNHM, UVM). Hincks (1950) and earlier authors also record it from the Caucasus. We have
not seen this species from the Caucasus, and believe that records from there are misidentified C. marginicolle.

Clinidium ( Arctoclinidium ) marginicolle Reitter 1889
(Fig. 107)

Clinidium marginicolle Reitter 1889: 23.

Clinidium ( Arctoclinidium ) marginicolle (Reitter) Bell and Bell 1978.

Type Material.- — We have not located authentic type material. According to the original description, the type
series was from Lenkoran (Azerbaijan), and was collected by Leder. A specimen in the Natural History Museum of
Vienna is labelled as a type, but is labelled: “PERSIA: Kopet Dagh, Siaret 1160 m., 5.99, Col. Hauser”. The label
probably indicates that Reitter compared it with his type series and considered it identical.

Description. — Length 5. 8-7. 5 mm. Antennal Segments VII-X with tufts of minor setae; basal setae few, limited to
Segment X; one temporal seta; pronotum more elongate than in C. canaliculatum, length/greatest width about 1.57;
widest near middle; base more narrowed than in C. canaliculatum, nearly as narrow as apex; angular setae present; basal
impressions closed posteriorly or almost closed; outer marginal groove deep, complete, barely visible in dorsal view;
prosternum with precoxal setae; sternopleural groove absent.

Elytral striae punctate; sutural stria complete, shallowly impressed; parasutural impressed, abbreviated near apex,
sinuate opposite dilated portion of intercalary stria; latter very deeply impressed; portion between 0.55, 0.70 of length
deeper, wider than remainder; intratubercular stria shallowly impressed; supramarginal stria almost absent, represented by
a few minute punctures near middle; marginal stria effaced at apex, marginal stria effaced at base, middle portion not
impressed, represented by row of punctures; apex impressed; sutural interval of even width, nearly flat; Interval II similar

Quaest. Ent., 1985,21 (1)

Bell and Bell

to sutural interval in anterior 0.5, posteriorly much narrower than sutural interval, convex, slightly sinuate opposite
depressed part of intercalary stria; Interval III slightly convex, as broad as sutural interval, depressed opposite depressed
part of intercalary stria; Interval IV broad, slightly convex except opposite depressed part of intercalary stria, where
narrowed, depressed; elytron without setae or one in apex of sutural stria, one seta in intercalary stria opposite apex of
parasutural stria, one near apex of intratubercular stria, two to four apex of marginal stria.

Metasternum not sulcate, transverse sulci of abdominal sterna rather narrowly separated in midline, without median
pollinosity in either sex; Sternum VI coarsely punctate; female with lateral pits on abdominal Sternum IV; Sternum VI of
female without impression; male with anterior, posterior trochanters pointed; male with small, obtuse ventral tooth on
anterior femur; male without proximal tooth on anterior tibia; middle calcar small, acute, triangular; hind calcar small,
with dorsal margin angulate, ventral margin straight, well above spurs; male with prosternum, abdominal sterna glabrous.

This species differs from C. canaliculatum in having an angular seta, in having a longer,
narrower pronotum, and in the shape of the calcars. Also, the subapical enlargement of the
intercalary stria is more conspicuous, and alters the neighbouring intervals and striae.

Range. — Caucasus Mountains of the southern U.S.S.R. and the Kopet Dagh range of

northeastern Iran. We have studied specimens from the following localities: IRAN, Siaret, Kopet
Dagh (NMW; NMHB, CNHM); Astrabad (MNHB); U.S.S.R. Paleton, 1200', Astar. R„ Talysh. (LEN).

Clinidium ( Arctoclinidium ) calcaratum LeConte 1875
(Figs. 105, 117, 123)

Clinidium calcaratum LeConte 1875: 164.

Clinidium ( Arctoclinidium ) calcaratum (LeConte) Bell 1970.

Type Material. — HOLOTYPE sex not recorded, labelled: “Vane.” (MCZ 6831). In the original description the
type locality is cited as “Vancouver Island”.

Description. — Length 5.8-8. 1 mm. Antennal Segments VII-X with tufts of minor setae; basal setae sparse,
limited to Segments IX, X; temporal seta absent; pronotum relatively elongate, length/greatest width about 1.59; widest
posteriorly, base slightly narrowed, apex strongly so; sides of pronotum only slightly curved; angular seta absent; basal
impressions relatively large, open posteriorly; outer marginal groove scarcely visible in dorsal view, close to inner marginal
groove, separated by narrow nearly linear marginal carina; precoxal setae absent; sternopleural groove present.

Elytral striae deep, broad, inconspicuously punctate; supramarginal stria shallower than the others; intervals narrow,
costate; intercalary not abbreviated posteriorly in most specimens, in a few specimens very shortly abbreviated on one
elytron; Intervals II, III not united posteriorly except as a unilateral aberration; elytron entirely without setae; cauda
small, rounded, present in both sexes.

Prosternum glabrous in both sexes; metasternum not sulcate; transverse sulci widely separated at middle; male with
pollinosity of Sternum II a narrow rectangle; male with median pollinose areas present on Sterna I-IV, absent from
Sternum V; Sternum VI varying geographically (see below, under variation)', female with lateral pits bn Sternum IV;
anterior, posterior trochanters of male pointed; male without ventral tooth on anterior, posterior femora, without proximal
tooth on anterior tibia; hind calcar very large, 0.5 as long as hind tibia.

This is the only member of the genus in western North America. The male is easily recognized by the very large hind
calcar, while the female differs from all species with carinate intervals other than C. veneficum in having the lateral pits in
abdominal Sternum IV but not Sternum III. The narrow marginal carina and the absence of a tuft of minor setae on
antennal Segment VI will separate it from the latter species.

Range. — Substantially as listed by Bell (1970), in the Coast Range and Sierra Nevada of
California, from Mendocino and Tuolumne Counties northwards. In Oregon, known from a
number of localities in Klamath and Jackson Counties, near the California state line, and from
a few spots near the Columbia River, both in the Coast and Cascade Ranges, but not known
from the remainder of Oregon. In Washington, known from the Puget Sound area, including
the Olympic Peninsula, the San Juan Islands, and the lowlands east of the Sound; in British
Columbia, known from southern half of Vancouver Island, north to Comox, and also from the
mainland. The record from Mt. Garibaldi, collected by Virginia Anderson, is significantly
north of previously recorded mainland localities.

The following locality records are in addition to those published by Bell (1970): BRITISH
COLUMBIA: Goldfield (CMP); Goldstream (UK); Mount Garibaldi (UVM); Vancouver (CAS). CALIFORNIA:
Denny (Bell Creek), 2500 ft. (LA); Georgetown (CAG); Happy Camp (Siskiyou Co.) (CAG); Lake Alamanor (Plumas
Co.) (CAG); Maple Creek (Humboldt Co.) (CAG); Placer Co. (PU); Siskiyou Co. (MO; CNHM; BMS); Uncle Toms

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

(CAG); Yuba Pass (Sierra Co.) (OS; LA). OREGON: Ashland Peak, Siskiyou Mts., 7000 ft. (CNHM); Beaver-Sulfur
(Jackson Co.) (OS); Forest Grove (MN); Huckleberry Mts. (Jackson Co.), 5500 ft. (CNHM); Josephine Co. (CNHM);
Merlin (Josephine Co.) (OS); Oregon Caves (Jackson Co.) (CNHM).

Variation. — In most specimens from Puget Sound and vicinity, Sternum VI is impunctate
in both sexes, and the submarginal groove is widely interrupted in the midline. In a few from
Puget Sound, the base of Sternum VI has a few punctures. In a majority of northern females,
Sternum VI is slightly impressed posteriorly. In this population the pronotum is shorter and
more oblong than is usual for California specimens. Most California specimens have numerous
punctures of Sternum VI, and the submarginal groove is scarcely interrupted. The female lacks
an impression on Sternum VI, and the pronotum in both sexes is more slender and elongate.

These differences might indicate separate subspecies. However, it is also possible that the
variation has a clinal nature. The absence of specimens from central Oregon makes it
impossible to decide at present.

Bionomics. — Recorded many times from douglas-fir ( Pseudotsuga ) logs (Bell 1970).

Clinidium ( Arctoclinidium ) valentinei Bell 1970
(Figs. 108, 116, 122)

Clinidium ( Arctoclinidium ) valentinei Bell 1970: 313.

Type Material. — HOLOTYPE male, labelled: “Gorgas, Walker Co., Ala., April 17, 1949, coll. Barry
Valentine” (OSU). PARATYPES one male, four females, collected with type (OSU; UVM).

Description. — Length 5. 4-6. 4 mm. Antennal Segments VII-X with tufts of minor setae; basal setae present on
Segments VII-X; temporal lobe present; pronotum relatively short, length/greatest width 1.42; widest near middle; base,
apex only slightly narrowed; lateral margins feebly curved; angular seta absent; basal impression large, 0.33 as long as
prontum, open posteriorly; outer marginal groove clearly visible in dorsal view, separated from inner groove by broad
marginal carina; precoxal setae absent; sternopleural groove present.

Elytral striae broad, deep, scarcely punctate; supramarginal stria impressed in most specimens, reduced to row of
coarse punctures in smallest specimens; intervals narrow, costate; intercalary stria not abbreviated posteriorly; Intervals II,
III not united posteriorly; marginal stria without setae or one or two near apex; cauda of female small, rounded.

Prosternum glabrous in both sexes; metasternum not sulcate; transverse sulci of abdominal sterna narrowly separated
in midline; male with median pollinose areas on Sterna I-IV, that of II narrowed anteriorly, its lateral margins straight or
concave; Sternum V without pollinosity; Sternum VI of female with posterior 0.5 deeply impressed, impression bounded
anteriorly by straight transverse scarp; disc with a few punctures anterior to scarp; female with large lateral pits on
Sternum III; anterior trochanter of male pointed, posterior one rounded; anterior, posterior femora of male each with
prominent ventral tooth; anterior tibia of male with large proximal tooth; hind calcar acute, smaller than in C. calcaratum,
0.40 as long as tibia.

This is the only member of the genus in the eastern U.S. to lack the median sulcus of the
metasternum. The species can also be recognized by the presence of a temporal seta, though the
latter is very small and often hard to see. The male is unique in the subgenus in having a ventral
tooth on the posterior femur. The female resembles C. baldufi and C. apertum in having lateral
pits on Sternum III but not Sternum IV. The rounded cauda separates it from C. baldufi, and
the impression of Sternum IV from C. apertum.

Blanchard (1889) recognized this species as “Form B” of C. sculptile.

Range. — An Appalachian species, known from three widely scattered regions; north central
Alabama; the mountains of eastern Tennessee, northeastern Georgia, and western North and
South Carolina, and southwestern Pennsylvania, near Pittsburgh. It is not clear whether the
range is really broken into relict areas or whether these merely record infrequent collection.

In addition to the localities listed by Bell (1970), we have seen specimens from the following

localities: NORTH CAROLINA: Highlands (BSRI); PENNSYLVANIA: Jeanette (CMP), Wall (CMP; UVM);
TENNESSEE: Chimneys C. Gr., Gt. Smoky Mt. Nat. Pk. 2800 ft. (CU), Gregory Bald, Gt. Smoky Nat. Pt. (CU);
SOUTH CAROLINA: Clemson (UVM).

Quaest. Ent., 1985,21 (1)

Bell and Bell

TABLE 1

POLYMORPHISM IN Clinidium veneficum LEWIS

Locality Sex Sternum VI Stria III

Miyanoshita

Riga

“Japan”

Nagasuki

Chiuzenji

Oyayama

Higo

Hakone

Mt. Kohtsu

Mitsugi

(continued on next page)

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Table 1 (continued)

Locality

Sex

Sternum VI

Stria III

Mitsugi

“Japan”

Explanation of abbreviations:

Sternum VI - “b” indicates sternum tuberculate on both sides; “r” indicates tuberculate on
right side only “1” indicates tuberculate on left side only “a” indicates tubercules absent.
Stria III = “t” indicates intercalary stria of both elytra contain tubercles; “u” indicates
tubercle present on one side only; “s” indicates stria simple, without tubercle.

TABLE 2

COMBINATIONS OF POLYMORPHIC CHARACTERS CLINIDIUM VENEFICUM

LEWIS

Simple Stria

Double Stria
Unilateral

Bilateral

Sternum VI

bituberculate

1 m, 1 f

1 f

5 f

right tubercle

1 m

left tubercle

2 f

no tubercles

9 m, 2 f

1 f

m = male, f = female

TABLE 3

PROPORTION OF POPULATION WITH EACH CHARACTER STATE

Sternum VI

males

females

both sexes

bituberculate

58%

34%

right tubercle

left tubercle

17%

no tubercles

82%

25%

53%

II.

Intercalary Stria
simple

100%

45.5%

70%

unilateral double stria

bilateral double stria

45.5%

26%

Quaest. Ent., 1985, 21 (1)

Bell and Bell

A specimen labelled (CANADA) “North West Territories, N. Karts Camp, S. Nahanni
River, 61° 34' N, 124° 2 ' W, 28-VI-19-VIII, 1972, S. Peck, R. Syme” (BSR1) must surely be
mislabelled.

Clinidium ( Arctoclinidium ) veneficum Lewis 1888
(Figs. 109, 110, 111)

Clinidium veneficum Lewis 1888: 83.

Clinidium ( Arctoclinidium ) veneficum (Lewis) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) male, labelled: “Japan, Miyanoshita, 20-XII-23-XII, 80, G.
Lewis 1910-320, Clinidium veneficum Lewis” (BMNH). PARALECTOTYPES four females, one male, same data as
lectotype (BMNH); two females, labelled: “Higo, Japan, G. Lewis” (BMNH).

Description. — Length 5. 1-7.2 mm. Antennal Segments VI-X with tufts of minor setae; basal setae sparse, limited
to Segments IX, X; temporal seta absent; pronotum varied in proportions, length/greatest width 1.15 to 1.55; widest near
middle; base, apex nearly equally narrowed; lateral margins curved; angular seta absent; basal impressions relatively small,
length about 0.25 of length of prontoum, oblique, closed posteriorly; outer marginal groove prominent in dorsal view; inner
marginal groove relatively distant from it; marginal carina broad at middle, narrowed to base, apex; precoxal setae absent;
sternopleural groove present.

Elytral striae broad, deep, inconspicuously punctate; elytral intervals costate; intercalary stria complete; in female,
dimorphic, either doubled for short distance behind middle, isolating small tubercle which looks like remnant of additional
interval, or else entire (Fig. 110); intercalary stria of male entire; supramarginal stria scarcely impressed, represented by
row of punctures, abbreviated posteriorly; marginal stria impressed near apex, more anteriorly represented by row of
punctures; apex of marginal stria with one or two setae, or else these setae on apical tubercle just above marginal stria ;
cauda dimorphic, trapezoidal in some females, rounded in others.

Prosternum glabrous in both sexes; metasternum not sulcate; transverse sulci of abdominal sterna narrowly interrupted
in midline; that of Sternum II not interrupted in some females; Sterna II-V in male with median pollinose area, that of
Sternum II trapezoidal; Sternum VI impressed in posterior 0.5, polymorphic, either with large tubercle near lateral margin
anterior to impression on either side, or with only one such tubercle on right side, or on left side, or entirely without
tubercles; female with deep lateral pits on Sternum IV; all trochanters of male rounded; anterior femur of male with small
ventral tooth; hind calcar acute, narrow, rather long, about 0.33 of length of tibia, not raised above level of spurs.

This is the only member of the species known from Japan. It is similar to C. calcaratum of
western North America, but has a broader pronotum with broader marginal carinae, and has
the basal impressions closed posteriorly. It is the only member of the subgenus to have a tuft of
minor setae on Antennal Segment VI.

Range. — Japan, islands of Kyushu and Honshu. On the latter island, north to Nikko. In

addition to type material, we have studied the following specimens: one female, labelled: “Chiuzenju,
19, VIII-24- VIII 81, Japan, G. Lewis, 1910-330” (BMNH); one female, labelled: “Hakone, Japan, Sharp Coll. 1905-313”
(BMNH); one male, labelled: “Kiga, Japan, G. Lewis, 1910-320” (BMNH); one male, labelled: “Mie Univ. Forest.
Ichishi-Gun Mie, 24-VM956, Coll. M. Sato” (SATO); four males, two females, labelled: “Mitsugi Mura, Mie Prf.,
ll-VI-1956, coll. Z. Naruso” (SATO); one female, labelled: “Mt. Kohtsu (Tokushima), 31-X-1965, M. Sakai leg.”
(SATO); one male, labelled: “Nagasaki, Japan, G. Lewis, 1910-320, 22-V-3-VI, 81” (BMNH); one female, labelled:
“Oyayama, 26-4-81, Japan, G. Lewis, 1910-320” (BMNH).

Variation. — This species exhibits remarkable polymorphism, and deserves detailed study.
The intercalary stria is either entire or is divided for a short distance behind the middle,
isolating a small tubercle. In all males it is entire, while the females are roughly 0.5 entire and
0.5 divided. Sternum VI is polymorphic. Many specimens have a pair of large tubercles (Fig.
1 1 1), a few have only the left tubercle, one has only the right tubercle, and many lack tubercles
entirely. These forms are not secondary sexual characters, though the relative numbers of each
morph are very different in the two sexes. Most males lack tubercles, while over half the
females have both tubercles. The elytral cauda of the female also seems to vary in shape from a
round to trapezoidal form, but detailed studies have not been completed.

The combination of characters seen in the 23 specimens which we studied are indicated in
Tables 1, 2, 3. Despite the diverse appearance of the individuals, it seems likely there is one

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

highly polymorphic species. The lectotype is a male with the intercalary stria entire, and
Sternum VI with the right tubercle, but without the left one. The other male from
Miyanoshima has both tubercles on Sternum VI. Thus neither agree with the majority of
males, which lack both tubercles.

Bionomics. — According to Lewis (1888) this species occurs in old beech trees.

Clinidium ( Arctoclinidium ) baldufi Bell 1970
(Figs. 104, 113, 119)

Clinidium ( Arctoclinidium ) baldufi Bell 1970: 313-314.

Type Material. — HOLOTYPE male, labelled: “Dayton, 111., May 17, 1946, coll. F. G. Werner” (MCZ 31748).
PARATYPES one male, one female, same label as holotype (UVM); two males, labelled: “Lasalle Co., 111.” (MCZ); one
male, labelled: “Putnam Co., 111.” (UI); two males, labelled: “Starved Rock State Park, 111.” (ISNHS).

Description. — Length 5. 9-7.4 mm. Antennal Segments VII-X with tufts of minor setae; Segment VIII with basal
setae few to absent; Segments IX-X with basal setae; temporal seta absent; pronotum relatively short, subquadrate,
length/greatest width about 1.42; widest near middle; base, apex slightly narrowed; basal impression closed posteriorly;
outer marginal groove evident in dorsal view; marginal carina relatively broad; angular seta absent; precoxal setae absent;
sternopleural groove present.

Elytral striae deep, broad, inconspicuously punctate; supramarginal striae shallower than others; intervals narrow,
costate; intercalary stria complete; setae absent or one or two present near tip of marginal stria; cauda of female
prominent, trapezoidal.

Prosternum glabrous in both sexes; metasternum sulcate; transverse sulci rather broadly interrupted in midline; male
with median pollinose areas on Sterna I-V, that of V narrowly divided in midline by glabrous carina; Sternum IV of male
with median carina posterior to middle, but with pollinosity continuous anterior to carina; pollinosity of Sternum II a
narrow rectangle; female with lateral pits on Sternum III; Sternum VI of female with apical 0.5 impressed.

Male with all trochanters rounded; male with small ventral tooth on anterior femur; anterior tibia of male without
proximal tooth; middle, hind calcars pointed; hind one moderately large, raised slightly above level of spurs.

The male of this species resembles C. apertum in lacking pollinosity on the prosternum. It
can be separated from the latter by the presence of a median carina on Sternum V and the
absence of a proximal tooth on the anterior tibia. The female, like that of C. apertum , has
lateral pits on Sternum III but not on Sternum IV. The female of C. baldufi has a prominent,
trapezoidal cauda on the elytra. In both sexes, the basal impressions are closed posteriorly,
while in C. apertum they are open.

This is probably “Form C” of C. sculptile according to Blanchard (1889).

Range. — More extensive and less discontinuous than indicated by Bell (1970). South to
northern Florida and southern Mississippi; northeast to Delaware River Valley of New Jersey;
north to vicinity of Pittsburgh, Pennsylvania, northern Illinois and central Iowa; western limit
central Iowa, southeastern Missouri and northwestern Mississippi.

We have seen specimens from the following localities, in addition to those listed by Bell

(1970): ALABAMA: Blount Springs (CMP), Mt. Cheaha State Park (Cleburne Co.) (TB; CAS); ILLINOIS: Crab
Orchard Lake (Williamson Co.) (SI), Gorham (CAS), Olive Branch (CAS), Peoria (CNHM), Wedron (WS), Wolf Lake
(Union Co.) (WRS); INDIANA: Ellettsville (Monroe Co.) (IU), Jefferson Co. (IO), Lafayette (CNHM), New Harmony
(CNHM), Turkey Run State Park (Parke Co.) (CNHM; WRS); IOWA: Ames (IO), Burlington (MNHB), Ledges State
Park (Boone Co.) (IO); KENTUCKY: Blue Lick St. Park (Nicholas Co.) (RCG), Cumberland Falls (Whitley Co.) (UL),
Hematite Lake (RCG), Henderson (WRS), Pine Mountain (Harlan Co.) (PA), Whitley City (McCreary Co.) (RCG);
MARYLAND: Elk Neck St. Pk. (UD); MISSISSIPPI: Charleston (IO), Lucedale (CU); MISSOURI: Creve Coeur Lk.
(St. Louis Co.) (MO), Cape Girardeau (MO), Reynolds Co. (MO), Rockwoods Res. (St. Louis Co.) (MO), St. Charles
(CAS; UW); NEW JERSEY: Phillipsburg (CAS); NORTH CAROLINA: Black Mts. (CAS; PA), Cove Creek
(Haywood Co.) (PK), Macon Co. (RCG), Wine Spring Bald (LS); OHIO: Cincinnati (CAS; UM); PENNSYLVANIA:
Allegheny Co. (CMP; CU), Darby (CAS), E. Park (CAS), Easton (CAS), Ogontz (CAS), Vella Novo (Montgomery Co.)
(CAS); SOUTH CAROLINA: Clemson College (WS); TENNESSEE: Blount Co.. Thunderhead Mt. 6000 ft. (PA),
Cades Cove, Gt. Smoky Mt. Nat. Pk., 2000 ft. (CNHM), Chimneys Campground, Gt. Smoky Mt. Nat Pk., 2800 ft.
(CU), Clarksville (UK), Gatlinburg (CNHM), Model (Stewart Co.) (DY), Newfound Gap, Gt. Smoky Mt. Nat. Pk.,
5000 ft. (CU), Quinland Lk. (Putnam Co.) (TB); VIRGINIA: Nelson Co. (NMNH).

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Bionomics. — Recorded from American Chestnut, and white oak logs (Bell 1970).
Clinidium ( Arctoclinidium ) apertum Reitter 1880

Clinidium apertum Reitter 1880: 29-30.

Clinidium ( Arctoclinidium ) allegheniense georgicum Bell and Bell 1975.

Clinidium ( Arctoclinidium ) apertum (Reitter) Bell and Bell 1978.

Type Material. — ( apertum ) HOLOTYPE male, labelled: “Himalaya, Clinidium apertum India, Reitt.” (with
red “typus” label) (NMW). The locality data on this specimen must be erroneous; ( georgicum ) HOLOTYPE male,
labelled: “GEORGIA, Cartersville, 26-III-39, P. W. Fattig” (NMNH); PARATYPES one male, same data as holotype
(GA); two females, same data as holotype (CNHM); one male, labelled “Athens, Georgia, 6-X-54, K. Parrish” (GA); two
males, labelled “West Pace’s Ferry X, Marietta Hgy (Dekalb Co.), 12-IX-54, W. H. Cross” (UVM); one female, labelled:
“Dallas, 16-IV-44, P. W. Fattig” (UVM).

Description. — Length 5. 5-7.0 mm. Antennal Segments VII-X with tufts of minor setae; basal setae present on
Segments VII-X or VIII-X; temporal seta absent; pronotum moderately long, length/greatest width about 1.48; widest
near middle, base scarcely narrowed, apex moderately so; basal impressions oblique, relatively long, about 0.38 of length of
pronotum, widely open posteriorly; outer marginal groove clearly visible in dorsal view; marginal carina relatively broad;
angular seta, precoxal setae absent; sternopleural groove present.

Elytral striae deep, broad, inconspicuously punctate; supramarginal stria shallower than the others; intervals narrow,
costate; intercalary stria complete; setae absent or one or two present near apex of marginal stria; cauda of female elytra
very small, rounded.

Prosternum glabrous in both sexes; metasternum sulcate; transverse sulci rather narrowly interrupted in midline; male
with median pollinose areas on Sterna II-IV, in some specimens also on Sternum V, latter not carinate; pollinosity of
Sternum II resembling letter “T”, or with stem of “T” disconnected or absent, leaving curved transverse bar; female with
lateral pits on Sternum III; Sternum VI not impressed; male with all trochanters rounded, anterior femur with large but
obtuse ventral tooth, anterior tibia with large but obtuse proximal tooth; middle calcar narrow, pointed; hind calcar
triangular, acute, not raised above level of spurs, much smaller than in C. baldufi.

The broadly open basal impressions will separate this species from all others of the eastern
U.S. except for C. valent inei (and some specimens of rosenbergi). The sulcate metasternum
separates it from valentinei. The male resembles C. baldufi in lacking pollinosity on the
prosternum, but differs in having a proximal tooth on the anterior tibia, in lacking the median
carina on Sternum V, and in having a smaller hind calcar. The female resembles C. baldufi and
C. valentinei in having pits in Sternum III but not Sternum IV. It differs from the former in
having a rounded cauda, and from the latter in lacking an impression on Sternum VI.

This species is divided into two subspecies, separated on the presence or absence of a median
pollinose area on Sternum V of the male. An isolated female specimen labelled “Mobile,
Loding” (MCZ) from Southern Alabama, cannot be identified to subspecies.

Clinidium ( Arctoclinidium ) apertum apertum Reitter 1880
(Figs. 114, 120)

Clinidium apertum Reitter 1880: 29-30.

Clinidium ( Arctoclinidium ) allegheniense georgicum Bell and Bell 1975.

Clinidium ( Arctoclinidium ) apertum apertum (Reitter) Bell and Bell 1978.

Description. — Male without median pollinose area on Sternum V; pollinosity of Sternum II of male in most
specimens forming a broken “T”, in a few (including holotype of C. apertum ), an unbroken “T”, in a few a curved
transverse bar, stem of “T” entirely absent.

Range. — Mountains of northern Georgia, recorded only from the type series.

Clinidium ( Arctoclinidium ) apertum allegheniense Bell and Bell 1975

Clinidium allegheniense allegheniense Bell and Bell 1975: 65-66.
Clinidium apertum allegheniense Bell and Bell 1978.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

Type Material. — HOLOTYPE male, labelled: “Allegheny, l-VI-24, coll. Chermock” (UK). This locality is now
the Northside district of Pittsburgh. PARATYPES two females, same data as holotype (UK); one male, one female, same
data (MCZ); 13 males, two females, labelled “PENNSYLVANIA:- Wall, VI-21, H. Klages Collection” (CMP).

Description. — As described for C. apertum s. str., except that median pollinose area is present on Sternum V.
Range. — Southwestern Pennsylvania, and possibly mountains of western North Carolina.
In addition to the type series, we have studied the following additional specimens from

Pennsylvania: one male, one female, labelled: “Squaw’s Run, Pa. i-VI-24, coll. Chermock, L. J. Milne collection”
(UNH). We provisionally assigned the following specimens from western North Carolina to this subspecies; one male, two
females, labelled: “Mount Mitchell, Black Mts., North Carolina” (CAS). These Specimens have the median

pollinose area on Sternum V as in C. allegheniense. They are atypical in having the inner
impression of Sternum VI longer and more deeply impressed, and in the male, the stem of the
“T” is dilated posteriorly.

Clinidium ( Arctoclinidium ) rosenbergi Bell 1970
(Figs. 103, 112, 118)

Rhysodes sculptilis Newman 1838: 666 (partim )

Clinidium ( Arctoclinidium ) rosenbergi Bell 1970: 315-316.

Type Material. — HOLOTYPE male, labelled: “Turkey Run State Park, Parke Co., Indiana, May 13, 1950, coll.
Ross T. Bell” (MCZ 31749). PARATYPES one female, same data as holotype (UVM); one male, same locality, coll.
Buser, May 3, 1953 (UI); three females, labelled: “Parke County, Indiana, coll. N. M. Downie, May 8, 1965” (WR).

Description. — Length 6. 2-7. 8 mm. Antennal Segments VII-X with tufts of minor setae; basal setae present on
Segments VII-X or VIII-X; temporal seta absent; pronotum moderately long, length/greatest width 1.44; widest near
middle, sides nearly parallel to weakly curved; basal impression closed posteriorly (open in a few specimens); outer
marginal groove evident in dorsal view; marginal carina relatively broad; angular seta, precoxal seta absent; sternopleural
groove present.

Elytral striae deep, broad, inconspicuously punctate; supramarginal stria shallower than the others; intervals narrow,
costate; intercalary stria complete; setae absent or one or two present in or just above apex of marginal stria; cauda of
female small, rounded.

Prosternum of male with medial pollinose area; metasternum sulcate; male with median pollinose areas on Sterna
I-IV, median area of Sternum V glabrous; pollinose area of Sternum II relatively broad, constricted near middle, apex,
base equally broad; female with lateral pits on Sternum III, IV equally large; Sternum VI impressed in some female
specimens, not impressed in other ones.

Male with all trochanters rounded; anterior femur of male with ventral tooth; anterior tibia of male with strong angle
proximad to cleaning organ, representing poorly defined proximal tooth; calcars pointed; hind one very small, scarcely
longer than middle one.

Males from this species resemble those of C. sculptile in having a median pollinose area on
the prosternum, but lack pollinosity at the middle of Sternum V and have a very small hind
calcar. Females have equally large lateral pits in Sterna III and IV, while in C. sculptile the
pits of Sternum IV are smaller than those of III.

Range. — North to the Great Lakes in Pennsylvania, Ohio, and Indiana. West to the
Mississippi River in Tennessee and southern Illinois, and slightly west of the river near St.
Louis, Missouri. South to southwestern Tennessee and western North Carolina. East to the
Delaware River in Pennsylvania, but not known from east of the Appalachians south of
Pennsylvania. Bell (1970) discussed female specimen from Mobile, Alabama, which may be
this species, though the shape of the pronotum is unusual. It might be a distinct, though closely
related species, but males must be collected to be sure of its status. A specimen labelled:
“Treesbank, Manitoba” (BSRI) is likely to have an incorrect locality label.

We have studied the following specimens in addition to those listed by Bell (1970): ILLINOIS:
Wolf Lake (Union Co.) (WRS); INDIANA: Lafayette (CNHM); KENTUCKY: Anchorage (UL), Mammoth Cave N.
Park (WRS), Wolf Creek (Wayne Co.) (UL); MISSOURI: St. Charles (UW); OHIO: Cincinnati (UM), Clermont Co.
(UM), Cleveland (HL), Columbiana Co. (UD), Oxford (NC); PENNSYLVANIA: Blain (AP). Harmerville (CMP).
Jeanette (CMP).

Quaest. Ent., 1985, 21 (1)

Bell and Bell

Variation. — This species will merit additional study when more material is available.
Sternum VI is not at all impressed in some specimens, mostly from western parts of the range,
but is deeply impressed in others, especially ones from eastern localities. There is also
considerable variation in the shape of the pronotum. Most specimens from Kentucky have a
subquadrate pronotum with the sides nearly parallel. In specimens from other regions, the base
and apex of the pronotum are more narrowed.

Clinidium ( Arctoclinidium ) sculptile (Newman) 1838
(Figs. 115, 121)

Rhysodes sculptilis Newman 1838: 666 (partim )

Clinidium sculptile (Newman) Chevrolat 1873a.

Clinidium ( Arctoclinidium ) sculptile (Newman) Bell 1970.

Type Material. — LECTOTYPE male, labelled: “Wheeling, Virginia” (BMNH). This locality is now in West
Virginia. PARALECTOTYPES: Newman listed two localities. Wheeling and Mount Pleasant, Ohio. The latter was
represented by two females. One of these is not conspecific with the male, but are the species described by Bell as C.
rosenbergi, while the other is conspecific with the holotype.

Description. — Length 6. 5-7.6 mm. Antennal Segments VII-X with tufts of minor setae; basal setae present on
Segments VII-X or VIII-X; temporal seta absent; pronotum moderately long, length/greatest width about 1.45; pronotum
subquadrate, lateral margins nearly parallel; basal impression closed posteriorly; outer marginal groove evident in dorsal
view; marginal carina relatively broad; angular seta, precoxal setae absent; sternopleural groove present.

Elytral striae deep, broad, inconspicuously punctate; supramarginal stria impressed, shallower than the others;
intervals narrow, costate; intercalary stria complete; setae absent or one or two present in or just above apex of marginal
stria; cauda of female prominent, rounded.

Prosternum of male with median pollinose area; metasternum sulcate; male with median pollinose areas on Sterna
II-V; pollinosity of Sternum II constricted near middle, anterior margin broader than posterior one; female with large
lateral pits on Sternum III, smaller ones on Sternum IV: Sternum VI of female impressed in apical 0.5.

Male with all trochanters rounded; anterior femur of male with very small, obtuse ventral tooth or angle; anterior tibia
of male with small, indistinct angle in place of proximal tooth; calcars pointed; hind calcar rather large, about 1 .3 longer
than middle one.

Males of C. sculptile resemble those of C. rosenbergi, in having a median pollinose area on
the prosternum, but differ from the latter species in having median pollinosity on Sternum V
and in having a larger hind calcar. Females differ from all other members of the subgenus in
having large lateral pits on Sternum III and smaller ones on Sternum IV.

Range. — More extensive than supposed by Bell (1970). Primarily Appalachian from north
central Alabama to southern New York (Catskill Mts.), but also in the Piedmont from
northern Virginia northwards. Midwestern records are from central Kentucky, southern Ohio
and Indiana, and the vicinity of Saint Louis, Missouri. C. sculptile is the commonest member of
the genus in the eastern part of its range, but is much rarer than C. baldufi and C. rosenbergi in
the Midwest. There are several specimens labelled as coming from localities far beyond the
range as described below. Among these are some from unspecified localities in Florida (MO)
and Texas (CAS), and from Westview, Millvale, and Squaw’s River, in Manitoba (BSRI). We
regard these records as dubious.

We have studied specimens from the following localities in addition to those listed by Bell

(1970): ALABAMA: Sawdust ( WRS); DELAWARE: Newark (UD); DISTRICT OF COLUMBIA: Rock Creek Park
(AU); GEORGIA: Athens (UW), Clayton 2000-3700 ft. (CAS; CNHM; BMS); INDIANA: Turkey Run Park (Parke
Co.) (CNHM); KENTUCKY: Mammoth Cave Nat. Park (TB); MARYLAND: Elk Neck St. Park (UT; UD), Forest
Glen (NMNH; AU), Glen Echo (Montgomery Co.) (WRS), Catoctin Mtn. (Frederick Co.) (AU); MISSOURI: St.
Charles (UW), St. Louis (CU); NEW JERSEY: “N. J.” (CAS; CNHM); NORTH CAROLINA: Blue Ridge (LUN),
Transylvania Co. (RCG); OHIO: Cincinnati (CAS); PENNSYLVANIA: Allegheny (CMP; MO), Bethayres (UW),
Cook Forest (UVM), Cooksburg (WS), Charter Oak (AP), Harrisburg (SDA; AP), Ingelnook (AP), Jeanette (CMP),
Keystone St. Pk. (WRS), Montebello (AP), Rockville (CU; CAS; AP); SOUTH CAROLINA: Oconee Co. (CAS; WRS);
TENNESSEE: Cades Cove, Great Smoky Mts. Nat., Pk. 2000' (CNHM); VIRGINIA: Brush Mts. (Montgomery Co.)

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

(VP), Potomac River (Fairfax Co.) (CAS), Turkey Run, McLean (UVM); WEST VIRGINIA: Burlington (Mineral Co.)
(CMP), White Sulphur (CNHM).

Bionomics. — Recorded from pitch pine and tulip-tree ( Liriodendron ) Bell (1970).
SUBGENUS CLINIDIUM SENSU STRICTO

Type species. — Clinidium guildingii Kirby.

Description. — Antennal stylet relatively large; tufts of minor setae present on Segments IV-X, V-X, VI-X, or
VII-X, or else minor setae entirely absent except for Segment XI; temporal setae one to four; eye very narrowly crescentic
in most species, in a few species minute, round, resembling an ocellus, or bilobed, or divided into two ocellus-like parts, or
large, hemispheral. In many species, completely pigmented in some, probably old individuals; marginal groove of pronotum
single; pronotum with angular seta, and one or more marginals; in some species, also with discal or basal setae;
sternopleural grooves absent; elytral striation incomplete; marginal stria fourth or fifth from suture; supramarginal seta
absent; inner elytral intervals flat to convex, not carinate; elytral setae numerous in most species; metasternum with or
without median sulcus; base of anterior tarsus opposite cleaning organ.

The reduced striation makes this subgenus similar to Tainoa, from which it is distinguished
by the position of the cleaning organ and the complete parasutural stria which is not
abbreviated anteriorly.

The subgenus is large and the species are quite varied in appearance. Probably it is
restricted to the Neotropical Realm, from Guatemala southwards in Central America, and in
the Andean Region south to Ecuador. The only records from the Amazon Basin are from the
upper or western portion, while the easternmost record from the northern coast is from
Cayenne. The subgenus is widely distributed also in the Greater and Lesser Antilles.

Two of the four species of the C. beccarii group, C. beccarii and C. argus , are recorded from
Old World localities, the former species from New Guinea and the latter from the Philippines.
Each is known from only one specimen, and we suspect that both are mislabelled, as they are
closely related to Central American species.

Phytogeny. — We divide the subgenus into six species groups, most of which are probably
monophyletic. Possible exceptions are noted in the discussions of the groups in question.

I. impressum group. Tufts of minor setae on Antennal Segments IV-X; eye
large, broadly oval; temporal, pronotal setae absent. One species. This
group might really belong to Rhyzodiastes , and have secondarily lost the
anterior part of the paramedian groove.

II. granatense group. Tufts of minor setae VII-X. Three species.

III. insigne group. Tufts of minor setae VI-X. Four species.

IV. guildingii group. Tufts of minor setae on Segments V-X; anterior median
pit very small. 25 species.

V. cavicolle group. Tufts of minor setae V-X; anterior median pit greatly
enlarged. Nine species.

VI. beccarii group. Tufts of minor setae absent; eye constricted or divided.

Four species.

KEY TO SPECIES

1 Eye large, broadly oval; median groove of pronotum with middle 0.33

dilated, separated by constriction from anterior median pit; temporal,
pronotal setae absent ( impressum group)

C. impressum new species, p. 99

Quaest. Ent., 1985,21 (1)

Bell and Bell

I' Eye small, narrowly crescentic, or constricted, or ocelliform, or divided;

median groove not dilated; temporal, pronotal setae present 2

2 (T) Outer antennal segments with tufts of minor setae on Segments V-X,

VI-X, or VII-X; eye narrowly crescentic or ocelliform (may be concealed

by heavy pigmentation) 3

2 ' Outer antennal segments without tufts of minor setae; eye bilobed or

divided ( beccarii group) 43

3 (2) Tufts of minor setae present on Antennal Segments VI-X or VII-X, but

absent from Segment V 4

3' Tufts of minor setae present on Antennal Segments V-X 10

4 (3) Tufts of minor setae present on Segments VII-X, but absent from Segment

VI (granatense group) 5

4' Tufts of minor setae present on Segments VI-X ( insigne group) 7

5 (4) Eye narrowly crescentic; metasternum sulcate; head as wide as long; one

temporal seta; pronotum with one angular seta and without or with one
marginal seta, near angular; otherwise without pronotal setae; male
without proximal tooth on anterior tibia; male calcars triangular, not
notched above; female (where known) with lateral pits in Sternum III and
IV. Anterior median pit less enlarged 6

5' Eye small, round, ocelliform; metasternum not sulcate; head longer than
wide; two temporal setae; pronotum with one angular, two marginals,
anterior to middle, one basal, two discal setae; male with proximal tooth on
anterior tibia; male calcars notched above; female with lateral pits on
Sternum IV, not III; anterior median pit very large
C. incudis Bell, p. 1 14

6 (5) Intercalary stria complete; one marginal seta on pronotum; transverse

sulcus of Sternum V nearly complete in male; middle calcar obliquely

truncate at tip C. hammondi new species, p. 1 1 3

6' Intercalary stria abbreviated; marginal seta absent; transverse sulcus of

male Sternum V broadly interrupted; middle calcar acutely pointed

C. granatense Chevrolat, p. 1 1 3

7 (4') Temporal lobes convergent posteriorly; anterior median pit of pronotum

small to obsolete; antennal stylet very small; male protibia without

proximal tooth 8

7' Temporal lobes divergent posteriorly; anterior median pit very large, with
tubercle; antennal stylet very large; male protibia with proximal tooth

(metasternum sulcate; intercalary stria abbreviated posteriorly)

C. dubium Grouvelle, p. 1 15

8 (7) Metasternum with median sulcus; intercalary stria abbreviated posteriorly 9

8' Metasternum not sulcate; intercalary stria entire

C. boroquense Bell, p. 1 17

9 (8) Preapical tubercles truncate, medial angles of tubercles well separate; head

flattened, as wide as long; parasutural stria without setae

C. insigne Grouvelle, p. 115

9' Preapical tubercles sinuate, medial angles lobate; head convex, longer than
wide; parasutural stria with many setae

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby 95

10 (3')
10'

11 (10)
11'

12 (11)
12'

13 (12)
13'

14 (13)
14'

15 (13')
15'

16 (15)
16'

17 (16)
17'

18 (15')
18'

19 (18)
19'

20 (19')
20'

21 (18')
21'

22 (21')

C. howdenorum new species, p. 1 16

Anterior median pit of pronotum very small, not wider than median groove

(guildingii group)

Anterior median pit very large, much broader than median groove

(cavicolle group)

Intercalary stria entire; pronotum with discal seta

Intercalary stria abbreviated, ending blindly posteriorly; pronotum without

discal setae

Apex of intratubercular stria impressed, apical, preapical tubercles distinct
Apex of intratubercular stria not impressed; preapical, apical tubercles not

separated

Paramedian groove (basal impression plus discal stride), 0.6-0. 9 of length

of pronotum

Paramedian groove 0.5 or less of length of pronotum

Paramedian groove about 0.9 of length of pronotum; preapical tubercle

angulate posteriorly, angle overhanging subapical space

C. integrum Grouvelle, p. 1 1 9

Paramedian groove about 0.6 of length of pronotum; preapical tubercle not

angulate nor overhanging subapical space

C. pilosum Grouvelle, p. 1 19

Median lobe joined laterally to antennal lobes; frontal grooves not joined to

tentorial pits

Median lobe not joined laterally to antennal lobes; frontal grooves joined to

tentorial pits

Precoxal seta absent

Precoxal seta present C. alleni new species, p. 122

Median lobe short, ending opposite anterior or mid level of eye;

paramedian groove 0.5 length of pronotum

C. whiteheadi new species, p. 122

Median lobe longer, ending posterior to mid level of eye; paramedian

groove short, 0.2 length of pronotum C. haitiense Bell, p. 125

Precoxal seta present

Precoxal seta absent

Elytral humeri strongly narrowed; metasternum with deep median sulcus

C. oberthueri Grouvelle, p. 121

Elytral humeri weakly narrowed; median sulcus of metasternum very

shallow to absent

Discal stride of pronotum present; eye elongate

C. humboldti new species, p. 123

Discal stride absent; eye very small, short

C. trionyx new species, p. 1 24

Parasutural stria with complete series of setae; paramedian groove about
0.5 of length of pronotum C. jolyi new species, p. 1 20

Parasutural stria without setae; paramedian groove 0.3 or less of length of
pronotum

Median lobe elongate, extending posterior to middle of eye; frontal grooves

Quaest. Ent., 1985,21 (1)

Bell and Bell

deep, narrow, both margins equally sharp, both conspicuously pollinose

C. corbis Bell, p. 1 26

22' Median lobe short, ending opposite anterior part of eye; frontal grooves

shallow, lateral margin indistinct, margins not or but faintly pollinose 23

23 (22') Sutural interval narrow, convex; female with lateral pits distinct

C. jamaicense Arrow, p. 128

23' Sutural interval broad, flat; female with lateral pits indistinct

C. chiolinoi Bell, p. 1 28

24 (12') Metasternum with median sulcus; three or four temporal setae; occipital

setae absent 25

24' Metasternum without median sulcus; one temporal seta; one pair of crossed
occipital setae C. rossi Bell, p. 1 29

25 (24) Median lobe joined to antennal lobe; discal stride absent; precoxal setae

absent C. penicillatum new species, p. 1 3 1

25' Median lobe separate from antennal lobe; discal stride present; precoxal

setae present 26

26 (250 Antennal stylet very short, acute; subapical, apical tubercles one

continuous elongated lobe; median sulcus narrow 27

26' Antennal stylet long, slender, acute; tip of subapical tubercle abrupt,
truncate, not continuous with apical tubercle; median sulcus wider
C. kochalkai new species, p. 132

27 (26) Intratubercular stria entire, thin, pilose line anterior to tubercular

punctures; frontal groove deeper; median lobe narrower

C. segne new species, p. 1 3 1

27' Intratubercular stria abbreviated from tubercular punctures; frontal groove
shallow, median lobe wider C. dormans new species, p. 1 30

28 (11') Metasternum without median sulcus; male without proximal tooth on

anterior tibia; middle, hind tibiae with false spurs (West Indian species) 29

28' Metasternum with median sulcus; male with proximal tooth on anterior

tibia; false spurs absent (South American species) 32

29 (28) Apical tubercles barely touching in midline above a large space; female

with median tubercles on Sternum VI; discal stride 0.5 as long as

pronotum; temporal setae two to four C. guildingii Kirby, p. 133

29' Apical tubercles broadly contiguous in midline, without conspicuous space
below them; female (where known) with transverse scarp on Sternum VI;
discal stride in most specimens less than 0.5 of length of pronotum; two
temporal setae 30

30 (29') Intratubercular stria not impressed, represented only by row of fine

punctures; marginal stria incomplete anteriorly; male with ventral surface
of anterior femur with many tubercles; calcars not angulate dorsally

(female unknown) C. microfossatum new species, p. 134

30' Intratubercular stria, marginal stria impressed, complete; male without
ventral tubercles on anterior tibia; calcars angulate dorsally; female with
transverse scarp on Sternum VI 31

31 (30') Basal impression plus discal stride 0.35 to 0.40 of length of pronotum;

calcars weakly angulate dorsally; female with shallow impression posterior

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

to scarp of Sternum VI, forming obtuse angle in lateral view; impression
convex, glabrous in midline, with oval pollinose concavity on either side

C. smithsonianum new species, p. 1 34

31' Basal impression plus discal stride 0.25 or less of length of pronotum;

calcars strongly angulate dorsally; female with deep impression posterior to
scarp of Sternum VI, forming right angle in lateral view; impression

entirely pollinose C. planum (Chevrolat), p. 135

32 (28") Female with Sternum VI deeply impressed in posterior 0.33, with pair of

notches defining median lobe (male unknown) 33

32' Female with Sternum VI not impressed 34

33 (32) Parasutural stria without setae; median lobe of Sternum VI of female

narrow, trapezoidal C. pala new species, p. 139

33' Parasutural stria with several setae; median lobe of Sternum VI of female
broad, rectangular C. excavatum new species, 138

34 (32') Parasutural stria without setae; frontal space entirely pollinose; male with

calcars acute, proximal tibial tooth acute C. rojasi Chevrolat, p. 136

34' Parasutural stria with three or four setae; frontal space glabrous in middle;

male with calcars obtuse, proximal tooth of anterior tibia represented by
obtuse angle C. bechyneorum new species, p. 138

35 (10') Intercalary stria entire; anterior median pit with central tubercle 36

35' Intercalary stria abbreviated posteriorly; anterior median pit without

median tubercle C. mathani Grouvelle, p. 140

36 (35) Metasternum without median sulcus C. humile new species, p. 140

36' Metasternum with median sulcus 37

37 (36') Disc of temporal lobe without isolated or semi-isolated setiferous puncture;

notopleural suture without pollinosity 38

37' Disc of temporal lobe with one large setiferous puncture, either isolated or

in narrow contact with posterior pilosity; notopleural suture with pollinosity 41

38 (37) Basal impression plus discal stride 0.5 or less of length of pronotum;

antennal stylet long 39

38' Basal impression plus discal stride more than 0.5 of length of pronotum;

antennal stylet short 40

39 (38) Discal stride 0.45 of length of pronotum, curved; margin of median groove

curved evenly into anterior median pit; basal setae absent

C. curvatum new species, p. 141

39' Discal striole 0.20 of length of pronotum, scarcely curved; margin of
median groove sinuate opposite tubercle, latter compressed
C.foveolatum Grouvelle, p. 142

40 (38') Dorsal surface of femora glabrous, anterior median pit closed anteriorly,

round; female with transverse sulci of all sterna broadly interrupted in
midline; Sternum VI of female with submarginal groove reaching nearly to
anterior margin C. cavicolle Chevrolat, p. 142

40' Dorsal surface of femora pilose; anterior median pit open anteriorly,
sinuate laterally; sterna of female with transverse sulci narrowly
interrupted in midline; Sternum VI of female with submarginal groove not
extending anterior to middle C. crater new species, p. 143

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Bell and Bell

41 (370 Margin of median groove curved evenly into that of anterior median pit;

basal impression plus discal striole 0.30 of length of pronotum; discal seta
of temporal lobe in isolated pollinose puncture in most specimens anterior

to hind margin of eye C. centrale Grouvelle, p. 144

41' Margin of median groove sinuate near anterior median pit; basal
impression plus discal striole 0.45 or more of length of pronotum; discal
seta of temporal lobe at or posterior to posterior margin of eye, narrowly
contacting pilosity of occiput 42

42 (41') Margin of median groove shallowly sinuate posterior to tubercle of anterior

median pit, nearly oblique; basal impression plus discal striole about 0.55

of length of pronotum C. validum Grouvelle, p. 144

42' Margin of median groove strongly emarginate posterior to tubercle; basal

impression plus discal striole about 0.45 of length of pronotum

C. spatulatum new species, p. 145

43 (2') Eye constricted but not divided; median groove narrow, much narrower

than anterior median pit; sternopleural groove incomplete; postantennal

groove narrowly pollinose 44

43' Eye divided into two ocellus-like organs; median groove very broad, as
broad as anterior median pit; postantennal groove broadly pollinose;

sternopleural groove complete 45

44 (43) Median groove of pronotum closed posteriorly, slightly constricted at
midpoint; posterior 0.5 of sternopleural groove marked by three separate
pilose spots; male with pair of tubercles on either side of midline on

abdominal Sterna III, IV; transverse sulci of male with only traces of

pollinosity, but with pits at medial ends

C. moldenkei new species, p. 1 46

44' Median groove broadly open posteriorly, not constricted at middle;

posterior 0.5 of sternopleural groove continuously pollinose; male with deep
longitudinal groove at middle of Sterna I-III, shallower one on Sternum IV;
no tubercles on Sterna III, IV; transverse sulci of male abdomen pollinose,

interrupted at midline C. sulcigaster Bell, p. 1 47

45 (43') Paramedian grooves about 0.5 of length of pronotum; male with middle,
hind calcars cultrate; pollinosity of Sterna II, III extending anteriorly onto

medial part of Sternum I C. argus new species, p. 148

45' Paramedian grooves over 0.67 of length of pronotum; male with middle,
hind calcars triangular; Sternum III with transverse pollinose band, not
extending anteriorly to Sternum I C. beccarii Grouvelle, p. 148

THE IMPRESSUM GROUP

This group is characterized by the large, almost round eyes, and the absence of temporal
and pronotal setae. The median groove is dilated in the middle portion, and the dilation is
separated from the anterior median pit by a constriction. A very small tuft of minor setae is
present on Segment IV of the antenna, and a larger one on Segment V. The intercalary stria is
complete, while the intratubercular stria is abbreviated posteriorly.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

The large eyes are unique within the genus, and are reminiscent of those of Rhyzodiastes
subgenus Rhyzostrix. The dilation of the median groove is similar to that of Clinidium
kochalkai , but in the latter species the anterior median tubercle is not enlarged, the eyes are
small and narrow, and temporal and pronotal setae are present. Only one species belongs to this
group.

Clinidium ( sensu stricto ) impressum new species
(Fig. 124)

Type Material. — HOLOTYPE male, labelled: “Guyane Franc., St. Laurent du Maroni, colln. Le Moult,
Clinidium impressum Grouv.” (MNHN). The latter name was never published. Segments VI-XI of the antenna are
missing from the holotype.

Description. — Length 5.3 mm. Antennal Segment IV with small tuft of minor setae; Segment V with larger one;
Segments VI-XI missing from holotype; Segments III-V each with subapical ring of pollinosity; head slightly longer than
wide; frontal grooves shallow, linear, not pollinose; median lobe triangular, tip pointed, opposite middle of eye; frontal
space very narrow; temporal lobe rounded, nearly glabrous, small, Finely pollinose area near posterior margin; eye large,
nearly round, but short, less than 0.33 of length of temporal lobe; orbital groove absent; temporal setae absent; one pair of
postlabial setae.

Pronotum elongate, oval, length/greatest width 1.61, widest near middle, sides strongly curved, apex, base strongly
narrowed, rounded; median groove narrowly dilated, separated from median pits by constrictions; anterior median pit
large, round; posterior median pit narrow, elongate; basal impression open posteriorly, tapered anteriorly, preceded by very
short discal stride; combined length of basal impression, discal striole about 0.25 of length of pronotum; marginal groove
very narrow, invisible in dorsal view; pronotal setae absent; notopleural suture glabrous; sternopleural groove absent;
precoxal setae absent.

Elytra rather elongate; sutural interval nearly flat; Intervals II, III convex; sutural stria impressed, punctured,
complete; parasutural impressed, punctate, complete; intercalary impressed, punctures complete, joining parasutural
posteriorly; intratubercular effaced near base, for most of length not impressed, represented by row of punctures, slightly
impressed near apex, but apex effaced, preapical tubercle therefore not distinct from apical tubercle; apical tubercles
inflated, truncate posteriorly, nearly contiguous in midline; marginal stria entire, impressed, punctate; sutural stria with
one seta near apex; intercalary stria with Five setae in complete row; intratubercular stria with Five setae near apex, in row
of punctures on lateral face of apical tubercle; marginal stria eight or nine in complete row; metasternum not sulcate; male
with transverse sulci complete on Sterna III, IV, narrowly interrupted on Sternum V; Sternum IV of male with small,
round lateral pit; Sternum VI of male without transverse sulci, but with short submarginal sulcus, one pair of setae; male
without ventral tooth on anterior femur, without proximal tooth; calcars small, blunt; middle, hind tibiae with two equal
spurs, without false spurs; female unknown.

THE GRANATENSE GROUP

This group contains species in which tufts of minor setae are restricted to Antennal
Segments VII-X. There is one pair of postlabial setae. The anterior median pit is expanded,
several times wider than the median groove, but is not tuberculate. The paramedian grooves are
about 0.5 as long as pronotum. The sternopleural groove is present. In the species in which the
female is known, Sternum VI of the female has a median pit. The eye is either narrowly
crescentic or is ocelliform. This group contains three species, two from northern Colombia, and
the third from Puerto Rico.

Phytogeny. — C. granatense and C. hammondi share several characters, including the
presence of a metasternal sulcus and a proximal tooth on anterior tibia of the male, indicating
that they are closer to one another than to C. incudis. The presence of a median pit on Sternum
VI of the female in the latter species suggests a real relationship with the Colombian species,
rather than just a coincidence in the arrangement of tufts of minor setae.

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Plate 11. Figs. 124-137. Subgenus Clinidium sensu stricto. Figs. 124-131. Head and pronotum, dorsal aspect; Fig. 124, C.
(s. str.) impressum new species; Fig. 1 25, C. (s. str.) hammondi new species; Fig. 1 26, C. (s. str.) granatense Chevrolat;

Fig. 127, C. (s. str.) incudis Bell; Fig. 128, C. (s. str.) dubium Grouvelle; Fig. 129, C. (s. str.) boroquense Bell; Fig. 130, C.
(s. str.) howdenorum new species; Fig. 131, C. (s. str.) insigne Grouvelle; Fig. 132, Head, left lateral aspect, C. (s. str.)
incudis Bell; Figs. 133-134, Left elytron, dorsal aspect; Fig. 133, C. (s. str.) hammondi new species; Fig. 134, C. (s. str.)
granatense Chevrolat; Figs. 135-136, Sterna IV-VI, right half; Fig. 135, C. (s. str.) insigne Grouvelle; Fig. 136, C. (s. str.)
insigne Grouvelle (Cali specimen); Fig. 137, Left elytron, apex, dorsal aspect, C. (s. str.) howdenorum new species.

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Plate 12. Figs. 138-150. Subgenus Clinidium sensu stricto. Figs. 138-144, Head and pronotum, dorsal aspect; Fig. 138, C.
fa. str.) integrum Grouvelle; Fig. 139, C. fa. str.) pilosum Grouvelle; Fig. 140, C. fa. str .) jolyi new species; Fig. 141, C. fa.
str.) oberthueri Grouvelle; Fig. 142, C. fa. str.) alleni new species; Fig. 143, C. fa. str.) whiteheadi new species; Fig. 144, C.
fa. str.) humboldti new species; Fig. 145, Right elytron, apex, posterior aspect, C. fa. str.) integrum Grouvelle; Figs.
146-148, Sterna V-VI, right half; Fig. 146, C. fa. str.) pilosum Grouvelle, female; Fig. 147, C. fa. str.) jolyi new species,
female; Fig. 148, C. fa. str.) alleni new species; Figs. 149-150, Left elytron, apex, dorsal aspect; Fig. 149, C. fa. str.) alleni
new species; Fig. 150, C. fa. str.) humboldti new species, female.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

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Plate 13. Figs. 151-162. Subgenus Clinidium sensu stricto. Figs. 151-157, Head and pronotum, dorsal aspect; Fig. 151, C.
(s. str.) trionyx new species; Fig. 152, C. (s. str .) haitiense Bell; Fig. 153, C. (s. str.) corbis Bell; Fig. 154, C. (s. str.)
jamaicense Arrow; Fig. 155, C. (s. str.) chiolinoi Bell; Fig. 156, C. (s. str.) rossi Bell; Fig. 157, C. (s. str.) dormans new
species; Figs. 158-159, Elytra, posterior aspect; Fig. 158, C. (s. str.) jamaicense Arrow; Fig. 159, C. (s. str.) chiolinoi Bell;
Fig. 160, Sterna V-VI, right half, male, C. (s. str.) trionyx new species; Fig. 161, Head, left lateral aspect, C. (s. str.)
trionyx new species; Fig. 1 62, Left elytron, apex, dorsal aspect, C. (s. str.) dormans new species.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

105

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Plate 14. Figs. 163-178. Subgenus Clinidium sensu stricto. Figs. 163-169, Head and pronotum, dorsal aspect; Fig. 163, C.
(s. str.) penicillatum new species; Fig. 164, C. (s. str.) segne new species; Fig. 165, C. (s. str.) kochalkai new species; Fig.
166, C. (s. str.) planum (Chevrolat); Fig. 167, C. (s. str.) guildingii Kirby; Fig. 168, C. (s. str.) microfossatum new species;
Fig. 169, C. (s. str.) smithsonianum new species; Fig. 170, Hind leg (excluding tarsus), female, C. (s. str.) penicillatum
new species; Figs. 171-173, Left elytron, apex, dorsal aspect; Fig. 171, C. (s. str.) microfossatum new species; Fig. 172, C.
(s. str.) planum (Chevrolat); Fig. 173, C. (s. str.) segne new species; Figs. 174-175, Sternum VI, female; Fig. 174, C. (s.
str.) guildingii Kirby; Fig. 175, C. (s. str.) smithsonianum new species; Fig. 176, Sternum VI, lateral aspect, female, C. (s.
str.) guildingii Kirby; Figs. 177-178, Elytra, posterior aspect; Fig. 177, C. (s. str.) guildingii Kirby; Fig. 178, C. (s. str.)
smithsonianum new species.

107

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

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Plate 15. Figs. 179-193 Subgenus Clinidium sensu stricto. Figs. 179-183, Head and pronotum, dorsal aspect; Fig. 179, C.
(s. str.) rojasi Chevrolat; Fig. 180, C. (s. str.) bechyneorum new species; Fig. 181, C. (s. str.) humile new species; Fig. 182,
C. (s. str.) mathani Grouvelle; Fig. 183, C. ( s . str.) cavicolle Chevrolat; Fig. 184, Anterior leg (excluding tarsus), male, C.
(s. str.) rojasi Chevrolat; Fig. 185, Elytra, posterior aspect, C. (s. str.) rojasi Chevrolat; Figs. 186-191, Sternum VI; Fig.
186, C. (s. str.) rojasi Chevrolat; Fig. 187, C. (s. str.) bechyneorum new species; Fig. 188, C. (s. str.) excavatum new
species; Fig. 189, C. (s. str.) pala new species; Fig. 190, C. (s. str.) humile new species; Fig. 191, C. (s. str.) cavicolle
Chevrolat; Fig. 192, C. (5. str.) mathani Grouvelle; Fig. 193, Left elytron, dorsal aspect, C. (s. str.) mathani Grouvelle.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

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Plate 16. Figs. 194-205. Subgenus Clinidium sensu stricto. Figs. 194-199, Head and pronotum, dorsal aspect; Fig. 194, C.
(s. str .) curvatum new species; Fig. 195, C. (s. str.) foveolatum Grouvelle; Fig. 196, C. (5. str.) crater new species; Fig. 197,
C. (s. str.) centrale Grouvelle; Fig. 198, C. (s. str.) spatulatum new species; Fig. 199, C. (s. str.) validum Grouvelle; Figs.
200-205, Sternum VI, female; Fig. 200, C. (s. str.) centrale Grouvelle; Fig. 201, C. (s. str.) spatulatum new species; Fig.
202, C. (s. str.) validum Grouvelle; Fig. 203, C. (s. str.) curvatum new species; Fig. 204, C. (s. str.) foveolatum Grouvelle;
Fig. 205, C. (s. str.) crater new species;

Ill

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Plate 17. Figs. 206-213. Subgenus Clinidium sensu stricto. Figs. 206-209, Head and pronotum, dorsal aspect; Fig. 206, C.
(s. str.) moldenkei new species; Fig. 207, C. (s. str.) argus new species; Fig. 208, C. (s. str .) sulcigaster Bell; Fig. 209, C. (s.
str.) beccarii Grouvelle (redrawn from sketch by R. Poggi); Fig. 210, Head, left lateral aspect, C. (s. str.) moldenkei new
species; Fig. 211, Prothorax, left lateral aspect, C. (s. str.) moldenkei new species; Figs. 212-213, Metasternum, abdomen,
right half; Fig. 212, C. (s. str.) moldenkei new species, male; Fig. 213, C. (s. str.) sulcigaster Bell, male.

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113

Clinidium ( sensu stricto) hammondi new species
(Figs. 125, 133)

Type Material. — HOLOTYPE male, labelled: “Bogota, Rhyzophagus ?, Clinidium granatense Chaudoir”
(BMNH). The type is in poor condition, broken into several pieces, but all important parts are present.

Description. — Length 6.0 mm. Antennal stylet very long, 0.4 as long as Segment XI; minor setal tufts present on
Segments VII-X; basal setae present on Segments VIII-X, sparse on VIII; antenna devoid of pollinosity; head triangular,
clearly broader than long; frontal grooves very shallow, obsolete anteriorly; median lobe short, broadly triangular, tip
acute, opposite anterior part of eye; temporal lobes convergent posteriorly, forming rounded medial angles, which are
narrowly separated; temporal lobe rounded posteriorly; eye short, rather broadly crescentic; orbital groove pollinose,
complete, joined posteriorly to rather broad marginal band of pollinosity; one temporal seta, in prominent puncture near
occipital margin; one pair of postlabial setae.

Pronotum oval, rather short, length/greatest width 1.40; widest near middle; sides strongly curved, apex truncate, base
rounded; median groove deep, linear; anterior median pit enlarged, round, width about 0.20 of width of pronotum; basal
impression open posteriorly, laterally, narrowed anteriorly, connecting to slightly curved discal stride, latter extending to
middle of pronotum; medial part of disc sloped towards median groove; lateral part of disc convex; marginal groove
prominent, visible in dorsal view; angular seta present; one marginal seta, just anterior to angular; notopleural suture
glabrous; sternopleural groove nearly complete, interrupted near coxa; precoxal setae absent.

Elytra rather elongate; intervals convex; striae impressed, punctured, pollinose; sutural, parasutural striae complete,
anastomosing posteriorly; intercalary intratubercular, marginal striae entire; preapical tubercle strongly inflated, rounded;
apical tubercles slightly inflated, contiguous (Fig. 133); sutural, parasutural striae without setae; intercalary stria with
complete row of four or five setae; one seta at apex of intratubercular stria; three or four setae in apical 0.2 of marginal
stria; one seta each on apical, preapical tubercles; metasternum with complete median sulcus; transverse sulci of abdomen
coarsely punctate, pollinose, those of Sterna III, IV continuous, that of V narrowly interrupted in midline in male; Sternum
VI with transverse sulci joined to submarginal sulcus; Sternum VI with two setae; male without ventral tooth on anterior
femur, without proximal tooth on anterior tibia; middle calcar narrow, triangular, its tip obliquely truncate; hind calcar
smaller than middle one, raised above level of spurs; tibial spurs equal, large, false spur absent; female unknown.

This species is similar to C. granatense Chevrolat, but differs in having the intercalary stria
entire and deeply impressed, and the middle calcar truncate. It is a pleasure to name this
species for Peter Hammond, of the British Museum of Natural History, in gratitude for his aid
in this project.

Clinidium ( sensu stricto ) granatense Chevrolat 1873a
(Figs. 126, 134)

Clinidium granatensis Chevrolat 1873a: 216

Clinidium granatense (Chevrolat) Grouvelle 1903 (grammatical correction).

Clinidium ( sensu stricto) granatense (Chevrolat) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) female, labelled: “Nov. Gren., Clinidium granatense , Chev.
type” (MNHN). PARALECTOTYPES one female, labelled “Neu Granada, Madellin, Typus, granatense ” (NMW); one
male, labelled: “Bogota, granetense , Chevrolat, Typus” (NMW).

Description. — Length 5. 3-6.8 mm. Antennal Stylet very long, 0.4 as long as Segment XI; minor setal tufts present
on Segments VII-X; basal setae present on Segments VII-X or VIII-X; antenna devoid of pollinosity; head triangular,
clearly broader than long; frontal grooves very shallow, obsolete anteriorly; median lobe short, broadly triangular, tip
acute, opposite anterior part of eye; temporal lobes convergent posteriorly, forming rounded medial angles, latter narrowly
separated; temporal lobe rounded posteriorly; eye short, rather broadly crescentic; orbital groove pollinose, complete,
joined posteriorly to rather broad marginal band of pollinosity; one temporal seta, in prominent puncture at margin of
pollinosity near posterior margin of temporal lobe; one pair of postlabial setae.

Pronotum slightly more elongate than that of C. hammondi , length/greatest width about 1.48; widest near middle,
sides strongly curved; apex truncate, base rounded; median groove deep, linear; anterior median pit enlarged, width about
0.20 of width of pronotum; basal impression open posteriorly, laterally, narrowed anteriorly, connecting to discal striole,
latter extending to middle of pronotum; marginal groove prominent, visible in dorsal view; angular seta present; marginal
setae absent; notopleural suture glabrous; sternopleural groove nearly complete, interrupted near coxa; precoxal setae
absent.

Elytra rather elongate; intervals convex; striae impressed, coarsely punctured; pollinosity less continuous than in C.
hammondi ; sutural, parasutural striae complete, anastomosing near apex; intercalary stria abbreviated, ending blindly at
anterior part of preapical tubercle; intratubercular, marginal striae entire; preapical tubercle inflated, rounded; apical

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tubercles inflated, contiguous (Fig. 134); sutural, parasutural striae without setae; intercalary stria with complete row of
four or five setae; one or two setae at apex of intratubercular stria; four or five setae in apical 0.2 of marginal stria; one seta
each on apical, preapical tubercles; metasternum with complete median sulcus; transverse sulci of abdominal sterna
coarsely punctate, pollinose, continuous on Sternum III in both sexes, continuous on IV in male, narrowly interrupted in
female; widely separated on V in both sexes. Sternum VI with transverse sulci joined to submarginals; Sternum VI with
two setae; lateral pits present on both III, IV in both sexes very small in male; female with small median pit on Sternum
VI; male without ventral tooth on anterior femur, without proximal tooth on anterior tibia; middle calcar triangular, acute;
hind calcar broadly triangular, tip acute, raised above level of spurs; tibial spurs large, equal, false spur absent.

The abbreviated intercalary stria and the different shape of the middle calcar separate this
species from C. hammondi, to which it is otherwise strongly similar.

Clinidium ( sensu stricto) incudis Bell 1970
(Figs. 127, 132)

Clinidium ( sensu stricto) incis Bell 1970: 319.

The original spelling, incis, is incorrect. The name is derived from incus (anvil), based on
the type locality, El Yunque, the Spanish word for anvil. The genitive singular form of the word
is incudis (“of the anvil”).

Type Material. — HOLOTYPE male, labelled: “El Yunque, Puerto Rico, May, 1938, coll. P. J. Darlington”
(MCZ 31756). PARATYPES two females with same data as type (MCZ); two males, two females, from the same
locality, coll. T. B. Hlavac, L. Herman, Jr., 2200-3200 ft., Feb. 15-24, 1969 (MCZ).

Description. — Length 6. 1-7.5 mm. Antennal stylet very long, about 0.5 as long as Segment XI; tufts of minor
setae present on Segments VII-X; basal setae entirely absent; Segments I-V each with subapical pollinose ring; head longer
than broad; frontal grooves linear, nearly glabrous, deeper than in preceding species; median lobe triangular, tip acute,
posterior to eye; temporal lobes divergent posteriorly, not forming medial angles; temporal lobe rounded posteriorly,
broadly bordered with pilosity; eye minute, round, protruding, resembling an ocellus, located near middle of length of
head; orbital groove deeply impressed, pollinose, complete; two temporal setae; one pair of postlabial setae.

Pronotum rather short, length/greatest width 1.40; widest anterior to middle, lateral margins curved anteriorly,
oblique posteriorly; margin shallowly sinuate anterior to hind angle; median groove deep, rather broad, parallel posteriorly,
anteriorly gradually broadened to anterior median pit; latter large, about 0.25 of width of pronotum at level of pit; basal
impression small, triangular, closed posteriorly; discal stride curved, extending to middle of pronotum; marginal groove
linear, visible in dorsal view; angular seta present; two marginal setae, anterior to middle of pronotum; two pairs of discals,
opposite anterior part of anterior median pit; one pair of basals, medial to basal impressions; notopleural suture glabrous;
anterior part of sternopleural suture very shallow, incomplete; precoxal setae absent.

Elytra rather elongate; intervals convex; striae impressed, pollinose, inconspicuously punctate; all striae, including
intercalary, complete; preapical tubercle strongly inflated; apical tubercles inflated, nearly contiguous, but separately
rounded; sutural stria without setae; parasutural stria with two or three setae, anterior one near or anterior to middle;
intercalary stria with complete row of four or five setae; one seta near apex of intratubercular stria; marginal stria with two
to six setae in posterior 0.5; preapical tubercle with one seta; apical tubercle with one to three setae; metasternum without
median sulcus; abdominal Sterna III-VI each with uninterrupted transverse sulcus, latter consisting of row of very coarse
punctures; that of Sternum VI not joined to submarginal groove; Sternum VI of female with median pit; female with deep
lateral pit on Sternum IV; male without ventral tooth on anterior femur, but with small triangular proximal tooth on
anterior tibia; calcars triangular, dorsal margin separated from tibia by deep notch; tibial spurs large, equal; false spur
absent.

The form of the eye is unique within the genus though it recalls those of Shyrodes dohertyi
(Grouvelle) and Srimara planicollis Bell and Bell. This is the only species from the West Indies
which has the anterior median pit enlarged. The eye, the anterior median pit, and the divergent
temporal lobes easily separate this species from the only other Rhysodine from Puerto Rico,
Clinidium ( sensu stricto) boroquense Bell.

Range. — Puerto Rico. We have seen additional specimens from El Yunque, and have
collected it there ourselves. In addition, we have seen a specimen labelled “Puerto Rico:
Villalba, C. M. Matos, VI-30-1938” (MAY).

Bionomics. — Host species have not been recorded. Bell (1970) quotes observations by
Hlavac (in litt.) on this species in the field and its behavior in the laboratory.

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115

THE INSIGNE GROUP

In this group, the tufts of minor setae are present on Segments VI-X. The eye is crescentic.
The form of the anterior median pit varies among the species. This group contains four species,
two from northwestern South America, one from the “continental” island of Trinidad, and one
from Puerto Rico.

Phytogeny. — The Ecuadorian species, C. dubium, is very different from the remaining
ones, and possibly had an independent origin from the cavicolle group, which it resembles
except in the absence of the tuft of setae from antennal Segment V. The rest of the species
share the following characters: temporal lobes convergent posteriorly; antennal stylet rather
small; anterior tibia of male without proximal tooth; anterior median pit medium to small,
without tubercle. C. boroquense stands apart from the remaining two, in having the intercalary
stria complete, the metasternum not sulcate, and the intratubercular stria not impressed except
at the apex. The Ecuadorian (and possibly Colombian) species, C. insigne, is obviously closely
related to C. howdenorum of Trinidad, despite the wide geographic separation of the two.

Clinidium {sensu stricto) dubium Grouvelle 1903
(Fig. 128)

Clinidium dubium Grouvelle 1903: 129-130.

Clinidium ( sensu stricto) dubium (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Loja, Ecuador, C. dubium type” (MNHN).

Description. — Length 5.7 mm. Antennal stylet very long, 0.4 as long as Segment XI, acuminate; tufts of minor
setae present on Segments VI-X; basal setae present on Segments VI-X, sparse on VI: Segments I-VII with subapical
pollinose bands; head with length, width approximately equal; frontal grooves rather fine, shallow; median lobe triangular,
tip acute, opposite middle of eye; temporal lobe divergent posteriorly, posterior margin rounded; bordered with pilosity; eye
small, narrowly crescentic, about 0.25 of length of temporal lobe; orbital groove complete, pollinose; one or two temporal
setae; two pairs of postlabial setae.

Pronotum rather short, length/greatest width 1.36; widest slightly anterior to middle, lateral margins curved,
becoming oblique posteriorly; median groove deep, rather broad, slightly constricted near middle, anteriorly broadened
gradually, slightly sinuate where joined to anterior median pit; latter large, about 0.25 of width of pronotum at level of pit;
anterior median pit with round pollinose central tubercle; basal impression elongate, triangular, closed posteriorly, about
0.25 as long as pronotum; discal stride not distinct; marginal groove visible in dorsal view; three or four marginal setae;
angular seta absent; notopleural suture glabrous; sternopleural groove complete, pollinose; precoxal setae absent.

Elytra moderately elongate; striae impressed, pollinose, inconspicously punctate; intercalary stria abbreviated
posteriorly, ending blindly at level of anterior end of preapical tubercle; other striae entire; preapical tubercle inflated;
apical tubercles inflated, contiguous; intercalary stria with complete row of three or four setae; intratubercular stria with
one seta near apex; marginal stria with six or seven setae in apical 0.2; metasternum with complete, deep median sulcus;
hind coxa with conspicuous pollinose area on lateral margin; male with complete transverse sulci on Segments II- VI;
submarginal groove of Sternum VI not connected to transverse sulcus; male without ventral tooth on anterior femur, but
with acute proximal tooth on anterior tibia; calcars narrow, acute; tibial spurs equal, false spur absent.

This is the only species in the group that has a large anterior median pit with a central
tubercle. It is similar to members of the cavicolle group except in lacking a tuft on Segment V.
It will not key to any member of the cavicolle group, since the only member of the latter group,
C. mathani, to have the intercalary stria abbreviated posteriorly, lacks a central tubercle in the
anterior median pit and has long discal strioles on the pronotum.

Clinidium ( sensu stricto ) insigne Grouvelle 1903
(Figs. 131, 135, 136)

Clinidium insigne Grouvelle 1903: 132.

Clinidium ( sensu stricto) insigne (Grouvelle) Bell and Bell 1978.

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116

Bell and Bell

Type Material. — According to the original description, the type was from Ecuador, and was in the Oberthiir
collection. We were unable to locate a specimen labelled as a type. Possibly the description was based on a female
specimen, labelled: “Ecuador, Slemiradski 1882-1883, Clinidium insigne Grouv.” (MNHN), though this was not labelled
as a type, it is the only specimen of this species among the material studied by Grouvelle.

Description. — Length 7.0-7.4 mm. Antennal stylet conical, acuminate, moderately long, about 0.25 of length of
Segment XI; tufts of minor setae present on Segments VI-X; basal setae present on Segments VII-X; Segment I with
pollinose subapical band; pollinosity otherwise absent from antenna; head with length, width almost equal; frontal grooves
narrow, deep, pollinose; median lobe triangular, narrow, tip acute, just behind level of anterior margin of eye; temporal
lobes strongly convergent posteriorly, forming lobate medial angles, latter very narrowly separated; posterior margin
rounded, bordered with pollinosity; eye crescentic about 0.5 length of temporal lobe; orbital groove complete, pollinose; one
temporal seta arising from large pollinose puncture touching posteriolateral pollinose border of temporal lobe; two pairs of
postlabial setae.

Pronotum long, length/greatest width 1.60; widest near middle, sides curved; base rounded; median groove deep,
narrow, sides parallel except at slight expansion at basal 0.33 of length; anterior median pit elongate, oval, about 0.15 of
width of pronotum opposite the pit; central tubercle absent; basal impression narrow, oblong, open posteriorly; discal striole
deep, nearly straight, extending anteriorly beyond middle of pronotum; marginal groove deep, visible in dorsal view; six
marginal setae, angular seta absent; notopleural suture glabrous; sternopleural groove nearly complete, narrowly
interrupted near coxa; precoxal setae absent.

Elytra moderately elongate; striae impressed, pollinose, inconspicuously punctate; intercalary stria abbreviated
posteriorly, ending blindly at level of anterior end of preapical tubercle; other striae entire; preapical tubercle strongly
inflated, truncate posteriorly; apical tubercles strongly inflated, rounded, contiguous; parasutural striae without setae;
intercalary stria with complete row of three to five setae; intratubercular stria with one or two setae near apex; marginal
stria with six or seven setae in apical 0.33; preapical tubercle with one seta; apical tubercle with one seta; metasternum
with deep, complete median sulcus; female with transverse sulci complete on Sterna III-IV, interrupted on midline on V,
VI; female with transverse sulci of Sternum VI joined to submarginal groove; Sternum VI with two setae; female with
large lateral pit on Sternum IV (Fig. 135); tibial spurs slightly unequal; false spur absent.

We provisionally assign a male, labelled: “Cali, Cauca, Colombia, VI-30-38, C. H. Seevers”
(CNHM) to this species. It differs from the female holotype in having the transverse sulcus of
Sternum V continuous and the submarginal groove of Sternum VI widely separated from the
transverse sulci (Fig. 136). The first of these characters is likely to be a secondary sexual
difference, but the second is not. It might be a separate, but closely allied species. The male
from Cali has the following secondary sexual characters: anterior femur without ventral tooth;
anterior tibia without proximal tooth; both calcars angulate dorsally; middle calcar longer than
hind one; tips of calcars obtuse.

This species is closest to C. howdenorum of Trinidad, which has a narrower head, more
elytral setae, and a differently shaped preapical lobe on the elytra.

Clinidium ( sensu stricto) howdenorum new species
(Fig. 130)

Type Material. — HOLOTYPE male, labelled: “Morne Blue, 2700' TRINIDAD, W.I., Aug. 19, 1969, H. & A.
Howden” (BSRI).

Description. — Length 6.0 mm. Antennal stylet flattened, narrowly, obliquely truncate, 0.2 of length of Segment
XI; tufts of minor setae present on Segments VI-XI; basal setae present on Segments VI-X; Segment I with pollinose
subapical band; pollinosity otherwise absent from antennae; head longer than wide; narrower, more convex than in C.
insigne-, frontal grooves narrow, deep, pollinose; median lobe very narrow, tip acute, posterior to level of anterior margin of
eye; temporal lobes strongly convergent posteriorly, forming lobate medial angles, latter very narrowly separated; posterior
margin rounded; eye crescentic about 0.15 of length of temporal lobe; orbital groove complete, pollinose; three temporal
setae, respectively anterior to, opposite, posterior to eye; each seta base surrounded by pollinosity, latter extensively
“scalloping” lateral margin of temporal lobe; orbital groove complete; three pairs of postlabial setae.

Pronotum elongate, but less so than in C. insigne-, length/greatest width 1.50; widest near middle, sides curved; base
rounded; median groove deep, narrow, sides parallel except at slight expansion at basal 0.33 of length; anterior median pit
elongate, oval, about 0.20 of width of pronotum at pit; central tubercle absent; basal impression triangular, open
posteriorly; discal striole deep, curved, extending anteriorly beyond middle of pronotum; marginal groove deep, visible in
dorsal view; angular seta present; eight marginal setae; notopleural suture glabrous; sternopleural groove nearly complete,
narrowly interrupted opposite coxa; precoxal setae absent.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

117

Elytra moderately elongate; striae impressed, pollinose, inconspicuously punctate; intercalary stria abbreviated
posteriorly, ending blindly at level of anterior end of preapical tubercle; remaining striae entire; preapical tubercle strongly
inflated, medial angles lobate; medial margin of tubercle emarginate opposite tip of intercalary stria (Fig. 137); apical
tubercles inflated, rounded, contiguous; parasutural stria with 10 setae; intercalary with nine setae; intratubercular stria
with three or four setae near apex; marginal stria with 10-12 setae; preapical tubercle with three setae; apical tubercle with
one seta; metasternum with deep, complete median sulcus; male with transverse sulcus narrowly interrupted on Sternum
III, V, VI, complete on Sternum IV; male with distinct lateral pit on Sternum IV; Sternum VI with submarginal sulcus
broadly separated from transverse sulcus; Sternum VI with four setae; tibial spurs equal; false spur absent; male without
ventral tooth on anterior femur, without proximal tooth on anterior tibia; middle calcar acute, smaller than hind calcar;
latter narrowly truncate.

This species is obviously closest to C. insigne, but differs in the form of the antennal stylet,
in the head being narrower and more convex, the elytral setae being much more numerous, and
in the preapical tubercles being lobate.

It is a pleasure to name this species, the first described from Trinidad, after the collectors,
Anne and Henry Howden.

Clinidium {sensu stricto) boroquense Bell 1970
(Figs. 129, 137)

Clinidium ( sensu stricto) boroquense Bell 1970:321.

Type Material. — HOLOTYPE male, labelled: “El Yunque, Puerto Rico, May, 1938, coll. P. J. Darlington”
(MCZ 31757). PARATYPES, one male, one female, same label as holotype (MCZ); one male, from same locality, coll. T.

B. Hlavac, L. Herman, Jr., 2200-3200 ft., Feb. 15-24, 1969 (MCZ).

Description. — Length 4.0-5. 9 mm. Antennal stylet minute; tufts of minor setae present on Segments VI-X; basal
setae present, but sparse on Segments IX-X; Segment I with pollinose subapical band; pollinosity otherwise absent from
antenna; head slightly longer than wide; frontal grooves Fine, scarcely pollinose; median lobe triangular, short, tip opposite
anterior 0.2 of eye; temporal lobes convergent posteriorly, forming rounded, nearly contiguous medial angles; frontal space
very narrow, posterior margin of temporal lobe rounded, very narrowly margined with pilosity; lateral margin of head only
slightly curved; eye very narrowly crescentic, about 0.8 of length of temporal lobe; orbital groove complete, pollinose but
very narrow; two or three temporal setae, one anterior to eye, one opposite middle of eye; posterior one near posterior
margin of temporal lobe; two pairs of postlabial setae.

Pronotum rather long, length/greatest width 1.55; widest near middle, sides curved, abruptly rounded, narrowed to
apex, more gradually rounded to base; latter curved; margin shallowly sinuate anterior to hind angle; median groove deep,
nearly linear, margins parallel; anterior median pit very small, margins of median groove not at all expanded opposite it;
basal impression small, triangular, closed posteriorly; discal stride Fine, slightly curved, extending to middle of pronotum;
marginal groove Fine, visible in dorsal view; angular seta present; one or two marginals or absent, location differing among
the specimens; notopleural suture glabrous; sternopleural groove absent; precoxal setae absent.

Elytra moderately elongate; striae impressed, pollinosity scant; striae punctate; all striae complete; intratubercular
stria shallower than the others; preapical tubercle slightly inflated, apex of intratubercular stria shallowly impressed, so
preapical, apical tubercles not strongly separated; apical tubercles inflated, rounded, slightly separated; sutural striae with
one to four setae near base; intercalary stria with complete row of four or Five setae; intratubercular stria with one to three
setae near apex; marginal stria with three to Five setae in apical 0.20; preapical tubercle without setae; apical tubercle with
one seta; metasternum without median sulcus; transverse sulci of abdominal sterna impressed laterally, medial portion
replaced by row of several very coarse punctures; Sternum VI on each side with two punctures in place of transverse sulcus;
submarginal sulcus short; Sternum VI with two setae; female with lateral pits on Sterna IV, V; male without ventral tooth
on anterior femur, without proximal tooth on anterior tibia; calcars small, triangular, acute; hind calcar raised above level
of spurs; tibial spurs large, equal; false spur absent.

This species shows many points of similarity to C. insigne and C. howdenorum, especially in
the distribution of tufts of minor setae and in having convergent temporal lobes. The latter
species, however, has the anterior median pit enlarged, and lacks discal setae on the pronotum.

C. boroquense is not likely to be confused with the other known Puerto Rican species, C.
incudis , as the latter has a strongly enlarged anterior median pit, divergent temporal lobes, and
an ocelliform eye.

Range. — Puerto Rico. We have seen four additional specimens from the type locality,

labelled: “El Yunque, Puerto Rico, Luquillo Exp. For., Rte. 915, 1.5 mi. off Rte. 988, Mar. 29, 1976, A. Gillogly, H.
Harlan” (UVM). We have seen specimens from four additional localities: “Utuado, 11-15-1935, A. Ramirez" (MAY);

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118

Bell and Bell

“Adjuntas, VIII-1933, R. G. Oakley” (MAY); “5 mi. s. of Utuado, 3 July, 1979, coll. M. A. Ivie” (MAI); “Aguas Buenas,
forest at Aguas Buenas cave, 7-17-V-73, 250 m., S. Peck et al” (BSRI).

THE GUILDINGII GROUP

In this group, the tufts of minor setae are found on Segments V-X of the antenna, and the
anterior median pit is very small. In most species, the median groove is not widened at the
anterior median pit, while in a few species it it slightly widened. Most species have false spurs
on the middle and hind tibiae. A false spur is a rigid tooth projecting from the apical margin of
the tibia. In size and shape it resembles the true spurs. The eyes are narrow and crescentic.
Some species have a broad tooth or a slight cusp in place of the false spur.

This group is the largest in the subgenus, with 25 species. The range is substantially that of
the subgenus, except that members of this group are not known at present from Puerto Rico,
French Guiana, or Guatemala.

Phytogeny. — We provisionally divided the group into five sections. The oberthueri section
has both intercalary and intratubercular striae complete, and has the metasternum sulcate,
sometimes only very shallowly so. The jamaicense section has similar striation, but lacks the
metasternal sulcus. The rossi section has the intercalary stria complete, but has the
intratubercular not impressed near the apex, so that the preapical and apical tubercles are
fused. The metasternal sulcus is absent in C. rossi, but present in the other members of the
section. The guildingii section has the intercalary stria abbreviated posteriorly, and the
metasternum not sulcate. The rojasi section has the intercalary stria abbreviated and the
metasternum sulcate.

The interrelationships among these sections can be analyzed in various ways, depending on
which character states are regarded as derived. We regard the features of the oberthueri
section as being primitive within the subgenus. Although the absence of a metasternal sulcus is
probably a primitive character in the Rhysodini as a whole, it appears to us that a sulcus was
present in the common ancestor of Clinidium sensu stricto, and has been secondarily lost three
times in the guildingii group, in C. rossi, and in the ancestors of the jamaicense and guildingii
sections. All but C. rossi are West Indian species. The metasternal sulcus is also absent in West
Indian species of the insigne and granatense groups. This seems to be an unusual example of
convergent evolution, comparable to the strongly narrowed outer pronotal carinae in members
of various genera and subgenera of Omoglymmiina from the Andaman Islands. An alternative
theory would be that the presence of a median metasternal sulcus is a derived character, which
arose independently in several lines in the Andean Region.

THE OBERTHUERI SECTION

This section contains species with the intercalary and intratubercular striae entire and the
metasternum with a median sulcus. There are seven species, probably occupying two disjunct
areas. Two species are found in Panama. Four others are on the eastern side of the Andes, in
Ecuador, Colombia, Venezuela, and the western part of Amazonas State, Brazil. They
approach the sea only in Merida State, Venezuela, south of Lake Maracaibo. The locality of C.
humboldti is ambiguous, as Nueva Granada included both Colombia and Panama.

Phytogeny — The two Panamanian species, C. alleni and C. whiteheadi, have the basal
impression round, and sharply separated from the discal striole, which is linear. They contrast
strongly with the South American species, and perhaps are not closely related to them. The

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119

latter species have the basal impression small, triangular, and poorly separated from the dilated
discal striole.

C. humboldti is closest to the two Panamanian species, but differs from both in having the
antennal lobes separated from the median lobe. The common ancestor of the Panamanian
species was probably the sister species of C. humboldti.

The four South American species can be grouped into two pairs. C. integrum and C.
pilosum have the paramedian groove more than 0.5 the length of the pronotum and the frontal
groove relatively broad and deep, while in C. oberthueri and C. jolyi the paramedian groove is
less than 0.5 the length of the pronotum and the frontal grooves are relatively shallow and
narrow.

Clinidium ( sensu stricto) integrum Grouvelle 1903
(Figs. 138, 145)

Clinidium integrum Grouvelle 1903: 127-128.

Clinidium ( sensu stricto) integrum (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE female, labelled: “St. Paulo d’Olivenca, M. de Mathan, Mai, 1883” (MNHN).
The type locality is on the Amazon River in western Amazonas State, Brazil, about 100 kilometers east of the Colombian
border.

Description. — Length 6.8 mm. Antennal stylet moderately long, about 0.25 of length of Segment XI; minor setal
tufts on Segments V-X; basal setae present though sparse on Segments VII-X; Segments I-X each with subapical pollinose
band; head distinctly longer than wide; frontal grooves deep, rather wide; median lobe short, triangular, tip opposite
anterior margin of eye; medial margins of temporal lobes curved, closest together near middle of head, divergent
posteriorly; posterior margin rounded, with wide pilose border; eye narrowly crescentic, very short, about 0.33 of length of
temporal lobe; head margins oblique behind eyes; orbital groove complete, pollinose; five or six temporal setae in orbital
groove; two pairs of postlabial setae.

Pronotum rather short, length/greatest width 1.44; widest near middle, base only slightly narrowed, apex more
strongly narrowed; lateral margins only slightly curved; apex truncate; base curved; median groove dilated, widest
posterior to middle, where 0.09 of width of pronotum, slightly narrowed near apex, more suddenly narrowed near base;
median groove not at all dilated at anterior median pit; basal impression narrow, open posteriorly, only moderately wider
than discal striole, latter extended anteriorly 0.9 of length of pronotum; marginal groove visible in dorsal view; angular seta
present; seven to 10 marginal setae; one or two discal setae; sternopleural groove absent; precoxal setae absent.

Elytra elongate; striae impressed, pollinose; all striae entire; apex of intratubercular stria deep; preapical tubercle
inflated, apex angular in posterior view, overhanging preapical impression; apical tubercles inflated, rounded, contiguous;
sutural stria with complete row of five or six setae; preapical impression with three setae aligned with those of sutural stria;
parasutural stria with one or two setae near apex; intercalary stria with complete row of eight or nine setae; intratubercular
stria with three or four setae near apex; marginal stria with complete row of about 10 setae; metasternum with deep
median sulcus; abdominal sterna with transverse sulci widely interrupted in midline; female with deep lateral pit on
Sternum IV; Sternum VI with eight setae; false spurs absent; male unknown.

This species is easily recognized by the great length of the discal striole of the pronotum.
Within the section, the angulate preapical tubercle is also distinctive (Fig. 145).

Vulcano and Pereira (1957b) figured and described a species under this name. They did not
study the type of C. integrum. Judging by their figure, they studied a quite different species,
perhaps referable to Rhyzodiastes , subgenus Rhyzostrix.

Clinidium ( sensu stricto) pilosum Grouvelle 1903
(Figs. 139, 146)

Clinidium pilosum Grouvelle 1903: 126-127.

Clinidium ( sensu stricto) pilosum (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE female, labelled: “Venezuela, Dr. Moritz, 1858, Clinidium pilosum ty.
Grouvelle” (NMW). There was no red “typus” label on the specimen, but the specimen was labelled in Grouvelle’s hand,
and the locality and collector agree with those cited in the original description.

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120

Bell and Bell

Description. — Length 5. 6-6.0 mm. Antennal stylet moderately long, about 0.20 of length of Segment XI; minor
setal tufts on Segments V-X; basal setae present on Segments IX-X; Segments I-VIII with subapical pollinose bands; head
distinctly longer than wide; frontal grooves deep, rather wide; median lobe short, triangular, tip opposite anterior margin of
eye; medial margins of temporal lobes nearly straight, subparallel; posterior margin rounded, with wide pilose border; eye
narrowly crescentic, longer than in C. integrum, over 0.5 of length of temporal lobe; orbital groove complete, pollinose; four
or five temporal setae in orbital groove; two pairs of postlabial setae.

Pronotum longer than in C. integrum-, length/greatest width 1.65; widest near middle, sides nearly parallel, only
slightly curved; apex truncate-, base curved; median groove narrow, margins nearly parallel except for dilation opposite
middle of discal stride and another at base; median groove not at all dilated opposite anterior median pit; basal impression
narrow, closed posteriorly but open laterally, only slightly wider than discal striole; latter extended anteriorly about 0.66 of
length of pronotum; marginal groove visible in dorsal view; angular seta present; six to 10 marginal setae; one or two
discals on either side anteriorly, one or two on each side in basal 0.25 of length or absent; sternopleural groove absent;
precoxal setae absent.

Elytra elongate, striae impressed, pollinose, coarsely punctate; all striae entire; apex of intratubercular stria rather
shallow; preapical tubercle inflated, rounded; apical tubercles inflated, contiguous; sutural stria with complete row of five
setae, apical impression with two aligned with those of sutural stria; parasutural stria with one seta near base; intercalary
stria with complete row of nine or 10 setae; intratubercular stria with four setae near apex; marginal stria with about 18
setae in complete row; apical tubercle with about six setae; metasternum with shallow median sulcus; transverse sulci of
abdominal Sternum III not interrupted in midline; that of Sternum IV not interrupted in male, interrupted in female; those
of Sterna V-VI broadly interrupted; Sternum VI with submarginal groove bent inward at base, not connected to transverse
sulci (Fig. 146); Sternum VI with six to eight setae, two to four in transverse row between anterior ends of submarginal
sulcus, four in curving row along submarginal sulcus; female with lateral pit in Sternum IV; tibiae with false spurs present,
though small; male without ventral tooth on anterior femur, nor proximal tooth on anterior tibia; calcars triangular; middle
one narrow, longer, more pointed than hind one; latter with dorsal margin slightly angulate.

The very long, nearly parallel-sided pronotum is distinctive. C. jolyi differs in having a
shorter pronotum, with the paramedian grooves less than 0.5 the length of the pronotum. Also,
the parasutural stria has a complete row. of setae, the submarginal groove of the sixth sternite is
absent and the frontal grooves much shallower. C. oberthueri differs in having a much more
oval pronotum, strongly narrowed at both base and apex. The eye is shorter, the frontal grooves
are shallower, and precoxal setae are present.

Range. — Merida State, in western Venezuela, and possibly in adjacent parts of Colombia.

In addition to the holotype, we have seen one male and two females, labelled: “Venezuela, Merida,
La Azulita, 2000 m., 5,6-X-69, J. and B. Bechyne leg.” (VEN), and a male, with a handwritten label which is difficult to
interpret, but which appears to us to read “Cae Lun, N., Columb., Mor. 8129” (MNHB).

Clinidium ( sensu stricto) jolyi new species
(Figs. 140, 147)

Type Material. — HOLOTYPE male, labelled: “VENEZUELA, Merida, La Azulita, 2000 m„ 6-X-69, J. & B.
Bechyne, leg.” (VEN). PARATYPES two males, labelled: “Venezuela, Merida, La Mucuy, 30-VIII- 1 956, C. J. Rosales
col.” (VEN); one male, one female, labelled: “Venezuela, Merida, Carbonera, 2600 m, 3-X-69, J. & B. Bechyne” (VEN).

Description. — Length 5.0-6.0 mm. Antennal stylet rather small, about 0.16 of length of Segment XI; minor setal
tufts on Segments V-X; basal setae present on Segments IX, X; Segments I-IV with subapical pollinose bands; head
slightly longer than wide; frontal grooves very shallow, linear, glabrous; median lobe short, triangular, ending just posterior
to anterior margin of eye; medial margins of temporal lobes slightly divergent posteriorly; posterior margin transverse;
posterior margin with very broad pilose border, this extending anteriorly along medial margin; eye narrowly crescentic,
shorter than in C. pilosum, about 0.4 of length of temporal lobe; orbital groove complete, pollinose; three temporal setae,
two opposite eye, the other behind eye, distant from margin, its base included in dilation of marginal pilose band; two
postlabial setae.

Pronotum shorter than in C. pilosum-, length/greatest width about 1.48; widest near middle, sides curved; base, apex
nearly equally narrowed; apex truncate; base curved; median groove narrow, margins nearly parallel; groove scarcely
dilated opposite anteriomedian pit; basal impression narrow, closed posteriorly, open laterally; discal striole relatively wide,
extending to middle of pronotum; marginal groove visible in dorsal view; angular seta present; three or four marginal setae,
mostly anterior to middle; one or two anterior discal setae; no posterior discals; sternopleural groove absent; precoxal setae
absent.

Elytra elongate; striae impressed, pollinose, coarsely punctate; apex of intratubercular stria rather shallow; preapical
tubercle inflated, rounded; apical tubercles inflated, contiguous; sutural striole with complete row of five to seven setae;

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

121

parasutural stria with complete row of five or six setae; intercalary stria with about 13 setae; intratubercular stria with
about five setae in apical 0.33; marginal stria with complete row of about 15 setae; apical tubercle with two setae;
metasternum with very shallow median groove; Sterna II-VI with shallow transverse sulci, each containing row of coarse
punctures; transverse sulci broadly interrupted in midline; Sternum VI without submarginal sulcus; four setae near
posterior margin of Sternum VI (Fig. 147); in most specimens two setae near middle of Sternum VI, one specimen with
three, another with four; female with deep lateral pit on Sternum IV, shallower one on Sternum V; false spur present, 0.67
as long as true spurs; male without ventral tooth on anterior femur, without proximal tooth on anterior tibia; calcars short,
broadly triangular, hind one deeper but not longer than middle one; dorsal margins of calcars not angulate.

The very shallow frontal grooves on the head and the absence of the submarginal sulcus of
Sternum VI are characteristic of this species. The shorter, more rounded pronotum with shorter
discal strides also separates it from the sympatric C. pilosum. The complete series of setae in
the parasutural stria also differentiates the type series from the latter species. However, this
character is lacking in a specimen from Trujillo State, which may be a variant of C. jolyi (see
below, under “variation”).

Range and variation. — The type series is known only from Merida State. We have studied
a female specimen, labelled: “Venezuela-Trujillo, La Pena, 3000 m., 5-IX-1968, J. & B.
Bechyne” (VEN). This specimen is closer to C. jolyi in most respects, including the shape of
the pronotum and the sculpture and chaetotaxy of Sternum VI. However, the elytral striae are
distinctly finer, and the chaetotaxy of the pronotum and elytra differ in important respects:
four temporal setae, six to eight marginal setae on the pronotum; parasutural stria without
setae, and pilose apical bands only on antennal Segment I. This specimen might be an extreme
variant of C. jolyi , although we suspect it is more likely to be a closely allied but distinct form.
A final decision will need to await the collection of more specimens.

Clinidium ( sensu stricto) oberthueri Grouvelle 1903
(Fig. 141)

Clinidium oberthueri Grouvelle 1903: 128-129.

Clinidium ( sensu stricto ) oberthueri (Grouvelle) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) male, labelled: “Ecuador, coll. Slemiradsky 1882-1883 ”
(MNHN). PARALECTOTYPES, seven specimens, sexes not recorded, same label as lectotype (MNHN); one male, one
female, same data as lectotype, labelled “Co-type” (BMNH).

Description. — Length 6. 0-6. 3 mm. Antennal stylet slender, acuminate about 0.20 of length of Segment XI; latter
long, conical; minor setal tufts on Segments V-X; basal setae present, but very sparse, on Segments VI-X; Segment 1 with
broad subapical band of pollinosity; antenna otherwise without pollinosity; head distinctly longer than wide; frontal
grooves narrow, shallow, margins inconspicuously pollinose; median lobe short, triangular, tip slightly behind anterior
margin of eye; medial margins of temporal lobes slightly divergent posteriorly; posterior margin rounded, with narrow
pollinose margin; eye narrowly crescentic, short, about 0.25 of length of temporal lobe; orbital groove complete, pollinose;
lateral margin of head oblique posterior to eye; four temporal setae in orbital groove; two pairs of postlabial setae.

Pronotum elongate, length/greatest width 1.67, oval, base, apex strongly narrowed, lateral margin strongly curved,
base nearly evenly rounded; apex narrowly truncate; median groove narrow, margins nearly parallel, dilated very slightly
opposite anterior median pit; basal impression very narrow, closed posteriorly, open laterally; impression only slightly
wider than discal striole, latter extending nearly to middle of pronotum; marginal groove visible in dorsal view; angular
seta present; seven or eight marginals; one anterior discal or absent; notopleural suture dilated, pollinose near middle;
sternopleural groove present; precoxal seta present.

Elytra moderately elongate, humeral region very strongly narrowed compared to related species; striae impressed,
pollinose, coarsely punctate; apex of intratubercular stria rather shallow; all striae entire; preapical tubercle strongly
inflated, rounded; apical tubercles weakly inflated, contiguous; sutural striae with complete row of four or five setae;
parasutural stria without setae; intercalary stria with complete row of five to setae setae; intratubercular stria with four or
five setae in apical 0.20; marginal stria with 10-13 setae in complete row; preapical tubercle with four or five setae; apical
tubercle with two setae; metasternum with deep median sulcus; transverse sulcus of Sternum III continuous in both sexes;
that of Sternum IV narrowly interrupted in both sexes; V narrowly interrupted in male, widely interrupted in female; that
of VI widely interrupted in both sexes; submarginal groove short, broadly separated from transverse sulci; female with
eight setae on Sternum VI, four in curved lines near hind margin, four in transverse line near middle; male with six setae,
inner pair of transverse line absent; female with very large lateral pit on Sternum IV; false spur absent, replaced by obtuse

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Bell and Bell

angle; male without ventral tooth on anterior femur or proximal tooth on anterior tibia; middle calcar broadly triangular,
dorsal margin straight; hind calcar larger, its apex obtuse, dorsal margin angulate.

The oval pronotum and very narrow elytral humeri of this species are distinctive. The precoxal setae also differentiate
it from all members of the section except for C. alleni. The latter species has a shorter pronotum with more parallel
margins, and linear discal strides.

Range. — Ecuador. The only specimens with a definite locality are three labelled:
“Papallacta, Napo-Pastaza Prov., 30 January 1958, R. W. Hodges, 10500 ft. elev.” (MSU;
UVM). This is on the eastern side of the Andes.

Clinidium ( sensu stricto) alleni new species
(Figs. 142, 148, 149)

Type Material. — HOLOTYPE male, labelled: “Panama, Cerro Jefe, 9° 12' N-79° 21'W, 700-750 m„ May 20,
1972, beating and under bark. R. T. Allen, ADP 11544” (NMNH). This locality is in the Cordillera de San Bias, on the
eastern, or South American side of the Panama Canal.

Description. — Length 6.4 mm. Antennal stylet very slender, inconspicuous, about 0.2 of length of Segment XI;
latter short, nearly spherical; minor setal tufts on Segments V-X; basal setae entirely absent; Antennal Segments I-VI with
pollinose subapical bands; more distal segments with pollinosity limited to bases of setae, forming broken bands; head 1.5
times longer than wide; frontal grooves narrow, deep, partly pollinose; median lobe short, triangular; joined to antennal
lobe, tip opposite anterior margin of eye; medial margins of temporal lobes divergent posteriorly; posterior margin
narrowly rounded; posterior and posteriomedial margins broadly bordered with pilosity, so glabrous part of temporal lobe
is strongly narrowed posteriorly; eye narrowly crescentic, about 0.33 of length of temporal lobe; orbital groove complete,
pollinose; lateral margin of head nearly straight posterior to eye; three temporal setae in orbital groove; two pairs of
postlabial setae.

Pronotum rather short, length/greatest width 1.40, widest just posterior to middle; base, apex moderately narrowed;
lateral margin curved, apex truncate; base rounded; median groove narrow, margins pollinose, anterior median pit very
small, but distinctly wider than median groove; basal 0.3 of median groove shallow, linear, glabrous; basal impression very
small, deep, punctiform, sharply distinct from discal striole, latter linear, slightly curved, extending slightly anterior to
middle of pronotum; marginal groove fine, visible in dorsal view; angular seta present; nine or 10 marginal setae; one or
two anterior discal setae; sternopleural groove absent; precoxal setae present.

Elytra rather short; humeri much less narrowed than in C. oberthuerr, intratubercular stria impressed only at base,
apex, in middle represented only by row of punctures (Fig. 149); remaining striae impressed, entire, pollinose, coarsely
punctured; Intervals II, III forming prominent swelling just posterior to base of elytron (this asymmetrical, and possibly
the result of an injury); preapical tubercle inflated, rounded; apical tubercles inflated, rounded, slightly separated; sutural
stria with complete row of five setae; parasutural stria with one or two setae near base; intercalary stria with complete row
of six or seven setae; intratubercular stria with two or three setae near apex; marginal stria with complete row of about 1 2
setae; preapical tubercle without setae; apical tubercle with three or four setae; metasternum with very shallow median
sulcus; transverse sulci of all abdominal sterna interrupted in midline, submarginal sulcus of Sternum VI widely separated
from transverse sulci (Fig. 148); Sternum VI with four setae in submarginal row; one or two on each side in transverse row;
middle, hind tibiae each with false spur; male without ventral tooth on anterior femur, without proximal tooth on anterior
tibia; calcars triangular, middle one narrow, with dorsal margin straight; hind one broader, dorsal margin nearly straight.
Female unknown.

This species resembles C. oberthueri in having precoxal setae, but differs from the latter in
having a shorter, less rounded pronotum, elytral humeri much less narrowed, and discal strides
and frontal grooves much shallower and narrower. C. whiteheadi is a similar species, found
nearby, but to the west of the Panama Canal. It lacks the precoxal setae, has basal setae on the
outer antennal segments, and has straight discal strioles and a more parallel-sided pronotum.

We take pleasure in naming this species for Dr. R. T. Allen, whose collections have helped
greatly in making known the beetle fauna of lower Central America.

Clinidium ( sensu stricto) whiteheadi new species
(Fig. 143)

Type Material. — HOLOTYPE male, labelled: “PANAMA: Panama, Cerro Campana 8° 40' N, 79° 56' W, 29
June 74, 850 ms; T. L. Erwin, D. R. Whitehead, under loose bark of log; Exped #1 23, notebook #3, ADP 25285”

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123

(NMNH). This locality is to the west of the Panama Canal, towards Central and North America. PARATYPES two
males, three females, same label as holotype (NMNH); one female, same locality, collected by T. L. Erwin and J. L.
Lawrence, 22 Feb. 1975 (in heartwood) (NMNH); one male, one female, same locality as holotype, labelled: “19-VIII-78,
ex Stemonitis, Q. D. Wheeler 7867” (CU).

Description. — Length 5.0-5. 8 mm. Antennal stylet minute, less than 0.1 of length of Segment XI; latter only
slightly longer than wide; minor setal tufts on Segments V-X; basal setae present on Segments VIII-X; Segments I-IV with
subapical pollinose bands; head slightly longer than wide, less elongate than in C. alleni; frontal grooves narrow but deep,
pollinose; median lobe short, triangular,joined to antennal lobe; tip opposite anterior 0.25 of eye; medial margins of
temporal lobes divergent posteriorly; posterior margin more broadly rounded than in C. alleni, posterior, posteriomedial
margins broadly bordered with pilosity; eye narrowly crescentic. 0.33 of length of temporal lobe; orbital groove complete,
pollinose; lateral margin of head nearly straight posterior to eye; two or three temporal setae in orbital groove: two pairs of
postlabial setae.

Pronotum rather short, length/greatest width 1.54, widest near middle, lateral margins nearly straight; base, apex less
narrowed than in C. alleni', apex truncate; base rounded; median groove narrow, though broader in C. alleni, widest in
middle 0.33, slightly constricted near apex; not broadened at anterior median pit; posterior 0.33 very shallow, finely
pollinose; basal impression very small, deep, punctiform, sharply distinct from discal striole; latter linear, straight,
extending to middle of pronotum; marginal groove fine, visible in dorsal view; angular seta present; seven to nine
marginals; one or two anterior discal setae; notopleural suture glabrous; sternopleural groove absent; precoxal setae absent.

Elytra rather short, humeri only slightly narrowed; intervals less convex than in C. alleni', intratubercular stria
impressed only at base, apex; middle part, a row of fine punctures; remaining striae impressed, entire, pollinose, coarsely
punctured; no swelling in Intervals II, III near base; preapical tubercle scarcely inflated, rounded; apical tubercles scarcely
inflated, rounded, slightly separated; sutural stria without setae in most specimens, with one seta on one elytron in one
female specimen; parasutural striae with one seta near base; intercalary stria with complete row of six or seven setae;
intratubercular stria with three setae near apex; marginal stria with complete row of about 10 setae; preapical tubercle
without setae; apical tubercle with two or three setae; metasternum with shallow median sulcus; transverse sulci of
abdominal Sterna II- VI widely interrupted in midline; submarginal sulcus of Sternum VI widely separated from transverse
sulci; Sternum VI with six to eight setae, four in row along submarginal sulcus, two on disc, in one male, with four on disc,
lateral ones more anterior than medial ones; female with large lateral pit in Sternum IV; middle, hind tibiae each with
large false spur; male without ventral tooth on anterior femur nor proximal tooth on anterior tibia; calcars triangular,
dorsal margins slightly curved; penis of holotype mounted separately on point, distal 0.5 straight, apex abruptly deflexed.

This species has linear, straight discal strides and a parallel-sided pronotum. This and the
absence of precoxal setae separate it from C. alleni. C. dormans, another similar Panamanian
species, has the apex of the intratubercular stria not impressed, so that preapical and apical
tubercles are not separate.

We take pleasure in naming this species for Dr. Whitehead, one of the ablest and most
productive of the students of Latin American beetles.

Bionomics. — The specimens were collected by Mr. Wheeler (in. litt .) in the fruiting bodies
of the slime mold Stemonitis. To our knowledge, this is the first record of a rhysodine in a
fruiting body, and the first linkage of a particular species of rhysodine with a particular genus
of slime mold.

Clinidium ( sensu stricto) humboldti new species
(Figs. 144, 150)

Type Material. — HOLOTYPE female, labelled: “nov. Granad., 43693 (MNHB).

Description. — Length 6.4 mm. Antennal stylet about 0.25 of length of Segment XI, larger than in related species;
Segment XI distinctly longer than wide; tufts of minor setae on Segments V-X; basal setae present on Segments VI-X;
Segments I-VI with subapical pollinose bands; head distinctly longer than wide; frontal grooves rather broad, moderately
deep, glabrous except for medial margins; median lobe short, triangular, tip opposite anterior 0.25 of eye; median lobe
separated from antennal lobe by frontoclypeal groove; medial margins of temporal lobes slightly divergent posteriorly;
glabrous area of temporal lobe oval, widest posterior to eye; posterior, posteriomedial margins broadly bordered by
pollinosity; eye narrowly crescentic, elongate, 0.67 of length of temporal lobe; orbital groove complete, broadly pollinose; 4
temporal setae; two pairs of postlabial setae.

Pronotum rather short, length/greatest width 1.47, widest near middle, lateral margins strongly curved; base, apex
strongly narrowed; apex truncate; base rounded; median groove slightly dilated; 0.75 deep, posterior 0.25 shallow; median
groove not widened at anterior median pit; latter far posterior to pronotal apex; basal impressions small, deep, sharply

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Bell and Bell

distinct from discal stride; latter linear, slightly curved, extending to middle of pronotum; basal impression broadly joined
laterally to marginal groove; latter fine, visible in dorsal groove; angular seta present; seven or eight marginal setae; one
pair of anterior discal setae; notopleural suture glabrous; sternopleural groove absent; precoxal setae present.

Elytra rather short, humeri scarcely narrowed; elytral intervals nearly flat; intratubercular stria impressed at apex;
remainder scarcely impressed, represented by row of fine punctures; remaining striae impressed, entire, pollinose, coarsely
punctate; no swelling on Intervals II, III near base; preapical tubercle strongly inflated, medial margins sinuate, apex
subtruncate (Fig. 150); apical tubercle inflated; sutural stria with three or four setae in posterior 0.67; parasutural stria
without setae; intercalary stria with complete row of nine or 10 setae; intratubercular stria with four or five setae near
apex; marginal stria with 11-12 setae; preapical tubercle with one seta on medial margin; apical tubercle with three or four
setae; metasternum with very shallow glabrous median impression, latter containing two elongate pits, near anterior,
posterior margins, respectively; transverse sulci of Sterna III- VI linear, widely separated in midline, those of Sternum V
slightly oblique; female with lateral pit on Sternum IV; transverse sulci of Sternum VI widely separated from submarginal
sulcus; Sternum VI with eight setae; four in transverse row anterior to submarginal sulcus, four near posterior margin;
middle, hind tibiae with short, triangular cusp, which does not resemble a spur. Male unknown.

This species resembles C. whiteheadi and C. alleni in appearance. It differs from both in
having the median lobe not connected to the antennal lobes. It differs from C. alleni in the
presence of basal setae on the antennae, in the shape of the temporal lobes, and in the presence
of anterior discal setae. C. whiteheadi lacks precoxal setae, and has the pronotum more
elongate, with the margins less curved.

We dedicate this species to Alexander von Humboldt and to the museum named for him, in
gratitude for the loan of their valuable collection of Rhysodini.

THE JAMAICENSE SECTION

Like the preceding section, this one contains species with both intercalary and
intratubercular striae with apices complete. However, there is no trace of a median sulcus on
the metasternum. There are five species, two in Jamaica, and three in Hispaniola.

Phytogeny. — C. trionyx of the Dominican Republic contrasts strongly with the four
remaining species. It has precoxal setae and false spurs, both probably plesiomorphic
characters, in which it resembles some members of the oberthueri section. The eye is reduced to
a small vestige, the discal strides are absent, and the intratubercular stria is virtually absent
except for its impressed apex. These are specialized features in comparison to the character
states in the remaining species. The latter are very close to one another, and can be regarded as
a species complex.

In the jamaicense complex, the two Haitian species clearly form one unit, and the two
Jamaican ones, another unit.

Clinidium ( sensu stricto) trionyx new species
(Figs. 151, 160, 161)

Type Material. — HOLOTYPE male, labelled: “Rep. Dominic, J. & S. Klapperich, Cazabita 1250 m. 30-VI-74”
(BSL).

Description. — Length 6.0 mm. Antennal stylet minute; minor setal tufts on Segments V-X; basal setae absent;
antennal Segments I-IV with subapical pollinose bands; head slightly longer than wide; frontal grooves narrow, shallow,
glabrous; median lobe triangular, rather long, tip opposite posterior margin of eye; medial margins of temporal lobes
slightly convergent posteriorly; posterior margin rounded; posterior, posteriomedial margins bordered by pollinosity; eye
minute, oblong, 0.2 of length of temporal lobe, eye 2.5 longer than deep (Fig. 161); orbital groove complete, pollinose;
lateral margin of head slightly oblique posterior to eye; 3 temporal setae in orbital groove; one pair of postlabial setae.

Pronotum rather short, length/greatest width 1.47; widest just posterior to middle, oval, lateral margins curved; base
curved; apex narrowly truncate; median groove narrow, margins pollinose, groove slightly dilated in middle 0.33; groove
not at all dilated at anterior median pit; basal 0.33 of median groove shallow, pollinose; basal impression small, deep,
triangular, closed posteriorly; discal stride absent; marginal groove fine, visible in dorsal view; angular seta present; five or
six marginals, one pair of anterior discal setae; sternopleural groove absent; precoxal setae present.

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125

Elytra relatively short, broad; three inner striae impressed, pollinose, entire; intratubercular stria with apex impressed,
separating preapical from apical tubercle, remainder of intratubercular stria nearly absent, represented by faint impression
visible by oblique lighting, without punctures or pollinosity; marginal stria entire, impressed, pollinose; preapical tubercle
only slightly inflated; apical tubercles inflated, contiguous, meeting in straight median suture; sutural striae with complete
row of five setae (most posterior one in apical impression just posterior to apex of sutural stria); parasutural stria with one
seta at base; intercalary stria with complete row of six setae; apical impressed part of intratubercular stria with three setae,
anteriormost one in conspicuous pollinose puncture; marginal stria with one seta near humerus, six or seven setae in apical
0.67, three or four in conspicuous punctures; metasternum without median sulcus; transverse sulci of abdominal Sterna
III-VI widely separated in midline, also from margin, forming pairs of oval impressions, most with one or two small
punctures medial to them; Sternum VI with submarginal groove widely separated from transverse sulci (Fig. 160);
Sternum VI with four pairs of setiferous punctures, anterior ones in recurved row, posterior ones in procurved row along
submarginal groove; middle, hind tibiae each with false spur; male without ventral tooth on anterior femur nor proximal
tooth on anterior tibia; middle calcar narrowly triangular, close to spurs; hind calcar more broadly triangular, raised above
level of spurs.

This species differs from the two Haitian members of the section in having slightly
convergent temporal lobes, much more finely punctate striae, the intratubercular stria almost
absent except for the impressed apex, a much smaller eye, precoxal setae and false spurs
present. In the oberthueri group, it is closest to C. alleni, but differs in having only one pair of
postlabial setae, metasternum not at all sulcate, and the discal stride absent.

THE J A M A ICE NS E COMPLEX

This includes C. haitiense, C. corbis, C. jamaicense and C. chiolinoi , in short, all members
of the jamaicense section, excepting C. trionyx. The species are so similar that it is convenient
to present a description for the complex before listing the distinctive features of each species.

Description. — Antennal stylet minute; minor setal tufts on Segments V-X; head distinctly longer than wide;
orbital groove complete; eye narrowly crescentic, 0.5-0.67 of length of temporal lobe; cornea clear in some (younger?)
specimens, completely darkly pigmented in other specimens; three temporal setae in orbital groove in most specimens, in
some, one side has two or four setae; one or two pairs of postlabial setae, pronotum rather short, widest in middle, lateral
margins curved, base rounded; apex truncate; median groove narrow, oval, pollinose, not at all dilated opposite anterior
median pit; basal impression small, deep, triangular; closed posteriorly; discal stride varying among the species; marginal
groove Fine, visible in dorsal view; angular seta present; marginals two to six; one to three anterior discal setae;
sternopleural groove, precoxal setae absent.

Elytra shorter, broader than in C. trionyx ; striae coarsely punctured; inner three striae deeply impressed, pollinose,
entire; intratubercular stria with apex impressed, remainder very shallowly impressed, but with coarse, conspicuous row of
punctures; marginal stria entire, impressed; preapical tubercle scarcely inflated; apical tubercles slightly inflated,
contiguous, meeting in straight median suture; metasternum not sulcate; transverse sulci moderately separated in midline,
reaching to lateral margins of abdominal sterna; Sternum VI with submarginal sulcus widely separated from transverse
sulci; Sternum VI with six to eight setiferous punctures; middle, hind tibiae without false spurs; male without ventral tooth
on anterior femur nor proximal tooth on anterior tibia.

These four species differ strongly from C. trionyx in the absence of false spurs and precoxal
setae, and in having the intratubercular stria coarsely punctate.

Clinidium ( sensu stricto ) haitiense Bell 1970
(Fig. 152)

Clinidium ( sensu stricto) haitiense Bell 1970: 322.

Type Material. — HOLOTYPE male, labelled: “La Visite, La Selle Range, Haiti, 5000-7000 ft., coll. P. J.
Darlington, Sept. 16-23, 1934” (MCZ 31755). PARATYPES one male, two females, same label as type (MCZ; UVM).

Description. — Length 5. 7-6.4 mm. Antennae very thick; basal setae present on Segments VII-X or VIII-X;
Segments I-II with complete subapical pollinose rings; Segments III, IV with rings interrupted; outer segments without
pollinosity; median lobe of head long connected laterally to antennal lobes, tip of medial lobe opposite posterior margin of
eye; frontal grooves deep, narrow, medial, lateral margins both sharp; margins of frontal grooves not pollinose; occipital
pilosity short; orbital groove very narrow; head lateral to orbital groove behind eye glabrous; labium not pollinose medially.

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Pronotum relatively elongate; length/greatest width 1.52; apex of pronotum more narrowed anteriorly than in C.
corbis ; discal setae in conspicuous punctures; in most specimens, two pairs of discals, forming rectangle, in one specimen
one on one side, in another, three on one side; three to seven marginal setae; basal impression very narrow; discal stride in
most specimens extending to posterior 0.25 of pronotum, but in one specimen, almost absent; pollinosity of discal striole
much reduced.

Elytra relatively elongate with humeri slightly narrowed; Interval I slightly less convex than Intervals II, III; marginal
coarsely punctate; sutural stria with three or four setae, mostly posterior to middle; parasutural without setae; intercalary
with complete row of five to seven, intratubercular with two or three near apex; marginal stria with nine or 10 setae, one at
humerus, others in posterior 0.5; apical tubercle with two or three setae; anterior femur with small pollinose spot at base of
each seta; legs relatively long, slender; calcars very small; hind calcar slightly obtuse, lower margin evenly curved.

This species is larger and more elongate than the closely related C. corbis, also from Haiti.
It has the pilosity much reduced, with the dorsal surface of the femora with isolated pollinose
spots at the bases of the setae, and the side of the head laterad to the orbital groove glabrous.

Range. — Probably restricted to high elevations in the Morne La Selle, south of
Port-au-Prince. Bell (1970) recorded it from Furcy and Mandeville in addition to the type
locality.

Clinidium ( sensu stricto ) corbis Bell 1970
(Fig. 153)

Clinidium ( sensu stricto) corbis Bell 1970: 322-323.

Type Material. — HOLOTYPE male, labelled: “Tardieu, Morne La Hotte, Haiti, 3000 ft., coll. P. J. Darlington,
October 14, 1934” (MCZ 31754). PARATYPES one female, same label as male (MCZ); one male, labelled: “Roche
Croix, 5000 ft., Morne La Hotte, coll. P. J. Darlington, Oct. 13, 1934” (MCZ); two males labelled: “northeast foothills,
Morne La Hotte, 2000-4000 ft., coll. P. J. Darlington, Oct. 10-24, 1934” (MCZ).

Description. — Length 4.4-5. 5 mm. Antennae less thick than in C. haitiense, with the segments more nearly round;
basal setae present, though sparse on Segments IX, X; Segments I- VIII in most specimens with subapical band of
pollinosity; in a few specimens, pollinosity restricted to I-VI or I-VII; median lobe of head long, separated laterally from
antennal lobes; tip of median lobe opposite posterior margin of eye; frontal grooves deep, narrow, margins pollinose;
medial, lateral margins equally pollinose; orbital groove relatively broad; occipital pilosity prominent; head laterad to
orbital groove posterior to eye pilose; labium with median pollinose band.

Pronotum relatively elongate, length/greatest width about 1.51; discal setae in smaller punctures than in C. haitiense ,
one or two pairs, varying geographically; four to six marginal setae; basal impression very narrow; discal striole longer,
more pollinose than in C. haitiense , 0.30-0.50 of length of pronotum except in specimens from Dajabon, P.R.

Elytra in most specimens shorter than those of C. haitiense, Interval I slightly less convex than Interval II; marginal
stria coarsely punctate; sutural stria with two or three setae posterior to middle; parasutural stria in most specimens
without setae, in a few, with one seta at base; intercalary stria with complete row of five or six setae; intratubercular stria
with one to three setae near apex; marginal stria with five- 10 setae, including one at humerus; apical tubercle with two or
three setae; anterior femur with dorsal pollinose stripe containing most of the setal bases; legs shorter, thicker than in C.
haitiense-, calcars slightly larger than in C. haitiense-, hind calcar acute, ventral margin abruptly bent.

This species is most easily separated from C. haitiense by the long pilosity laterad to the
posterior part of the orbital groove.

Range. — This species is probably not confined to the Morne La Hotte, as supposed by Bell
(1970). Specimens apparently belonging to this species have been found in the following

additional localities: HAITI, Lebrun, near Miragoane, coll. R. T. Bell, R. Sette, over fifty specimens (UVM); Morne
Grand Bois, 3780', coll. M. Langworthy, five specimens (UVM); Catiche, 2700', coll. M. Langworthy, one specimen
(UVM); lie de la Tortue, Aux Basin, w. of Aux Palmiste, coll. M. Langworthy, T. Dowhan (UVM). All localities except
the last are from the Southern Peninsula: DOMINICAN REPUBLIC: Dajabon, Mariano Cestero, 650 m. 1 2-VIII- 1 980,
A. Norrbom (CMP), two specimens. Probably this species was formerly throughout Hispaniola at low and medium
elevations. One male and two females labelled:“WM 5958, Cinnamon Bay, June 6, 1980, between buttresses of large
kapok tree” appear to represent this species, although they are unusually large, 5.0-5.6 mm. This locality is in the
(American) Virgin Islands, on Saint John. They may represent an introduction by human agency, or a very recent natural
invasion. The latter seems unlikely, in view of the apparent absence of the species from Puerto Rico. The specimens were
collected by William Muchmore and sent to us by Kenneth Cooper.

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127

Variation. — On the basis of the limited material available, the species appears to vary
geographically, and might represent more than one taxon. In the type series, from Morne La
Hotte, three have the discal setae on the pronotum 1-1 (one on each side), while two have them
1-2. The long series from Lebrun mostly have more discal setae. They are distributed as
follows:

Discal setae

Number of s

1-1

1-2

2-2

2-3

3-3

2-1

3-1

In addition, the series from Lebrun appear to have the lateral margins consistently
straighter and more parallel than specimens from other regions. The calcars appear identical to
those from Morne La Hotte.

The series from lie de la Tortue resemble those from Morne La Hotte in the shape of the
pronotum and in having few discal setae. These are distributed as follows:

0- 1 1 specimen

1- 1 5

1-2 6

The hind calcar is scarcely raised above the level of the spurs.

The specimens from Dajabon, Dominican Republic, have the calcars as in the series from
Morne La Hotte, which they also resemble in the shape of the pronotum. The discal setae are
1-1 and 0-1. The discal strioles are much shorter than in other specimens of C. corbis , being
scarcely longer than those of C. haitiense.

The specimens from Saint John have three or four discal setae, and have the calcar strongly
raised above the tibial spurs. The length of the body is greater than those of the other series.

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128

Bell and Bell

The differences among these populations may represent subspecific characters. However,
they might be merely the result of individual variation. The large series from Lebrun were
almost entirely from a single log. Two individuals were from a smaller piece of wood only a few
meters away. Thus, the entire series could represent the offspring of a single mating. The same
is true of the series from lie de La Tortue.

Bionomics. — The series from Lebrun were mostly in small rotten areas in a large, dry, hard
log. The local name of the tree is “marron”, but we were unable to find its scientific name. Two
of the beetles were in a stick of Cecropia located a few meters away.

The series from lie de La Tortue were in an unusual habitat, within a log in a sink hole in a
large cave system.

Clinidium (sensu stricto) jamaicense Arrow 1942
(Figs. 154, 158)

Clinidium jamaicense Arrow 1942:181.

Clinidium ( sensu stricto) jamaicense (Arrow) Bell and Bell 1978.

Type Material. — LECTOTYPE (here designated) male, labelled: “Jamaica, Dr. M. Cameron, BM- 1936-555”
(BMNH). According to the original description, the type series was collected at Newcastle, Jamaica, under rotting bark.
PARALECTOTYPES one male, two females, same label as lectotype. (BMNH).

Description. — Length 4. 6-5. 7 mm. Antenna with basal setae very few, confined to Segment X, or IX, X;
Segments I-IV with subapical bands of pollinosity; median lobe of head short, blunt, opposite anterior 0.25 of eye; frontal
grooves very shallow, glabrous; lateral margin of frontal groove ill-defined; occipital pilosity long; orbital groove broadly
pollinose; labium pollinose medially.

Pronotum slightly shorter than in Haitian species, length/greatest width about 1.48; discal setae in most specimens one
on each side, a few specimens with 2-1 or 0-1; marginal setae two or three, in most specimens with two near anterior angle,
one near middle or absent; basal impression triangular, discal striole very short, shorter than basal impression, in many
specimens obsolete.

Elytra relatively short, broad; Interval I narrow, convex, not broadened posteriorly; sutural stria deeply impressed,
medial border as distinct as lateral border; parasutural stria deeply impressed, coarsely punctate; Intervals I-III of nearly
equal height, convexity; intercalary stria deeply impressed; intratubercular stria with apex impressed, remainder a row of
coarse punctures; marginal stria complete in all specimens; sutural stria with two or three setae near apex; parasutural
stria without setae; intercalary stria with two to five setae, when numerous forming complete row; intratubercular stria
with two or three setae in impressed apex; marginal stria with six or fewer setae, most anterior one at humerus; apical
tubercle with two or three setae; calcars very small.

The shallow frontal grooves and short medial lobe distinguish this species from the two
Haitian ones, while the convex sutural interval separates it from C. chiolinoi.

Range. — Mountains of Jamaica above 2000 ft. elevation. Bell (1970) recorded it from
Portland Gap, Cinchona, Hardwar Gap, Blue Mountain Peak, Whitfield Hall, and Belmore
Castle.

Clinidium ( sensu stricto) chiolinoi Bell 1970
(Figs. 155, 159)

Clinidium ( sensu stricto) chiolinoi 1970: 323-324.

Type Material. — HOLOTYPE male, labelled: “Mount Diablo, St. Ann Parish, Jamaica, coll. R. T., J. R. Bell,
B. B. Chiolino, Jan. 2, 1967” (MCZ). PARATYPES two males, five females, same label as type (MCZ; UVM).

Description. — Length 4. 4-5.6 mm. Antenna with basal setae on VI-X, VII-X, or VIII-X, sparse, mostly lateral;
all antennal segments with subapical pollinose bands, those of distal segments very narrow; median lobe of head short,
blunt, tip opposite anterior end of eye; frontal grooves very shallow, very finely margined with inconspicuous pollinosity;
lateral margin of frontal groove ill-defined; occipital pilosity long; orbital groove broadly pollinose; labium pollinose
medially.

Pronotum proportion as in C. jamaicense, length/greatest width 1.48; discal setae in most specimens one on each side,
in a few specimens 2-1 or 1-0; marginal setae 2-3, in most specimens one or two near anterior angle, one near middle;

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

129

angular seta present or absent; basal impression triangular; discal stride very short, shorter than basal impression, in many
specimens obsolete.

Elytra relatively short, broad, Interval I broad, flat, depressed below level of other intervals, wider posteriorly; sutural
stria deep, punctate, medial margin lower than lateral margin; parasutural stria deeply impressed, punctate; intercalary
striae shallowly impressed; Interval III more convex, higher than other intervals; intratubercular stria with apex
impressed, remainder row of very fine punctures, in many specimens almost absent anteriorly; marginal stria entire
(western specimens) or interrupted near humerus (eastern specimens); chaetotaxy of elytra identical to C. jamaicense
except for five or six setae on apical tubercle; hind calcars very small.

This species resembles C. jamaicense but is easily separated by the strongly depressed
Interval I (Fig. 159).

Range. — Jamaica, at elevations of 2000 ft. or less. Bell (1970) lists the following localities
in addition to the type locality: John Crow Mountains, Port Antonio, Ocho Rios (Fern Gulley),
Cornpuss Gap, Belmore Castle.

Variation. — There is considerable variation among collections from various localities, and
perhaps two or three subspecies will be recognized when more material is available. Specimens
from western localities, Mt. Diablo, Fern Gulley, and Belmore Castle, have the parasutural and
intercalary striae relatively shallow, and the marginal stria entire or nearly so. Specimens from
the John Crow Mountains and Cornpuss Gap have the parasutural and intercalary striae
relatively deep and the marginal stria interrupted near the humerus. Specimens from Port
Antonio resemble those from the John Crow Mountains in striation, but have first interval
extremely flat and more widened posteriorly than in other populations.

Bionomics. — Bell (1970) collected this species in logs and sticks in relatively moist forest
between sinkholes in the Karst Plateau of Mount Diablo. The locality at Fern Gulley is a very
moist ravine only a few meters above sea level.

THE ROSSI SECTION

In this section, the intercalary stria is complete, but the apex of the intratubercular stria is
absent, so that the preapical tubercle is not distinct from the apical tubercle. The median sulcus
of the metasternum is variable. Except in C. kochalkai , the eye is reduced to a narrow line.
There are five species, ranging from Costa Rica to western Venezuela.

Phytogeny. — C. rossi is probably the most isolated member of the section. In the absence of
a metasternal sulcus, and the presence of only one temporal seta and a pair of crossed occipital
setae, it differs from the remaining species.

The latter consists of two pairs of similar species. C. dormans and C. penicillatum lack
discal setae, have short discal strides, only three temporal setae and two or three marginals,
while the other pair, C. segne and C. kochalkai , have discal setae, long discal strides, four
temporal setae and five or more marginals. Each of these pairs has one species with a linear
median groove and precoxal setae present. (C. dormans, C. segne) and another with a dilated
median groove and without precoxal setae (C. penicillatum , C. kochalkai ) . Possibly these
characters indicate the real phylogeny, and the preceding characters are the result of
convergence. On balance, we believe that the converse is more likely true, that C. dormans is
related to C. penicillatum and C. segne to C. kochalkai.

Clinidium ( sensu stricto) rossi Bell 1970
(Fig. 156)

Clinidium ( sensu stricto) rossi Bell 1970:321-322.

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130

Bell and Bell

Type Material. — HOLOTYPE male, labelled: “Golfito, Costa Rica, Oct. 30, 1950, coll. E. S. Ross” (CAS).

Description. — Length 4.1 mm. Antennal stylet minute; antennae short, stout; tufts of minor setae on Segments
V-X; basal setae not studied; head 1.5 longer than wide; frontal grooves very shallow, anterior portion obsolete; median
lobe relatively narrow, triangular, tip opposite middle of eye; medial margins of temporal lobes parallel posteriorly;
posteriomedial, posterior margins of temporal lobe broadly margined with pilosity; orbital groove complete; eye very
narrow, linear, pigments in holotype; one temporal seta, in orbital groove posterior to eye; one pair of occipital setae
present, crossed; one pair of postlabial setae.

Pronotum moderately long, length/greatest width 1.53; lateral margins parallel in middle 0.33, obliquely narrowed to
base, apex; base rounded, apex truncate; median groove linear, not at all dilated at anterior median pit, narrowly dilated
between middle and basal 0.25; basal impressions elongate, triangular, closed posteriorly; discal striole relatively short,
reaching basal 0.33 of pronotum; marginal groove linear, visible in dorsal view; angular seta present; five marginal setae;
one pair of anterior discals; sternopleural groove distinct, though shallow; precoxal setae absent.

Elytra relatively short, broad; sutural, parasutural, intercalary striae complete, impressed, finely punctate;
intratubercular stria with apex not impressed, so preapical tubercle not distinct from apical tubercle; remainder of
intratubercular stria coarsely punctate, very shallowly impressed; marginal stria complete, deeply impressed; sutural striae
with four or five setae in complete row; parasutural stria without setae; intercalary stria with five or six setae in complete
row; intratubercular stria without setae; marginal stria with 10 setae, forming complete row; apical tubercle with three
setae, forming complete row; apical tubercle with three setae in line with intratubercular stria, one ventrad to them, near
suture; metasternum without median sulcus; transverse sulci of abdominal Sterna III- VI broadly interrupted in midline;
submarginal groove of Sternum VI widely separated from transverse groove; Sternum VI with six setae, four in transverse
row in middle, two near submarginal groove; male without ventral tooth on anterior femur nor proximal tooth on anterior
tibia; calcars very small; female with lateral pit on Sternum IV, shallower pit on Sternum V.

This species is easily recognized by the very shallow frontal grooves, the crossed occipital
setae and the presence of only one temporal seta.

In addition to the type specimen we have seen one male, one female labelled: “Golfito, Costa
Rica, July 7, 1957, Truxal & Menka” (LA).

Clinidium ( sensu stricto) dor mans new species
(Figs. 157, 162)

Type Material. — HOLOTYPE male, labelled: “Finca Lerida, near Boquete, Chiriqui Prov., Panama, Mar. 15,
1959, G. A. Salem leg. CNHM Panama Zoo. Exped. (1959) ADP 06974, La Barca, 5650'” (NMNH).

Description. — Length 5.2 mm. Antennal stylet minute; tufts of minor setae on Segments V-X; basal setae present
on Segments VII-X; Segments I-III with subapical pollinose rings; head longer than wide; frontal grooves very shallow,
glabrous; median lobe short, broad, triangular, tip even with middle of eye; medial margins of temporal lobes slightly
divergent posteriorly; anterior part of frontal space glabrous, shallow; posteriomedial, posterior margins of temporal lobe
margined with pilosity; orbital groove complete; eye very narrow, linear, 0.4 of length of temporal lobe; heavily pigmented
in holotype; three temporal setae, two opposite eye, one posterior to eye; occipital setae absent; postlabial setae apparently
absent (but possibly lost from holotype).

Pronotum moderately long, length/greatest width 1.47; lateral margins curved; apex truncate; base rounded; margin
oblique anterior to hind angle; median groove narrow, margins nearly parallel, except for slight expansion at anterior
median pit, constriction posterior to posterior median pit; basal impression elongate, triangular, discal striole relatively
short; extending to posterior 0.33 of pronotum; marginal groove linear, visible in dorsal view; angular seta present; one or
two marginal setae near apex of pronotum, also one just anterior to angular seta; discal setae absent; precoxal setae
present, sternopleural groove absent.

Elytra moderately elongate, relatively broad; sutural, parasutural, intercalary striae complete, impressed, finely
punctate; intratubercular stria virtually absent, represented by scattered, irregular punctures, preapical tubercle not
separated from apical tubercle (Fig. 162); marginal stria impressed, apical 0.25 dilated; sutural stria without setae;
parasutural stria with one seta at base; intercalary stria with complete row of five setae; marginal stria with one or two
setae near middle, four in impressed apical portion; apical tubercle with two or three setae on medial margin, one in line
with those of intercalary stria, also four or five arising from line of coarse punctures on lateral surface, probably
representing posterior part of intratubercular stria; metasternum with shallow median sulcus; transverse sulci of abdominal
sterna narrow, shallow, broadly interrupted at midline; Sternum VI with submarginal sulcus widely separated from
transverse sulci; Sternum VI with two pairs of setae, one on disc, other on submarginal sulcus; male without ventral tooth
on anterior femur nor proximal tooth on anterior tibia; false spur absent, short, broadly triangular tooth in its place; middle
calcar narrowly angular; hind calcar broadly triangular; both small. Female unknown.

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131

This species is closest to C. penicillatum of Colombia, but the latter species has the median
groove dilated in the middle 0.33, the frontal grooves deeper, precoxal setae absent, the discal
striole scarcely evident, and the pronotum strongly narrowed anteriorly.

Clinidium ( sensu stricto) penicillatum new species
(Figs. 163, 170)

Type Material.— HOLOTYPE female, labelled: “Colombia: Dept. Valle, 1967, R. B. Root, W. L. Brown,
Represa Calima, below dam, 1200 m., 21 Mar., canyon bottom” (MCZ).

Description. — Length 6.0 mm. Antennal stylet small, 0.1 of length of Segment XI; tufts of minor setae on
Segments V-X; basal setae present on Segments V-X; Segments I- VII with subapical pollinose rings; head 1.5 longer than
wide; frontal grooves narrow, moderately deep, partly pollinose; median lobe short, broad, triangular, tip margined by
pollinosity, even with anterior margin of eye, joined to antennal lobe; medial margins of temporal lobes slightly convergent
posteriorly; posteriomedial, posterior margins of temporal lobe bordered with pollinosity; orbital groove complete; eye very
narrow, linear, 0.4 of length of temporal lobe; three temporal setae, two opposite eye, one posterior to eye; occipital setae
absent; one pair of postlabial setae.

Pronotum rather elongate, length/greatest width 1.57; greatest width near base; margins convergent anteriorly; apex
narrow, truncate; base slightly narrowed, rounded; anterior 0.1, posterior 0.33 of median groove linear, middle portion
narrowly dilated, conspicuously pollinose; basal impression elongate, triangular; discal striole scarcely evident; angular seta
present; two or three marginals present, near apex, one at base; discal setae absent; precoxal setae absent; stemopleural
groove absent.

Elytra moderately elongate; sutural, parasutural, intercalary striae complete, impressed, rather coarsely punctate;
intratubercular stria nearly absent, represented by row of very fine punctures; preapical tubercle not separate from apical
tubercle; marginal stria impressed, apical 0.25 dilated; sutural stria with one seta near apex; parasutural stria with one seta
at base; intercalary stria with complete row of five setae; marginal stria with one seta near middle, two behind middle, and
five or six near apex; apical tubercle with row of three or four setae on medial margin, row of six on lateral surface, in line
with intratubercular stria; metasternum with shallow, broad median sulcus; transverse sulci of Sterna III-VI broadly
interrupted in midline, each with row of fine punctures; Sternum VI with submarginal sulcus widely separated from
transverse sulci; Sternum VI with eight setae, four in transverse row, four in curved row posterior to submarginal sulcus;
female with shallow lateral pit on Sternum IV, very shallow one on Sternum V; false spurs absent; hind femur of female
with dense brush of long pilosity on dorsal aspect (Fig. 170); male unknown.

This species is closest to C. dormans but differs in having the median lobe connected to the
antennal lobe, in having the middle part of the median groove dilated, and the pronotum widest
near the base and strongly tapered anteriorly. The brush on the hind femur is unique, but may
be a secondary sexual character, as the male is unknown.

Clinidium ( sensu stricto) segne new species
(Figs. 164, 173)

Type Material. — HOLOTYPE female, labelled: “VEN. Edo. Aragua, Rancho Grande, 1500 m. (15 km n. of
Maracay), 21-11-1971, S. Peck” (BSRI). PARATYPE one female, same label as holotype (BSRI). The type locality is
near the north coast of Venezuela, a little west of Caracas.

Description. — Length 4. 6-5. 2 mm. Antennal stylet very slender, 0.25 of length of Segment XI; tufts of minor
setae on Segments V-X; Segments I- VII with subapical pollinose rings; basal setae on Segments IX-X or VIII-X, sparse;
head slightly longer than wide; frontal grooves deep, narrow, median lobe short, triangular, tip opposite anterior part of
eye; medial margins of temporal lobes slightly divergent posteriorly; posterior, posteriomedial margins of temporal lobe
very broadly bordered by pilosity; orbital groove complete; eye narrow, short, about 0.3 of length of temporal lobe; four
temporal setae, one anterior to eye, two opposite eye, one posterior to eye; occipital setae absent; two pairs of postlabial
setae.

Pronotum moderately long, length/greatest width 1.47; widest slightly behind middle, lateral margins curved; apex
narrowed, truncate; base moderately narrowed, rounded; median groove narrow, margins parallel, not at all expanded at
anterior median pit; posterior 0.20 very narrow, shallow; basal impression narrow, triangular; discal striole linear, straight
or slightly curved, extending 0.30 to 0.40 of length of pronotum; marginal groove slightly dilated, visible in dorsal view;
angular seta present; nine or 10 marginal setae; two pairs of discal setae, more anterior ones near anterior margin,
posterior ones slightly anterior to middle; precoxal seta present; anterior part of stemopleural groove faintly suggested;
middle part absent.

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132

Bell and Bell

Elytra moderately elongate; sutural, parasutural, intercalary striae complete, impressed, finely punctate;
intratubercular stria shallowly impressed, base entire, apex effaced, so preapical tubercle not separate from apical tubercle
(Fig. 173); marginal stria impressed, entire, apical 0.25 dilated; sutural stria with complete row of five setae; one seta in
conspicuous puncture at anteriomedial angle of Interval III; intercalary stria with complete row of nine setae;
intratubercular stria without setae; marginal stria with complete row of about 1 8 setae; apical tubercle with two setae on
medial margin, row of five setae in isolated punctures on lateral surface, in line with intratubercular striae; metasternum
with deep median sulcus; transverse sulci of Sterna III-VI narrow, impunctate, broadly separated in midline; submarginal
sulcus of Sternum VI widely separated from transverse sulci; Sternum VI with six setae, two on disc, four posterior to
submarginal sulcus; female with shallow lateral pit on Sternum IV, very shallow one on Sternum V; false spurs absent.
Male unknown.

In chaetotaxy, in having a long discal stride and an impressed intratubercular stria, this
species resembles C. kochalkai. In the latter species, the median groove is much more dilated,
and there is at most one seta in the sutural stria.

Clinidium ( sensu stricto) kochalkai new species
(Fig. 165)

Type Material. — HOLOTYPE male, labelled: “COLOMBIA 8860', J. A. Kochalka, Casa Antonia, Loma
Cebolleta, S(ierra) N(evada) de Santa Marta, V-8-1975” (to be deposited in NMNH). PARATYPE female, same label
as holotype (to be deposited in NMNH).

Description. — Length 6.0 mm. Antennal stylet very slender, long, 0.33 of length of Segment XI; tufts of minor
setae on Segments V-X; Segments I-IV with subapical pollinose rings, V-VI with small pollinose spots; a few basals on
Segment X; head slightly longer than wide; frontal grooves deep, broader than in C. segne; median lobe short, triangular,
tip opposite anterior margin of eye; medial margins of temporal lobes slightly convergent posteriorly; margins of temporal
lobe with much narrower pilose borders than C. segne ; orbital groove complete; eye narrow, 0.5 of length of temporal lobe;
three or four temporal setae in orbital groove, one or two at anterior margin of eye; one at its posterior margin, one
posterior to eye; occipital setae absent; two or three pairs of occipital setae.

Pronotum moderately long, length/greatest width 1.43; widest near middle, lateral margins curved; apex, base
moderately narrowed, apex truncate, base curved; median groove dilated, deep in middle, tapered anteriorly, not at all
dilated at anterior median pit; posterior 0.25 linear, shallow; basal impression triangular, tapered gradually into discal
striole, latter broader than in C. segne-, discal stride attaining middle of pronotum; marginal groove dilated, prominent in
dorsal view; six or seven marginal setae; one pair of anterior discal setae; precoxal setae absent; sternopleural setae absent.

Elytra rather broad; sutural, parasutural, striae complete, impressed, finely punctate; intercalary stria impressed, apex
entire, but base abbreviated posterior to level of humerus; intratubercular stria shallowly impressed, apex effaced so
preapical tubercle not separate from apical tubercle; marginal stria impressed, entire, apical 0.25 dilated; sutural stria
without setae, or with one seta anterior to middle; anteriomedial angle of Interval III without seta; intercalary stria with
complete row of seven or eight setae; intratubercular stria without setae; marginal stria with complete row of nine or 10
setae; medial margin of apical tubercle with five setae, lateral surface with four or five setae in conspicuous punctures
aligned with intratubercular stria; metasternum with deep median sulcus; transverse sulci of Sterna III-V linear, broadly
separated in midline; transverse sulci of Sternum VI dilated, oval broadly separated; submarginal sulcus of Sternum VI
dilated, well separated from transverse sulci; Sternum VI with six setae, four posterior to submarginal sulci, two on disc,
widely separated; anterior margin of submarginal sulcus evenly curved in male, angulate, slightly tuberculate in midline in
female; female with shallow lateral pit on Sternum IV; middle hind tibiae without false spur, but with short, triangular
tooth in its place; male without ventral tooth on anterior femur, without proximal tooth on anterior tibia; calcars raised
above level of spurs; obliquely truncate.

This species is easily recognized by the dilated middle portion of the median groove of the
pronotum and by the abbreviation of the base of the intercalary stria. It is named for the
collector, the able and enthusiastic arachnologist, John A. Kochalka, our friend and former
student.

THE GUILDINGII SECTION

In this section, the intercalary stria is abbreviated posteriorly, while the intratubercular stria
is complete to the apex, separating the preapical tubercle from the apical tubercle. The
metasternum lacks a median sulcus. The false spurs are well-developed. The section consists of

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133

four species, each confined to one island in the Lesser Antilles.

Phytogeny. — Of the four species, C. guildingii is the most isolated, with the apical tubercles
separated by a large, round space, and the female with Sternum VI with a median tubercle.
The three remaining species have a small pore beneath the apical tubercle, which are broadly
contiguous above it. Of these, C. microfossatum contrasts with the remaining two in the virtual
disappearance of the intratubercular stria and the abbreviation of the marginal stria at the
base. The femur of the male has many minute tubercles on the ventral surface, a feature not
found in other members of the genus. C. planum and C. smithsonianum are closely related to
one another, but differ in secondary sexual characters and in the length of the discal striole.

Clinidium {sensu stricto) guildingii Kirby 1835
(Figs. 167, 174, 176, 177)

Clinidium guildingii Kirby 1835: 8-10.

Rhysodes guildingii (Kirby) Newman 1838. Chevrolat (1873a) changed the spelling to “guildingi”, a practice followed by
most later authors.

Clinidium ( sensu stricto) guildingii (Kirby) Bell 1970.

Type Material. — We have not been able to locate type material for this species. The original description does not
indicate the location of types, and the latter are not in the British Museum of Natural History. According to the
description, the type locality is Mount Saint Andrews, Saint Vincent. We have studied two males and one female collected
by ourselves at the type locality. If the type series is really lost, these could serve as neotypes.

Description. — Length 5. 5-6.0 mm. Antennal stylet acuminate, 0.3 of length of Segment II; tufts of minor setae on
Segments V-X; basal setae present on Segments VII-X or VIII-X; Segments I-VI with subapical bands of pollinosity; head
slightly longer than wide; frontal grooves moderately deep, pollinose, of even width; median lobe short, broad, triangular,
tip even with anterior part of eye; frontal space moderately broad; medial margins of temporal lobes oblique, slightly
divergent posteriorly; posterior, posteriomedial margins of temporal lobe bordered with pollinosity; orbital groove narrow,
complete; eye narrowly crescentic, deeply pigmented in mature specimens; most specimens with three temporal setae, in
orbital groove; a few specimens with two or four temporals; two pairs of postorbital setae.

Pronotum elongate, length/greatest width 1.49; lateral margins curved; apex strongly narrowed, base moderately
narrowed; apex truncate; base rounded; margin oblique anterior to hind angle; median groove slightly dilated, margins
parallel in middle, groove slightly enlarged at anterior median pit; basal 0.33 of groove narrow, shallow; basal impression
narrow, triangular; discal striole long, attaining middle of pronotum; marginal groove visible in dorsal view; angular seta
present, anterior, medial to hind angle; about 10 marginal setae present; discal setae, precoxal setae absent; sternopleural
groove complete.

Elytra short, relatively broad; striae impressed, indistinctly punctate; intercalary stria abbreviated posteriorly;
intratubercular stria complete; marginal stria complete; apical tubercles inflated, truncate posteriorly, touching at
dorsoposterior points above large, round opening (Fig. 177); sutural stria with complete row of four or five setae;
intercalary stria with complete row of six to eight setae; intratubercular stria with one seta at base, three near apex;
marginal stria with about 1 7 setae; apical tubercle with three setae in prominent punctures; metasternum without median
sulcus; male with transverse sulci of abdominal Sterna III-V complete, not interrupted in midline, with those of VI
narrowly interrupted in midline, connected laterally to submarginal sulcus; female with sulci of Sterna III-IV complete,
that of V narrowly interrupted; that of VI broadly interrupted; Sternum VI in both sexes with one pair of setae in
submarginal sulcus; Sternum VI of female with median tubercle, disc sloped gradually posterior to it (Figs. 174, 176);
male anterior femur without ventral tooth; male anterior tibia without proximal tooth; false spurs present on middle, hind
tibiae; middle, hind calcars triangular, not notched on dorsal margin.

The large circular opening between the apical tubercles is distinctive. The male has calcars
similar to those of C. microfossatum , but does not have tubercles on the ventral side of the
anterior femur. The female differs from other known females in having a tubercle in the middle
of Sternum VI.

Range. — St. Vincent, in the Lesser Antilles. We have studied two males and one female,

labelled: “Checkley Level, Mount Saint Andrew, coll. J. R. Bell, Dec. 31, 1968" (UVM), and one female, labelled:
“Richmond Est., Oct. 31, open valley, sea level in rotten wood, Kingstown” (collector and year not given) (BMNH).

There is one female labelled “Guadeloupe”, which is clearly this species and not C. planum.
We suspect that the locality label is incorrect.

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Bionomics. — Bell (1970) describes in detail the situation of the specimens from Mount
Saint Andrews. These were collected in the stump and roots of Torrubia fragrans (Du Mont de
Courset), a member of the Nyctaginaceae.

Clinidium ( sensu stricto) microfossatum new species
(Figs. 168, 171)

Type Material. — HOLOTYPE male, labelled: “La Martinique, Dr. L. Pornain, 1901” (MNHN).

Description. — Length 5.8 mm. Antennal stylet acuminate, about 0.3 of length of Segment XI; tufts of minor setae
on Segments V-X; basal setae present on Segments VI-X or VII-X; Segments I-III with subapical bands of pollinosity;
head distinctly longer than wide; frontal grooves deep, narrower than in C. guildingii, of even width, margined with
pollinosity; median lobe short, broad, triangular, tip even with anterior part of eye; frontal space rather narrow; medial
margins of temporal lobes oblique, slightly divergent posteriorly; posterior, posteriomedial margins of temporal lobe with
very narrow pollinose borders; orbital groove very narrow, complete or with short interruption posterior to eye; eye
narrowly crescentic; orbital groove with two temporal setae, one opposite eye, the other near posterior margin; two pairs of
postlabial setae.

Pronotum elongate; length/greatest width 1.55; lateral margins curved; apex strongly narrowed, more so than in C.
guildingii-, base moderately narrowed; apex truncate; base rounded; margin oblique anterior to hind angle; median groove
narrow, less dilated than in C. guildingii, margins parallel; groove very slightly dilated at anterior median pit; basal 0.33
very narrow, shallow; basal impression small, oval, discal striole absent; marginal groove visible in dorsal view, finer than
in C. guildingii', angular seta medial, anterior to hind angle; five or six marginal setae; discal, precoxal setae absent;
sternopleural groove effaced anteriorly.

Elytra short, relatively broad; sutural, parasutural, intercalary striae impressed, pollinose; intercalary stria abbreviated
posteriorly; intratubercular stria not impressed, not pollinose, represented only by line of fine punctures, preapical tubercle
thus scarcely separated from apical tubercle (Fig. 171): marginal stria incomplete, basal 0.25 entirely effaced, next 0.25
represented only by row of fine punctures; apical 0.5 impressed; apical tubercles inflated, truncate, meeting in straight line
at suture, minute pore in midline below them; sutural stria with four or five setae in complete row; intercalary stria with
complete row of eight setae; intratubercular with one seta at base, one at apex; marginal stria with about nine setae; apical
tubercle with two setae in line with intratubercular stria; five setae in row ventrad to preceding; metasternum without
median sulcus; male with transverse sulci of Sterna III, IV complete, that of V narrowly interrupted in midline, that of VI
broadly interrupted in midline, widely separated from submarginal sulcus; Sternum VI with one pair of setae; male with
anterior femur without ventral tooth, but with many minute tubercles on ventral surface; male without proximal tooth on
anterior tibia; false spurs present; calcars triangular, dorsal margins straight, not notched. Female unknown.

The reduction of the intratubercular stria and of the base of the marginal stria are
distinctive. The virtual absence of the discal strides separates it from C. smithsonianum and C.
guildingii. Some individuals of C. planum have the strides equally reduced, but differ in the
shape of the frontal space and, in the male, in the absence of tubercles on the ventral side of the
anterior femur, and in the shape of the calcars.

Clinidium {sensu stricto) smithsonianum new species
(Figs. 169, 175, 178)

Type Material. — HOLOTYPE male, labelled: “Dominica: 2 mi. NW Pont Casse, X-26-1964, P.J. Spangler”
(NMNH). Twelve PARATYPES: two females, same data as holotype. (one specimen missing head and thorax) (NMNH);
four females labelled: “Dominica, 3.0 mi. E. of Pont Casse, VII-3 1-1964, T.J. Spilman” (NMNH); one female labelled:
“Dominica, 1.0 mi. E. of Pont Casse, VII-23-1964, T. J. Spilman” (NMNH); one male, one female labelled: “Dominica,
0.6 mi. W. of Pont Casse, VII-7-1964, T. J. spilman” (NMNH). (All the foregoing specimens also bear the label:
“Bredin-Archbold Smithsonian Survey”.); one male, one female, labelled: “Dominica, nr. Jean, 2000', 1 1-17-65, JFGC &
T. M. Clarke, in rotten log” (NMNH); one female, labelled: “Wet Area Exp. Sta. St. Joseph Parish, 800', 31 Dec. 1978,
M. A. & L. L. Ivie” (MAI).

Description. — Length 5.0-6. 1 mm. Antennal stylet acuminate, about 0.3 of length of Segment XI; tufts of minor
setae on Segments V-X; basal setae present on Segments V-X; Segments I-V with pollinose subapical bands; head
distinctly longer than wide; frontal grooves deep, rather narrow, of even width, margined by pollinosity; median lobe short,
broad, triangular, tip even with anterior part of eye; frontal space rather narrow; medial margins of temporal lobes oblique,
slightly divergent posteriorly; posterior, posteriomedial margins of temporal lobes very narrowly bordered with pollinosity;

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135

orbital groove very narrow, complete or with short interruption posterior to eye; eye narrowly crescentic; orbital groove
with two temporal setae, one opposite eye, the other near posterior margin; two pairs of postlabial setae.

Pronotum moderately elongate; length/greatest width 1.50; lateral margins curved; apex strongly narrowed; base
moderately narrowed; apex truncate; base rounded; margin oblique anterior to hind angle; median groove narrow, margins
parallel; groove very slightly dilated at anterior median pit; basal 0.33 very narrow, shallow; basal impression small,
triangular, discal striole well-developed, with basal impression about 0.35 to 0.40 of length of pronotum; marginal groove
fine, visible in dorsal view; angular seta medial, anterior to hind angle; seven or eight marginal setae; discal, precoxal setae
absent; sternopleural groove absent.

Elytra short, relatively broad; all striae impressed, pollinose; intercalary stria abbreviated posteriorly; marginal stria
complete to base; apical tubercles inflated, truncate, meeting in straight line at suture, minute pore in midline below them
(Fig. 178); sutural stria with complete row of four or five setae; intercalary with complete row of eight or nine setae;
intratubercular stria with one seta at base, three or four setae in apical 0.33; marginal stria with 13-14 setae; apical
tubercle with three setae in prominent punctures; metasternum without median sulcus; male with transverse sulci of Sterna
III- V complete; transverse sulci of VI widely separated at midline, widely separated from submarginal sulcus; female with
transverse sulci of Sterna III-IV complete; sulcus of V narrowly interrupted at midline; Sternum VI of female with
transverse scarp at middle of length, scarp bounded posteriorly by deep oval impression, latter with central convexity,
bounded laterally by longitudinal oval, pollinose cavity (Fig. 175); female with lateral pit on Sternum IV; both sexes with
one pair of setae on Sternum VI, posterior to submarginal sulcus; false spurs present; male without ventral tooth or
tubercles on anterior femur; male without proximal tooth on anterior tibia; calcars with dorsal margins weakly angulate.

The long discal striole gives this species a similarity to C. guildingii , but it differs from the
latter species in having only a minute pore beneath the apical tubercles. C. planum is more
closely related, but has shorter discal strides, and differs in secondary sexual characters, the
male having more strongly angulate calcars, and the female having the impression of Sternum
VI without a central tubercle.

In addition to type material, we have seen two males labelled: “Dominica, St. Peter
syndicate estate, under bark, 7-10-VII-1970, coll. J. H. Frank” (BMNH); one female, labelled:
“Dominica, Springfield Est., VI-20-25-69, P. J. Darlington, Jr.” (MCZ).

Clinidium ( sensu stricto) planum (Chevrolat 1844)

(Figs. 166, 172)

Rhyzodes planus Chevrolat 1844: 58.

Clinidium guildingii Kirby (wrongly synonymized by Chevrolat 1873a).

Clinidium planum (Chevrolat) Arrow 1942.

Clinidium ( sensu stricto) planum (Chevrolat) Bell 1970.

Type Material. — We have not been able to locate type material. We have studied a specimen from the type
locality, Point-a-Pitre, Guadeloupe. It is a male, labelled: “Point-a-Pitre, Guadeloupe, W.I., June 6, 1911” (AMNH). If
the type of C. planum is lost, this specimen could serve as a neotype. Another similar male specimen is labelled
“Guadeloupe, Vitrac” (GEN).

Description. — Length 5. 3-6. 3 mm. Antennal stylet acuminate, 0.3 of length of Segment XI; tufts of minor setae
on Segments V-X; basal setae present on Segments VI-X; Segments I-V with subapical bands of pollinosity; head slightly
longer than wide; frontal grooves narrow, rather shallow, pollinose, narrowed near junction with frontal space; median lobe
short, broad, triangular, tip even with anterior part of eye; frontal space rather narrow, anterior part with margins parallel,
separated by more or less distinct angles from posterior part with oblique margins; posterior, posteriomedial margins
bordered by pollinosity; orbital groove very narrow, complete or with short interruption posterior to eye; eye narrowly
crescentic, orbital groove with two temporal setae; one opposite eye; other near posterior end of orbital groove: two pairs of
postlabial setae.

Pronotum moderately elongate; length/greatest width 1.51; lateral margins curved; apex narrowed; base only slightly
less narrowed than apex; apex truncate; base rounded; margin oblique anterior to hind angle; median groove slightly
dilated near middle, narrowed anteriorly, scarcely dilated at anterior median pit; basal 0.33 very shallow, narrow; basal
impression small, triangular; discal striole relatively short, 0.1 to 0.25 of length of pronotum; marginal groove fine, visible
in dorsal view; angular seta medial, anterior to hind angle; seven or eight marginal setae; discal, precoxal setae absent;
sternopleural groove absent.

Elytra relatively short, broad; all striae impressed, pollinose; intercalary stria abbreviated posteriorly (Fig. 172);
marginal stria complete to base; apical tubercles inflated, truncate, meeting in straight line at suture, minute pore in
midline beneath them; sutural stria with complete row of four or five setae; intercalary stria with complete row of five to
seven setae; intratubercular stria with one seta at base, three in apical 0.33; marginal stria with six to eight setae in

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Bell and Bell

complete row, sparse near middle of length; apical tubercle with three setae in prominent punctures; metasternum without
median sulcus; male with transverse sulcus of Sternum III complete, those of III, IV complete in some specimens, narrowly
interrupted in others; that of V narrowly separated; female with transverse sulci of Sterna III-V not interrupted; female
with deep lateral pit on Sternum IV and shallow one on Sternum V; in both sexes, Sternum VI with submarginal sulcus
widely separated from transverse sulci; one pair of setae, posterior to transverse sulcus; female with transverse scarp on
Sternum VI, bounded posteriorly by deep, entirely pollinose impression without median convexity; scarp in lateral view
forming right angle with anterior part of disc; false spur on middle and hind tibiae present; male without ventral tooth or
tubercles on anterior femur, without proximal tooth on anterior tibia; calcars strongly angulate on dorsal margin; margin
distinctly notched between angle and shaft of tibia.

The discal strides of this species are intermediate in length, separating it from C. guildingii
and C. smithsonianum , which have long ones, and probably from C. microfossatum which
almost lacks them. The male is also distinguished by the strongly notched dorsal margin of the
hind calcar, and the female by the deep pollinose impression of Sternum VI.

Range. — Probably confined to Guadeloupe. We have seen several specimens labelled “W.
Ind.” (MNHN) that appear to belong to this species. In addition we have seen a specimen
labelled “Mexico, Bowditch” (MCZ) that is either C. planum or else another species closely
related to it. It is a female which resembles C. planum except that the impression of Sternum
VI is not pollinose, and the disc of Sternum VI has a broad, low tubercle anterior to it. The
frontal grooves are very narrow. The specimen probably bears an incorrect locality label. If not
an aberrant C. planum , it might belong to an undescribed species from one of the Lesser
Antilles, such as Grenada or Saint Lucia, from which no members of the section have yet been
described.

THE ROJASI SECTION

This section resembles the guildingii section in having the intercalary stria abbreviated
posteriorly and the intratubercular stria complete. However, the false spurs are absent. The
metasternum has the median sulcus well-developed, and the apical tubercle is strongly
emarginate. The male has a proximal tooth or angle on the anterior tibia. There are four, or
possibly more, species confined to the mountains near the coast of Venezuela, from Falcon
State eastward.

Phylogeny. — Unfortunately two of the species, C. pala and C. excavatum are known only
from the females. They show distinctive modifications of Sternum VI. This character suggests
that they are sister species. Non-sexual characters, however, suggest that C. pala is closest to C.
rojasi and C. excavatum, to C. bechyneorum. As noted under “variation”, there are possible
additional species in this section. An analysis of phylogeny must wait until more material is
collected.

Clinidium ( sensu stricto) rojasi Chevrolat 1873a
(Figs. 179, 184, 185, 186)

Clinidium rojasi Chevrolat 1873a: 211-215.

Clinidium ( sensu stricto ) rojasi (Chevrolat) Bell and Bell 1978.

Type Material. — We have been unable to study the types of this species. According to the original description,
there were two specimens, one collected by Rojas, the other by Salle. Both were labelled simply “Venezuela”. Vulcano and
Pereira (1975b) studied the types, borrowed from NMW. According to them, both syntypes are female. We studied a
female example, labelled: “type, Colonia Tovar, E. Simon, III-88, exemplaire typique” (MNHN). This species is not an
authentic type, as it was not listed in the original description, and was collected long after the publication of the name.
Thus it is not quite certain which member of the section was really described by Chevrolat. We follow Vulcano and Pereira
(1975b) in assigning it to this species, as their illustrations of the male legs show acute calcars and a sharp proximal tooth.
The Simon specimen also belongs to this species, as shown by the pollinose frontal space and the absence of setae on the

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137

parasutural stria.

Clinidium simplex Chevrolat 1873b: 378 is difficult to interpret. We have studied a specimen labelled: “Dr. Moritz,
1858, Venezuela, Clinidium simplex ” (NMW). This was lent to us as the type, but is not labelled as such, and is from the
wrong locality. The type locality in the original description is given as “Nova-Grenata”. The Vienna specimen is a male C.
rojasi. However, Vulcano and Pereira (1975b) describe and illustrate a different species under this name, and state that
the description is based on a specimen labelled as a type and as from Nova Grenata. It appears to belong to the rojasi
section, and has the emarginate apical tubercle typical of that section. According to the figure, it is longer and more
slender than C. rojasi , with the head more narrowed behind. Thus it may represent an additional species in the section.
This may also not be the real type, as it differs from the original description in an important feature, in not having double
marginal grooves on the pronotum. No South American species of Clinidium known to us has double marginal grooves, a
feature found in most species of subgenera Arctoclinidium and Mexiclinidium. At the present time we cannot interpret the
name C. simplex with any certainty.

Description. — Length 4.7-5. 8 mm. Antennal stylet acuminate, elongate, 0.3 of length of Segment XI; latter
elongate; tufts of minor setae on Segments V-X; basal setae present on Segments VII-X or VIII-X; Segments I-VIII with
pollinose subapical rings; head scarcely longer than wide; base rather broad; margin oblique posterior to eye; frontal
grooves deep, narrow, pollinose; median lobe short, broad, triangular, tip even with anterior part of eye; frontal space
rather narrow, completely pollinose; medial margins of temporal lobe oblique, slightly divergent posteriorly; posterior,
posteriomedial margins of temporal lobe bordered by pilosity; orbital groove complete; eye narrow, crescentic, heavily
pigmented in most specimens; most specimens with three temporal setae, one midorbital, one postorbital, one posttemporal,
one or two of these missing in some specimens; two pairs of postlabial setae.

Pronotum elongate; length/greatest width 1.59; lateral margins curved; apex strongly narrowed; base moderately
narrowed; apex truncate, base rounded; median groove nearly linear, expanded at anterior median pit, latter about 0.1 of
width of pronotum at apex; basal 0.2 of median groove very shallow, narrow; pollinosity of median groove connected to
transverse band at base of pronotum; latter occupying about 0.3 of width of base; basal impressions small, triangular,
closed posteriorly; discal striole slightly curved, extending to middle of length of pronotum; marginal groove visible in
dorsal view; angular seta anterior, medial to hind angle; nine to 12 marginal setae; discal, precoxal setae absent; anterior
part of sternopleural groove absent, posterior part barely indicated.

Elytra short, relatively broad; striae impressed, pollinose, indistinctly punctate; intercalary stria abbreviated
posteriorly; apical tubercles inflated, strongly emarginate (Fig. 185); sutural stria with complete row of four to six setae;
parasutural stria without setae; intercalary stria with complete row of nine to 1 1 setae; intratubercular stria with two setae
near base, two near apex; marginal stria with complete row of 11-14 setae; anterior medial angle of Interval III with one
seta in prominent pollinose pit; apical tubercle with three to six setae in prominent punctures; metasternum with complete
median sulcus; in both sexes transverse sulci of abdominal Sterna III-IV entire, those of V and VI interrupted in midline,
submarginal groove of Segment VI well separated from transverse sulci, deeply U-shaped in female (Fig. 186), transverse
in male; Sternum VI evenly convex in both sexes, with one pair of setae; female with lateral pit in Sternum IV; false spur
absent; male without ventral tooth on anterior femur, but with broad, obtuse proximal tooth on anterior tibia (Fig. 184);
calcars acute, small.

The acute calcars of the male and the unmodified Sternum VI of the female separate this
species from other members of the section.

Range. — Andes of northern Venezuela, from Falcon State (Cerro Galicia) on the west to

Aragua State (Tiara) on the east. We have studied the following specimens: one male, one female,
labelled: “Caracas, Silla” (MNHB); four males, four females labelled: “Cero Galicia, Venezuela, Falcon, 1500 m.,
22-XI- 1 97 1 , J. & B. Bechyne leg.” (VEN); one male, labelled: “Colonia Tovar, capacha bajo, Venezuela, Aragua,
24-IX-1968, en corteza de guamo” (VEN); one female, labelled: “Colonia Tovar, E. Simon, III-88, exemplaire typique”
(MNHN); one female, labelled: “Venezuela, Aragua, Tiara, 1 6- VI- 1 970, J. & B. Bechyne leg.” (VEN); one male,
labelled: “Venez., Fry Colin. 1905, 100, 18628. ” (BMNH). According to the ledger of the Fry Collection in the British
Museum, this last specimen was collected at Caracas.

Variation. — An additional female specimen from Tiara has the posterior 0.5 of Sternum VI
shallowly impressed and the transverse sulci of Sterna II- VI interrupted at the midline. This
specimen might represent an extreme variant of C. rojasi , but could also belong to a different
species. It was collected with the typical C. rojasi female from Tiara referred to above, and
bears an identical label.

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Clinidium ( sensu stricto) bechyneorum new species
(Figs. 180, 187)

Type Material. — HOLOTYPE male, labelled: “Hac. Montero, Montalban, Venezuela, Carabobo, 1300 m.,
18-IV-1968, J. & B. Bechyne leg.” (VEN). PARATYPES one male, one female, same label as holotype. (VEN)

Description. — Length 5. 5-6.0 mm. Antennal stylet acuminate, elongate, 0.3 of length of Segment XI, latter
elongate; tufts of minor setae on Segments V-X; basal setae present on Segments VII-X; Segments I-IV with subapical
pollinose rings; head as long as broad, base broad, margin not oblique posterior to eye; frontal groove deep, narrow, partly
pollinose; median lobe short, broad, triangular, tip opposite anterior part of eye; frontal space rather narrow, anterior 0.5
glabrous, posterior 0.5 pollinose; medial margins of temporal lobes oblique, slightly divergent posteriorly; posterior,
posteriomedial margins of temporal lobes bordered by pollinosity; orbital groove complete, wider than in C. rojasi; eye
narrow, crescentic, longer than in C. rojasi ; two or three temporal setae; two pairs of postlabial setae.

Pronotum elongate; length/greatest width 1.58; lateral margins curved, slightly sinuate anterior to hind angle; apex
strongly narrowed; base moderately narrowed; apex truncate; base rounded; median groove very shallow, narrow;
pollinosity of median groove connected to transverse band of pollinosity occupying median 0.3 of base of pronotum; basal
impressions small, triangular, closed posteriorly; discal stride slightly curved, extending to middle of length of pronotum;
marginal groove fine, visible in dorsal view; angular seta anterior, medial to hind angle; 7-10 marginal setae; discal,
precoxal setae absent; sternopleural suture absent.

Elytra short, relatively broad; striae impressed, pollinose, indistinctly punctate; intercalary stria abbreviated
posteriorly; apical tubercles inflated, strongly emarginate; sutural stria without setae or with one or two near apex;
parasutural stria with two to four setae; intercalary stria with complete row of four or five; intratubercular stria without or
with one basal and without or with one apical seta; marginal stria with complete row of eight to 15 setae; apical tubercle
with two to five setae in prominent punctures; metasternum with complete median sulcus; in both sexes, transverse sulci of
Sternum III, IV entire, those of V, VI interrupted in midline; submarginal sulcus of Sternum VI in both sexes short,
scarcely curved (Fig. 187); Sternum VI with one pair of setae; Sternum VI evenly convex, similar to that of male; false
spur absent; male without ventral tooth on anterior femur, with broad, obtuse proximal tooth on anterior tibia; calcars
obtusely rounded at apices.

The presence of setae in the parasutural stria, the rounded calcars of the male, and the
shape of the submarginal sulcus of Sternum VI of the female separate this species from C.
rojasi. The evenly convex Sternum VI of the female separate it from C. excavatum. We
dedicate this species to J. & B. Bechyne whose fine series of Clinidium have made the
Rhysodine fauna of Venezuela the best known of any South American country.

Variation. — A single male specimen, labelled: “Venezuela, Aragua, Rancho Grande, 1400
m., 26-VIII-70, J. & B. Bechyne leg.” (VEN) may represent this species. It is in poor condition,
with middle and hind legs missing, and with most setae of the head missing. It was probably
dead when found. The orbital groove is very narrow and the pollinosity is very reduced at the
posterior margin of the temporal lobe, exposing a distinct occipital angle. It is not clear whether
this represents a real difference from C. bechyneorum or is the result of abrasion after death.
The regular arrangement of the pollinosity suggests that the former is more probably. The
absence of the middle and hind legs prevents comparison of the calcars with those of C.
bechyneorum. We suspect that this specimen represents an additional species, but decline to
name it until better material is available.

Clinidium (sensu stricto) excavatum new species
(Fig. 188)

Type Material. — HOLOTYPE female, labelled: “Venezuela-Carabobo-Montalban Oeste 1800 mts.
26-VI-1968, C. J. Rosales, A. D. Ascoli” (VEN). PARATYPE one female, same label as holotype. (VEN)

Description. — Length 6. 5-6.8 mm. Identical to C. bechyneorum in most respects, but larger with more elytral
setae, and with Sternum VI strikingly modified. Setae within the ranges given for C. bechyneorum , except for intercalary
stria, with eight setae and intratubercular without or with one at base and three near apex. Sternum VI with deep
concavity in posterior 0.33, evidently representing a greatly enlarged submarginal sulcus; cavity bounded anteriorly by
scarp, latter with broad rectangular lobe in midline, bordered on each side by deep, prominent notch (Fig. 188). Male

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139

unknown.

C. pala has a similarly modified sternum, except that the notches are much closer together
and are convergent anteriorly.

Clinidium ( sensu strict o ) pala new species
(Fig. 189)

Type Material. — HOLOTYPE female, labelled: “VEN: ,Edo. Miranda Guatopo Nat. park, 50 km. SE Caracas,
5-6 III, 1971, 400m forest & hum dung” (BSRI). PARATYPE one female, same label as holotype. (BSRI)

Description. — Length 5.0-5. 5 mm. Antennal stylet acute, both stylet and Segment XI less elongate than in C.
rojasr, tufts of minor setae on Segments V-X; basal setae present on Segments VI-X or VII-X; Segments I- VIII ringed
with pollinosity; head distinctly longer than wide, longer, narrower, more parallel-sided than in C. rojasr, frontal groove
deep, narrow, pollinose, median lobe short, rather narrow, triangular, tip opposite anterior margin of eye; frontal space
narrow, anterior part scarcely pollinose; medial margins of temporal lobes oblique, slightly divergent posteriorly; posterior,
medial margins of temporal lobes bordered by pilosity; orbital groove complete; eye very narrow, crescentic; two temporal
setae; two pairs of postlabial setae.

Pronotum elongate; length/greatest width 1.53; lateral margins curved; apex strongly narrowed; base moderately
narrowed; apex truncate; base rounded; median groove nearly linear, scarcely expanded at anterior median pit; latter
smaller than in C. rojasr, basal 0.2 of median groove very narrow, shallow; pollinosity of median groove connected to
transverse band at base of pronotum, latter occupying about 0.3 of basal width; basal impressions small, triangular, closed
posteriorly; discal striae curved, extending almost to middle of length of pronotum; marginal groove visible in dorsal view;
angular seta anterior, medial to hind angle; eight or nine marginal setae; discal, precoxal setae absent; anterior part of
sternopleural groove shallow, posterior part deep, incomplete.

Elytra short, relatively broad; striae impressed, pollinose, indistinctly punctate; intercalary stria abbreviated
posteriorly; apical tubercles inflated, strongly emarginate; sutural stria with complete row of four or five setae; parasutural
without setae; intercalary stria with complete row of six to eight setae; intratubercular stria with one or two setae near
base, two near apex; marginal stria with complete row of 12-14 setae; anterior medial angle of Interval III with one seta in
prominent pollinose pit; apical tubercle with eight or nine setae in prominent punctures; metasternum with median sulcus;
transverse sulcus of abdominal Sternum III entire, those of IV-VI interrupted in midline; tibiae without false spurs; female
with lateral pit in Sternum IV, smaller one in Sternum V; female with Sternum VI with submarginal groove greatly
expanded, forming deep concavity occupying posterior 0.33 of sternum, limited anteriorly by scarp, latter interrupted by
pair of notches which are convergent anteriorly defining narrow, trapezoidal median lobe (Fig. 189); male unknown.

The absence of setae from the parasutural stria link this species to C. rojasi. The latter
species has a shorter, broader head, and does not have Sternum VI modified in the female. The
form of Sternum VI in C. pala is similar to that of C. excavatum except that the median lobe is
much broader and more rectangular in the latter.

THE CAVICOLLE GROUP

This group resembles the guildingii group in having tufts of minor setae present on
Antennal Segments V-X. It differs strongly in having the anterior median pit greatly enlarged.
In all species except C. mathani the pit contains a prominent median tubercle. False spurs are
absent. The form of the anterior median pit is strongly similar to that of C. dubium, in the
insigne group, and the latter species might really be more closely related to the cavicolle group
than to C. insigne.

There are nine species in the cavicolle group. They are restricted to southern Central
America and northwestern South America, from Costa Rica to Ecuador, eastern Colombia,
and the western part of Amazonas State, Brazil.

Phytogeny. — Phylogenetic relationships within this group are not clear. Of the nine species,
both sexes are known in only three. Our tentative conclusions about relationships are reflected
in the key. This arrangement might be altered substantially when both sexes of all species have
been studied. The most distinctive species is C. mathani , which has the intercalary stria

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abbreviated and lacks the tubercle in the anterior median pit. Both of these features, however,
might be derived characters, and C. mathani might not be the sister group to the remaining
species. Similarly, C. humile, the only species to lack a median sulcus of the metasternum,
might have lost the sulcus secondarily, and might be close to C. cavicolle. We placed C.
centrale and C. validum together because of the similar arrangement of temporal setae and the
presence of pollinosity along the notopleural suture. However, C. centrale resembles C.
curvatum, C. humile and C. cavicolle in having a round anterior median pit, while C. validum
has a pit which combines a truncate anterior margin, as in C. crater , and sinuate margins, as in
C. foveolatum. We provisionally attribute similarities in the anterior median pit to convergent
evolution. It is possible that we are wrong, and that the similarities in chaetotaxy and
pollinosity between C. centrale and C. validum are themselves the result of convergence.
Further conclusions will have to await the collection of more specimens of this excessively rare
group, of which we have studied only about 15 specimens.

Clinidium ( sensu stricto) mathani Grouvelle 1903
(Figs. 182, 192, 193)

Clinidium mathani Grouvelle 1903: 131.

Clinidium ( sensu stricto ) mathani (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “St. Paulo d’Olivenca, Amazonas, M. de Mathan” (MNHN).
The locality is on the upper Amazon in Brazil, close to the border with Peru and Colombia.

Description. — Length 6.3 mm. Antennal stylet conical, very large, about 0.4 of length of Segment XI, apex blunt;
tufts of minor setae present on Segments V-X; basal setae on Segments VII-X; Segment I with subapical pollinosity; head
slightly longer than wide; clypeal setae present, frontal grooves shallow, effaced anteriorly; median lobe narrow, tip
opposite anterior 0.3 of eye; frontal space narrow; medial margins of temporal lobe oblique; convergent posteriorly, nearly
contiguous at distinct medial angles; temporal lobe broadly bordered by pilosity posteriorly; eye crescentic, relatively
broad, about 0.5 of length of temporal lobe; orbital groove complete; one temporal seta, in orbital groove behind eye; two
pairs of postlabial setae.

Pronotum elongate; length/greatest width 1.50; pronotum widest slightly anterior to middle; lateral margins curved;
base moderately narrowed, curved; apex strongly narrowed, truncate; median groove dilated, basal 0.33 shallow but broad;
middle 0.33 deep, broad, margins parallel, apical 0.33 occupied by anterior median pit, latter elliptical, without tubercle,
0.25 of width of pronotum; basal impression small, triangular, open posteriorly; discal strides elongate, curved, 0.60 of
length of pronotum; marginal groove visible in dorsal view; angular seta present; three or four marginal setae; notopleural
suture not pollinose; sternopleural groove broadly interrupted.

Elytra moderately elongate; striae deep, pollinose, very coarsely punctate; intercalary stria abbreviated posteriorly
(Fig. 193); intratubercular stria strongly dilated near apex; preapical tubercle strongly inflated, truncate at apex, tubercles
separated by combined widths of sutural intervals; sutural stria without setae; parasutural stria with complete row of four
setae; intercalary with complete row of six setae; intratubercular stria with one seta near apex; marginal stria with five
setae in apical 0.2; preapical tubercle with three or four setae; apical tubercle without setae; metasternum with median
sulcus; abdominal Sterna III- VI each with transverse sulci complete, not interrupted in midline; submarginal sulcus of
Sternum VI broad, curved, not joined to transverse sulcus (Fig. 192); male without ventral tooth on anterior femur,
without proximal tooth on anterior tibia; middle calcar small, acute; hind calcar small, narrowly triangular, apex obtuse.
Female unknown.

This species is recognized by the abbreviated intercalary stria and by the elongate elliptical
anterior median pit, without a median tubercle.

Range. — Amazon Basin. Vulcano and Pereira (1975b) record it from Cerro de Nairo,
Amapa Territory.

Clinidium ( sensu stricto ) humile new species
(Figs. 181, 190)

Clinidium cavicolle Chevrolat 1873b: 388 {pars )

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141

Type Material. — HOLOTYPE male, labelled: “ cavicolle , New Granada, Chev. type” (NMW). This specimen
matches one discussed in the original description of C. cavicolle Chevrolat, as a possible representative of “the other sex” of
C. cavicolle. It is a male, while the lectotype of C. cavicolle is a female, but the two are not conspecific.

Description. — Length 6.8 mm. Antennal stylet conical, relatively small, about 0.15 of length of Segment XI; tufts
of minor setae on Segments V-X; basal setae sparse on Segments IX, X; Segments I-IX with subapical pollinose rings;
head short, broad, scarcely longer than wide, width behind eyes nearly equal to width across eyes; clypeal setae present;
frontal grooves deep, entire; median lobe narrow, tip opposite anterior 0.33 of eye; frontal space narrow, sides parallel;
medial margin of temporal lobe narrowly pilose, posterior margin broadly pilose; eye crescentic, narrow; orbital groove
complete, broad; three temporal setae all posterior to eye; three or four pairs of postlabial setae.

Pronotum rather short, length/greatest width 1.39; pronotum widest slightly posterior to middle, lateral margins
strongly curved; base moderately narrowed, curved; apex strongly narrowed, truncate; median groove dilated; gradually
broader anteriorly to anterior medial pit; latter rounded anteriorly, sides evenly curved, with large round median tubercle;
greatest width 0.33 of that of pronotum; basal impressions small, triangular, open posteriorly; discal strides long, curved,
0.6 of length of pronotum; marginal groove visible in dorsal view; angular seta present; five to seven marginal setae;
notopleural suture not pollinose; sternopleural groove absent.

Elytra moderately elongate; striae deep, pollinose, punctate; intercalary stria slightly dilated posteriorly; preapical
tubercles scarcely inflated, rounded at apex, widely separated; sutural stria without setae; parasutural stria with one seta at
base; intercalary stria with three setae in anterior 0.3; intratubercular stria with one seta at base; marginal stria with four
setae in anterior 0.25, four setae in posterior 0.25; preapical tubercle without setae; metasternum without median sulcus;
abdominal Sterna III-V with transverse sulci rather narrowly interrupted in midline; transverse sulcus of Sternum VI
reduced to small oval pit, widely separated from submarginal sulcus (Fig. 190); Sternum VI with two pairs of setae;
femora with dorsal pollinosity; male with large ventral tooth on anterior femur, without proximal tooth on anterior tibia;
middle calcar small, triangular, acute; hind calcar slightly larger, subtriangular, obtuse. Female unknown.

This species is the only member of the group in which the metasternum lacks a median
sulcus. It differs from C. cavicolle, in having a shorter, broader head; dorsal sides of femora
pollinose, and Sternum VI with submarginal sulcus separate from transverse sulci. The last two
characters might be secondary sexual differences, rather than species differences, since C.
cavicolle is represented only by females and C. humile only by males.

Clinidium ( sensu stricto) curvatum new species
(Figs. 194, 203)

Type Material. — HOLOTYPE male, labelled: “Oroque, Colombia, Santander del Norte, 10-VI-1965, J. & B.
Bechyne leg.” (VEN). PARATYPES one broken female, same label as holotype (VEN).

Description. — Length 6.2 mm. Antennal stylet elongate, acute, 0.4 of length of Segment XI; tufts of minor setae
on Segments V-X; basal setae absent; Segment I with subapical pollinose ring; head slightly longer than wide; frontal
grooves narrow, deep; clypeal setae present; median lobe short, broad, triangular, tip anterior to anterior margin of eye;
frontal space very narrow, sides parallel; medial margin narrowly pilose; posterior margin broadly pilose; eye crescentic,
narrow; orbital groove complete; two temporal setae, one medial to posterior margin of eye, other posteriomedial to it; two
pairs of postlabial setae.

Pronotum moderately elongate, length/greatest width 1.52; widest slightly behind middle, lateral margins strongly
curved; base moderately narrow, curved; apex strongly narrowed, truncate; median groove narrow, almost linear except at
anterior median pit; latter 0.33 of length of pronotum, about 0.3 of width of pronotum, margins divergent nearly to apex,
there strongly narrowed, with very large round median tubercle; basal impressions small, rounded, open posteriorly; discal
strides long, curved, about 0.45 of length of pronotum; marginal groove fine, visible in dorsal view; angular seta present,
medial to hind angle; five or six marginal setae; notopleural suture not pollinose; sternopleural groove absent.

Elytra rather short; striae deep, pollinose, punctate; intercalary stria not abbreviated posteriorly; intratubercular stria
slightly dilated posteriorly; preapical tubercles moderately dilated posteriorly, widely separated; sutural striae without
setae; parasutural stria with one seta at base; intercalary stria with complete row of six setae; intratubercular stria with one
seta at base, three near apex; marginal stria with complete row of about 1 5 setae; apical tubercle with one or two setae;
metasternum with median sulcus; abdominal Sterna III-V in both sexes with transverse sulci broadly interrupted in
middle; in female, large lateral pit on Sternum IV, smaller on Sternum III; Sternum VI of male with short, slightly oblique
transverse sulci, narrowly separated from submarginal sulcus, three or four pairs of setae; female with each transverse
sulcus of Sternum VI broken into two pits (Fig. 203); two pairs of setae; in both sexes, submarginal sulcus curved,
extending nearly to transverse sulci; middle, hind tibiae each with only one spur, femora not pollinose dorsally; male with
ventral tooth on anterior femur, without proximal tooth on anterior tibia; middle calcar triangular, apex obtuse; hind
calcar subtriangular, apex narrowly truncate; ventral margin with minute tooth anterior to spur.

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Bell and Bell

This species resembles C. cavicolle in general appearance, but differs in having shorter
discal strioles, distinct transverse sulci on Sternum VI and a narrower anterior median pit with
margins oblique and nearly straight.

Clinidium ( sensu stricto) foveolatum Grouvelle
(Figs. 195,204)

Clinidium foveolatum Grouvelle 1903: 130-131.

Clinidium ( sensu stricto) foveolatum (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE female, labelled: “Ecuador, Siemiradski 1882-1883, Clinidium foveolatum
Grouvelle, type” (MNHN).

Description. — Length 6.7 mm. Antennal stylet elongate, acute, 0.5 of length of Segment XI; tufts of minor setae
present on Segments V-X; basal setae present on Segments VI-X; subapical pollinose rings on Segments I-IX; head as long
as wide; clypeal setae absent; frontal grooves deep, rather broad; median lobe narrow, triangular, tip opposite middle of
eye; frontal space moderately wide, margins slightly convergent posteriorly; medial margin narrowly bordered with
pilosity; posterior margin widely bordered by pilosity; eye crescentic, rather short; orbital groove complete, rather broad;
three temporal setae, in orbital groove, one opposite posterior part of eye, one near occiput, one between them.

Pronotum oval, rather short; length/greatest width 1.36; widest near middle, lateral margins strongly curved; base
moderately narrowed, curved; apex very narrow, truncate; median groove with basal 0.5 moderately narrow, sides parallel;
apical 0.5 strongly dilated, margin sinuate, curved medially opposite tubercle of anterior median pit, dilated, rounded
anterior to constriction, tubercle transverse, oval; basal impression very small, triangular, open posteriorly; discal striole
short, 0.2 of length of pronotum; marginal groove visible in dorsal view; angular seta present; six or seven marginal setae;
sternopleural groove absent except near anterior margin.

Elytra elongate; striae deep, pollinose, indistinctly punctate; intercalary stria not abbreviated posteriorly;
intratubercular stria scarcely dilated near apex; preapical tubercles moderately dilated, widely separated posteriorly;
sutural stria without setae; parasutural stria with one seta near middle, one near apex; intercalary stria with complete row
of seven setae; intratubercular stria with one seta at base, one near apex; marginal stria with complete row of about 1 1
setae; apical tubercle with three or four setae; metasternum with median sulcus; transverse sulcus of abdominal Sternum
III not interrupted; those of IV-VI narrowly interrupted at midline; that of Sternum VI separated from submarginal sulcus
(Fig. 204); all transverse sulci coarsely punctate; female with shallow lateral pit on Sternum IV; middle, hind tibiae with
two small spurs; male unknown.

The very short discal strioles separate this species from all others except for C. centrale. The
latter species has the margin of the anterior median pit rounded. The shape of the anterior
median pit in C. foveolatum resembles that of C. spatulatum, but the latter species has a well
developed discal striole and a much smaller antennal stylet.

Clinidium ( sensu stricto) cavicolle Chevrolat 1873b
(Figs. 183, 191)

Clinidium cavicolle Chevrolat 1873b: 388.

Clinidium ( sensu stricto) cavicolle (Chevrolat) Bell and Bell 1978.

Type Material. — LECTOTYPE, here designated, female, labelled: “C. cavicolle, Colombia, Steinheil, Ocana
(Landolt)”, with red “typus” label (NMW). There is doubt as to whether this is the specimen described by Chevrolat, as
he gives the locality as “Nova-Grenata, Bogoto”. The latter is presumably a misspelling of “Bogota”. Nevertheless, this
specimen fits the original description better than the other syntype, described above as C. humile new species. The latter is
the specimen discussed by Chevrolat as “probably another sex” of C. cavicolle.

Description. — Length 6.0-7.0 mm. Antennal stylet conical, rather small, about 0.2 of length of Segment XI;
Segment XI somewhat compressed; tufts of minor setae on Segments V-X; basal setae present on Segments IX, X;
Segments I-X with subapical pollinose rings; head slightly longer than in C. humile, with lateral margins more parallel,
base more abruptly truncate; clypeal setae present; frontal grooves deep, entire; median lobe narrow, short, tip anterior to
eye; frontal space narrow, sides parallel; medial margin of temporal lobe narrowly pilose; posterior margin broadly pilose;
eye crescentic, narrow, rather short; orbital groove complete, broad; two temporal setae, one near posterior margin of eye,
the other near occiput; one or two pairs of postlabial setae.

Pronotum rather short, oval; length/greatest width 1.39; pronotum widest near middle, margins strongly curved; base
narrow, rounded; apex narrowly truncate; median groove dilated, margins parallel except in apical 0.25, opposite anterior
median pit; median groove pollinose near base, otherwise with margins pollinose, middle glabrous; anterior median pit

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143

rounded, about 0.33 of width of pronotum, lateral margins with long pilosity; median tubercle round; basal impression
small, triangular, open posteriorly; discal strides long, 0.6 of length of pronotum, strongly curved; marginal groove visible
in dorsal view; angular seta present, medial, anterior to hind angle; eight or nine marginal setae; notopleural suture not
pollinose; sternopleural groove incomplete.

Elytra moderately elongate; striae deep, broad, pollinose, punctate; intervals narrow, subcostate; intercalary stria not
abbreviated posteriorly; intratubercular stria slightly dilated posteriorly; preapical tubercle slightly dilated; sutural stria
without setae; parasutural stria with one seta at base, two near apex; intercalary stria with seven setae in complete row;
intratubercular stria with one seta at base, three or four near apex; marginal stria with complete row of about 1 5 setae;
apical tubercle with two or three setae; metasternum with complete median sulcus; abdominal Sterna III-V broadly
interrupted in midline; Sternum VI without transverse sulci, with submarginal sulcus long, curved, reaching nearly to
anterior margin (Fig. 191); one or two pairs of setae on Sternum VI; female with large lateral pit in Sternum III, smaller
one on Sternum IV; middle, hind tibiae each with one spur; femora not pollinose on dorsal surface. Male unknown.

The large, oval anterior median pit and the long, curved discal strioles make this species
resemble C. humile. The latter species, however, lacks the median sulcus on the metasternum,
has a broader head, and pollinosity on the dorsal surface of the femora.

Range. — Colombia. Hincks (1950) also lists it from Brazil, but without a definite locality.
We have been unable to find the source of this record. In addition to the lectotype, we have seen
two females, labelled: “Mesa Rica, Colombia, Santander del Norte, 2500 m., 2-VI-1965, J. &
B. Bechyne, leg.” (YEN).

Clinidium ( sensu stricto) crater new species
(Figs. 196, 205)

Type Material. — HOLOTYPE female, labelled: “PANAMA: Cerro Jefe, Azul Ridge, 9° 12' N, 79° 21' W,
700-750 m., cloud for., 20 May, 72, T. L., L. J. Erwin coll. Exped. #10, notebook #1, loose bark, log ADP01472”
(NMNH). PARATYPES two females, labelled: “PANAMA, Province of Panama, Cerro Jefe, 1000 m., 21-V-1977, coll.
Lloyd Davis, under dead bark, fallen hardwood” (UVM).

Description. — Length 5. 9-6. 9 mm. Antennal stylet near conical, small, about 0.2 of length of Segment XI,
slightly curved; Segment XI slightly compressed; tufts of minor setae on Segments V-X; basal setae on Segments VI-X;
subapical pollinose rings on Segments I-X; head slightly longer than wide; clypeal setae present; frontal grooves deep,
entire; median lobe short, narrow, tip anterior to eye; frontal space narrow, sides slightly divergent posteriorly; medial
margin of temporal lobe narrowly pilose; posterior margin very broadly pilose, to level of posterior margin of eye; eye
narrow, crescentic, rather short; orbital groove complete, rather broad; two temporal setae, one near posterior margin of
eye, other in middle of pollinosity near occiput; two pairs of postlabial setae.

Pronotum rather short, oval, length/greatest width 1.41; pronotum widest near middle, margins strongly curved, base
narrow, rounded; apex narrowly truncate; median groove broadened from base to apex; basal 0.2 narrow, parallel, margins
anterior to there oblique, divergent; anterior median pit over 0.5 of width of pronotum, margins sinuate opposite tubercle,
latter rounded; anterior margin of pit transverse; sides of anterior median pit long, pilose; basal impression small,
triangular, open posteriorly; discal strioles long, over 0.66 of length of pronotum, curved; marginal groove visible in dorsal
view; angular seta present, anterior, medial to hind angle; about nine marginal setae; notopleural suture not pollinose;
sternopleural groove incomplete.

Elytra moderately elongate; striae deep, broad, pollinose, truncate; intervals narrow, subcostate; intercalary stria not
abbreviated posteriorly; intratubercular stria slightly dilated posteriorly; preapical tubercles slightly inflated; sutural stria
without setae or with one or two setae near apex; parasutural stria with complete row of eight setae; intercalary stria with
complete row of eight setae; intratubercular stria with three setae near apex; marginal stria with about 1 5 setae; apical
tubercle with two or three setae; metasternum with median sulcus; abdominal Sternum III with transverse sulcus entire;
sulci of Sterna IV, V narrowly interrupted in midline; Sterna III, IV with shallow lateral pits in female; Sternum VI
without transverse sulci, submarginal sulcus shorter than in C. cavicolle restricted to posterior 0.5 of sternum (Fig. 205);
femora with dorsal surface pollinose; one spur on each middle, hind tibia. Male unknown.

The shape of the median groove of the pronotum in this species is closest to that of C.
validum. In the latter species, however, the anterior median pit is larger, the notopleural suture
is pollinose, and there are two tibial spurs.

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Clinidium ( sensu stricto) centrale Grouvelle 1903
(Figs. 197, 200)

Clinidium centrale Grouvelle 1903: 133-134.

Clinidium ( sensu stricto) centrale (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Costa Rica, C. centrale Grouv., type” (MNHN). Two
additional specimens, labelled as types (MNHN), must have been so marked by accident, as they bear collecting dates
later than 1903.

Description. — Length 6.0-7.4 mm. Antennal stylet slender, small, about 0.2 of length of Segment XI; tufts of
minor setae on Segments V-X; basal setae on Segments VII-X; subapical pollinose rings on Segments I-X; head scarcely
longer than wide, clypeal setae present; frontal grooves deep, entire, pollinose; median lobe short, broad, tip obtuse,
opposite anterior margin of eye; frontal space very narrow, sides parallel; medial margin of temporal lobe narrowly
pollinose; posterior margin very broadly pollinose, to level of posterior margin of eye; base of temporal lobe nearly
transverse, abruptly curved to lateral margin, latter nearly longitudinal behind eye; eye narrow, crescentic, rather short;
orbital groove complete; temporal lobe with one or two setae variously placed in large isolated pollinose punctures medial
to eye, three temporal setae in transverse row in pilosity of posterior margin; three pairs of postlabial setae.

Pronotum rather elongate, length/greatest width 1.58; widest near middle; margins strongly curved, base narrow,
curved; apex strongly narrowed, truncate; median groove moderately dilate, parallel-sided in basal 0.25; anteriorly, evenly
broadened to anterior medial pit; nearly evenly rounded anterior to pit; pit 0.33 as wide as pronotum; tubercle rounded;
sides of anterior median pit rather short, pilose; basal impression triangular, open posteriorly; discal stride short, about
equal in length to basal impression; length of impression plus stride about 0.33 of length of pronotum; marginal groove
visible in dorsal view; 1 angular seta, 8-11 marginal setae; notopleural suture pollinose (continuous in some specimens,
interrupted anterior to middle in others); sternopleural groove absent.

Elytra rather elongate; striae impressed, pollinose, punctate; intercalary stria not abbreviated at apex; apex of
intratubercular stria slightly dilate; preapical tubercle slightly inflated; sutural stria without setae; parasutural stria with
complete row of six or seven setae; intercalary stria with complete row of nine or 10 setae; intratubercular stria with one
seta at base, two near base; marginal stria with complete row of about 15 setae; preapical tubercle with one seta in
prominent puncture; apical tubercle with two or three setae; metasternum with median sulcus; transverse sulci of Sterna
III-VI entire in male; in female, sulcus of Sternum V narrowly interrupted, others entire; Sternum VI with submarginal
sulcus rather broadly separated from transverse sulcus (Fig. 200); Sternum VI with two or three pairs of setae; female with
deep lateral pit on Sternum IV; dorsal surface of femora with pollinosity; middle, hind tibiae each with two equal spurs;
anterior femur of male with prominent ventral carina; anterior tibia of male with proximal tooth; calcars acute, triangular.

The pollinose notopleural suture and isolated setose punctures on the temporal lobe separate
this species from all species except C. validum, which has a much larger, sinuate anterior
median pit. The discal strioles of the pronotum are only slightly shorter than those of C.
curvatum but the latter species has a smaller anterior pit and only one spur on each tibia.

Range. — Costa Rica. We have seen the following specimens with specific locality data: one
female, labelled: “Costa Rica, Cote de Tablazo, 1904, coll. P. Biolley” (MNHN); one specimen, sex not recorded,
labelled:“Sta. Maria de Dota, 1600 m., 1-1907” (MNHN) (The two preceding are incorrectly labelled as types.); two
males, four females, labelled: “Coronado, Costa Rica, VI-27- 1967, E. B. Fagan” (FLA); one female, same locality as
previous group but dated V-30-1967, elev. 5500 ft. (FLA). 20 males, 9 females, labelled: “Costa Rica: Cartago Prov., 5
km. S. El Empalme, VII- 14-73, J. Doyen & P. A. Opler Coll.” (UCB).

Clinidium ( sensu stricto) validum Grouvelle 1903
(Figs. 199, 202)

Clinidium validum Grouvelle 1903: 133.

Clinidium ( sensu stricto) validum (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Teffe (Ega, Amazonas, M. de Mathan 3me trimestre 1878, C.
validum Grouv.” (MNHN). The locality is in Brazil, several hundred kilometers west of Manaus.

Description. — Length 5. 8-6.4 mm. Antennal stylet slender, small, about 0.2 of length of Segment XI; tufts of
minor setae on Segments V-X; basal setae on Segments VI-X or VII-X; subapical pollinose rings on Segments I-X; head
slightly longer than wide; clypeal setae present; frontal grooves deep, entire, median lobe short, narrow, tip opposite
anterior margin of eye; frontal space moderately narrow, sides slightly divergent posteriorly; medial margin of temporal
lobe narrowly pilose; posterior margin very broadly pollinose to level of posterior margin of eye; eye crescentic, rather
broad of subgenus; orbital groove complete; one or two temporal setae medial to eye, in large punctures surrounded by
pollinose spots, latter in partial contact with pilosity of posterior margin; three or four smaller setae among long pilosity of

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145

occiput; three pairs of postlabial setae.

Pronotum rather elongate, length/greatest width 1.55, oval, widest near middle, sides curved; apex truncate, less
narrowed than in other members of group; base moderately narrowed, rounded; median groove dilated, posterior 0.5
parallel-sided except for slight dilation at posterior median pit; margins anterior to middle divergent to anterior median pit;
latter very large, margins oblique, divergent to level of tubercle; side suddenly broadened anterior to tubercle, apex broadly
rounded; margins of anterior median pit very long pilose; pit over 0.66 of width of pronotum opposite it; basal impressions
small, triangular, open posteriorly; discal stride very long, curved, 0.67 of length of pronotum; marginal groove visible in
dorsal view; angular seta present; eight to 10 marginal setae; notopleural suture pollinose; sternopleural groove incomplete.

Elytra rather elongate; striae impressed, pollinose, punctate; intercalary stria not abbreviated posteriorly;
intratubercular stria strongly dilated at apex; preapical tubercles strongly inflated, apex rounded, nearly contiguous;
sutural stria without setae; parasutural stria with complete row of six or seven setae; intercalary stria with complete row of
10 setae; intratubercular stria with one seta at base, three near apex; marginal stria with about 20 setae; preapical tubercle
with two setae; metasternum sulcate; transverse sulci of abdominal Sterna III-VI in male, very narrowly interrupted in
female; submarginal sulcus of Sternum VI rather broadly separated from transverse sulcus (Fig. 202); Sternum VI with
two to four pairs of setae; female with lateral pit on Sternum IV; dorsal surfaces of femora pollinose; middle, hind tibiae
with two equal spurs; male with anterior femur without ventral tooth or carina; male with prominent proximal tooth on
anterior tibia; calcars small, triangular, acute.

The greatly enlarged anterior median pit of this species resembles only that of C. crater in
shape. The latter species has the pit smaller, and the preapical tubercle of the elytron much less
prominent. In addition, the middle and hind tibiae have only one spur.

Range. — Widespread in the Amazon Basin. In addition to the type, we have seen one male,
three females from Ega (BMNH), and one specimen, sex not recorded, from Para (MNHN).
Vulcano and Pereira (1975b) record it also from Serro do Navio, Amapa Territory.

Clinidium (sensu stricto) spatulatum new species
(Figs. 198, 201)

Type Material. — HOLOTYPE female, labelled: “PANAMA:Colon Prov. Santa Rita Ridge, 300 m, 10-11,
VI-77, H.&A. Howden:” (BSRI).

Description. — Length 6.6 mm. Antennal stylet slender, small, about 0.2 of length of Segment XI; tufts of minor
setae on Segments V-X; basal setae on Segments VII-X; subapical pollinose rings on Segments I-X; head slightly longer
than wide; clypeal setae present; frontal grooves deep, entire, median lobe short, narrow, tip opposite anterior margin of
eye; frontal space moderately narrow, sides slightly divergent posteriorly; medial margin of temporal lobe narrowly
pillinose; posterior margin very broadly pollinose to level of posterior margin of eye; eye crescentic in lateral view, narrower
than in C. validum', orbital groove complete; four temporal setae as follows: one pair at posterior end of eye; one pair near
middle of temporal lobe opposite posterior margin of eye, in partially isolated tuft of pollinosity; two pairs posterior to eye;
two pairs of postlabial setae.

Pronotum rather elongate, length/greatest width 1.43, oval, widest near middle, sides curved; apex truncate, less
narrowed than in other species (except for C. validum)', base moderately narrowed, rounded; median groove dilated,
posterior 0.5 nearly parallel-sided except for slight dilation opposite posterior median pit; apical 0.5 strongly dilated,
margin sinuate, curved medially opposite tubercle of anterior median pit, strongly dilated both anterior and posterior to
constriction; tubercle nearly round, slightly transverse; basal impression small, open posteriorly; discal striole straight,
about 0.45 of length of pronotum; marginal stria visible in dorsal view; angular seta present; seven or eight marginal setae;
notopleural suture inconspicuously pollinose; sternopleural groove absent except for small pollinose spot near anterior
margin of prothorax.

Elytra rather elongate; striae impressed, pollinose, punctate; all striae complete; intratubercular stria dilated at apex;
preapical tubercle strongly inflated, apex rounded, well separated from opposite tubercle, medial margin appearing
“scalloped” by depressed pilose areas around setal punctures; sutural stria without setae; parasutural stria with complete
row of six setae; intercalary stria with complete row of 12 setae; intratubercular with one seta at base, and three near apex;
marginal stria with complete row of about 20 setae; preapical tubercle with two setae; metasternum sulcate; transverse
sulci of abdominal Sterna III-VI entire in female; submarginal sulcus of Sternum VI broadly separated from transverse
sulcus; Sternum VI with two pairs of setae (Fig. 201); female with minute lateral pit on Sternum IV; dorsal surfaces of
femora pollinose; middle, hind tibiae with two equal spurs; male unknown.

This species resembles C. foveolatum in having a very large anterior median pit with a
strong constriction opposite the tubercle. The latter species differs in the virtual absence of the
discal strides and in having a much larger antennal stylet. C. curvatum has discal strioles of
about the same length, as C. spatulatum but the strioles are curved, the anterior median pit is

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146

Bell and Bell

smaller with the constriction merely suggested, and the stylet is much larger. C. validum has a
large anterior median pit, but with the margin oblique posteriorly, not strongly sinuate, while
the discal stride is longer and the sternopleural groove is better developed.

THE BECCARII GROUP

These four species are unique in the tribe in entirely lacking minor setae on Segments IV-X.
As in all other Rhysodini, however, there are scattered minor setae on Segment XI. The
compound eye is either constricted at its middle or else entirely divided into two structures
resembling ocelli. The male has the midline of the anterior abdominal sterna modified with a
sulcus or paired tubercles. The antennal stylet is elongate and compressed, with the tip
obliquely truncate.

The range of the group is in doubt. C. moldenkei is the only species collected independently
by more than one collector. It is certainly from Costa Rica. There is no reason to doubt that the
single specimen of C. sulcigaster is from nearby Guatemala, as it was collected recently and
has exact data. The two remaining species are supposedly from the southwestern Pacific, C.
argus, from Negros, Philippine Islands, and C. beccarii from New Guinea. The holotype of C.
beccarii has exact locality data, and Beccari was a notably careful collector. According to Dr.
Poggi {in. litt.) not one of his specimens has previously been shown to be mislabelled. If it were
not for this, we would be almost certain that the specimen is mislabelled, as the distribution is
totally unexpected within a group of closely related species. Moreover, the entire subgenus is
otherwise strictly neotropical. It is still possible that Grouvelle or some other person
inadvertently interchanged labels between this and some other specimen. One should be
open-minded about the range of this group until more specimens of C. beccarii or C. argus
come to light, either from their supposed localities, or from somewhere in Central America.

Phytogeny. — The group clearly consists of two pairs of closely related species. In C.
moldenkei and C. sulcigaster , the eye is bilobed, the median groove of the pronotum is much
narrower than the anterior median pit, and the anterior part of the sternopleural groove is
absent. In C. beccarii and C. argus the eye is completely divided, the median groove is almost
as wide as the anterior median pit, and the anterior part of the sternopleural groove is present.

Clinidium ( sensu stricto) moldenkei new species
(Figs. 206,210,211,212)

Type Material. — HOLOTYPE male, labelled: “COSTA RICA, Rincon de Osa VII-15-66, A. R. Moldenke,
borrowed ex G. E. Ball ADP 38023” (NMNH). PARATYPES three males, two females, labelled: “Rincon de Osa,
Puntarenas, Costa Rica, 100 m., 3-X-1969, Halffter & Reyes” (MZSP).

Description. — Length 6. 8-7. 8 mm. Stylet elongate, 0.4 of length of Segment XI, compressed, apex obliquely
truncate; minor setae absent except for Segment XI, basal setae absent; Segments I-X each with subapical pollinose rings;
median lobe small, shield-shaped, frontal grooves broad, pollinose; temporal, antennal lobes separated by narrow
postantennal groove; temporal lobes slightly divergent posteriorly; medial, posterior margins of temporal lobes broadly
pilose; orbital groove complete; three temporal setae in transverse row near occiput; eye deeply bilobed, strongly
constricted at middle (Fig. 210); two pairs of postlabial setae.

Pronotum elongate; length/greatest width 1.61; sides curved; base moderately narrowed, curved; apex strongly
narrowed, truncate; median groove dilated, slightly constricted anterior to posterior median pit, closed at base; anterior
median pit four times wider than median groove at middle of its length; pit 0.36 of width of pronotum; basal impressions
open posteriorly; discal stride nearly straight, reaching middle of pronotum; marginal groove fine, visible in dorsal view;
angular seta present; eight or nine marginal setae; two or three basal setae just medial to basal impression; anterior 0.5 of
sternopleural groove absent, posterior part represented by three isolated pits (Fig. 211).

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

147

Elytra relatively short, broad; striae impressed, punctate; sutural interval depressed below level of others; intercalary
stria not abbreviated; intratubercular stria only slightly dilated near apex; preapical tubercle only slightly inflated; apical
tubercles more strongly inflated; sutural stria with one seta near apex, one seta in sutural interval posterior, medial to apex
of sutural stria; parasutural stria with complete row of eight setae; intercalary stria with complete row of eight setae;
intratubercular stria with three or four setae near apex; marginal stria with about 14 setae; apical tubercle with four or five
setae; metasternum not sulcate; transverse sulci glabrous, narrowed medially, medial end of each half with small deep pit;
sulci of Sternum VI pollinose, widely separated from one another, also from submarginal sulcus; female with slight lateral
pit on Sternum IV; male with pair of tubercles posterior to medial ends of transverse sulci on Sterna III, IV, without
median sulcus (Fig. 212); male without ventral tooth on anterior femur, without proximal tooth on anterior tibia; middle
calcar narrow, prominent, apex obtuse; hind calcar triangular, apex slightly obtuse.

This species is closest to C. sulcigaster, but differs in having the median groove of the
pronotum closed posteriorly, the posterior part of the sternopleural groove interrupted, and the
eye more strongly constricted. The male lacks the median sulcus on abdominal Sterna I-IV
which is characteristic of the latter species. We dedicate this species to the collector, Andrew
Moldenke, a dedicated coleopterist and our onetime co-worker on Vermont Carabidae.

Clinidium ( sensu stricto) sulcigaster Bell 1973
(Figs. 208,213)

Clinidium ( sensu stricto ) sulcigaster Bell 1973: 279-282

Type Material. — HOLOTYPE male, labelled: “GUAT. Finca Moca, Santa Barbara, Such., 3000', VI- 12- 1966,
J. M. Campbell” (BSRI, CNC no. 12,700). The locality is in Guatemala, Department of Suchitepequez, near Lake
Atitlan.

Description. — Length 5.5 mm. Stylet elongate, 0.5 of length of antennal Segment XI, compressed, apex slightly
oblique; minor setae absent except for Segment XI; basal setae absent; Segments I-IX each with subapical pollinose ring;
head longer than broad; median lobe small, shield-shaped; frontal grooves broad, pollinose; temporal, antennal lobes
separated by narrow postantennal groove; frontal space narrow; medial margins of temporal lobe parallel; medial margin
narrowly pollinose; posterior margin of temporal lobe broadly pilose; orbital groove complete; three temporal setae in
orbital groove; eye very small, heavily pigmented, less constricted at middle than in C. moldenkei\ three pairs of postlabial
setae.

Pronotum elongate, length/greatest width 1.68, widest slightly anterior to middle; sides curved anteriorly, oblique,
scarcely curved posteriorly; apex narrow, truncate; base moderately narrow, curved; median groove open posteriorly,
moderately dilated, about 0.1 of width of pronotum at middle; margins parallel in posterior 0.5 except for slight dilation at
posterior median pit, latter equidistant between middle and base of pronotum; anterior to middle, margins divergent to
anterior median pit, abruptly narrowed anterior to pit; anterior median pit about 0.33 of width of pronotum; basal
impressions narrow, open posteriorly; discal striole nearly straight, reaching nearly to middle of pronotum; marginal groove
fine, visible in dorsal view; angular seta absent; three marginal setae present in anterior 0.33 of marginal groove; one basal
seta just medial to basal impression; anterior 0.5 of sternopleural groove absent, posterior 0.5 entire.

Elytra relatively short, broad; striae impressed, punctate; sutural interval as convex as Interval II; intercalary stria not
abbreviated; intratubercular stria with basal 0.25 effaced; middle portion represented by row of punctures; apical 0.33
impressed; marginal stria entire; preapical tubercle slightly inflated; apical tubercles inflated, contiguous; sutural,
parasutural striae without setae; intercalary stria with one seta at base, one in basal 0.5 or absent, and two in apical 0.5;
intratubercular stria with two setae near apex; marginal stria with one seta near middle, six setae in apical 0.33; preapical
tubercle with two setae; metasternum not sulcate; transverse sulci interrupted medially, each 0.5 pollinose in medial 0.5,
glabrous in lateral 0.5; in male, abdomen with median sulcus bounded laterally by paired carinae; sulcus deep, distinct on
Sterna II, III, extending anteriorly to include medial part of Sternum I, posteriorly across Sternum IV to end on Sternum
V (Fig. 213); male without ventral tooth on anterior femur, without proximal tooth on anterior tibia; middle calcar
triangular, dorsal margin angulate, notched; hind calcar narrower, scarcely notched.

Female unknown.

This species differs from C. moldenkei in having the intratubercular stria incomplete
anteriorly, in having fewer elytral setae and a less constricted, smaller eye. Also, the male has a
median sulcus on the anterior 0.5 of the abdomen.

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148

Bell and Bell

Clinidium ( sensu stricto ) argus new species
(Fig. 207)

Type Material. — HOLOTYPE male, labelled: “Philippines, Horns of Negros”, date and collector not specified
(MCZ). The label is similar to that on the holotype of Omoglymmius ( sensu stricto ) crassicornis Bell and Bell. The label
on the latter specimen lists the collector as J. W. Chapman. The locality is a mountain on the Island of Negros. This
locality is at least questionable.

Description. — Length 6.6 mm. Stylet elongate, 0.4 of length of Segment XI, compressed, apex truncate; minor
setae absent except for Segment XI; basal setae absent; Segments I-X each with subapical pollinose ring; head 1.5 longer
than broad; median lobe rhomboid; frontal grooves broad, pollinose; antennal lobe small, triangular, separated from
temporal lobe by broad, pilose postantennal area; frontal space moderately broad; medial margins of temporal lobes
parallel; posterior margin of temporal lobe broadly pilose; orbital groove complete; four or five temporal setae in orbital
groove; eye divided into two portions resembling ocelli, anterior eye oval, posterior one smaller, round; two pairs of
postlabial setae.

Pronotum elongate; length/greatest width 1.72; widest near middle; sides curved; apex strongly narrowed, truncate;
base slightly narrowed, curved; median groove open posteriorly, strongly dilated, 0.16 of width of pronotum at middle;
margins parallel, anterior median pit only slightly wider than median groove; basal impression round, closed posteriorly,
but connected to lateral margin, median groove by depressed, pollinose areas; discal stride straight, extending to middle of
pronotum; marginal groove slightly dilated, visible in dorsal view; angular seta absent; eight to 10 marginal setae; three
basal setae medial to basal impression; sternopleural groove nearly complete, interrupted dorsad to coxa.

Elytra relatively long, narrow; sutural, parasutural striae impressed, narrow, conspicuously punctate; intercalary stria
wider, deeper than others; intratubercular stria fine, entire; marginal stria entire, strongly dilated posteriorly; preapical
tubercle scarcely inflated; apical tubercles strongly inflated, contiguous; sutural stria without setae; parasutural stria with
one seta at base, one or two in anterior 0.33; intercalary stria with two setae at base, one laterad to the other, complete row
of 10-11; intratubercular stria with four setae in apical 0.33; marginal stria with complete row of about 20 setae; apical
tubercle with three setae in conspicuous punctures; metasternum with incomplete median sulcus in anterior 0.5; abdominal
sterna with transverse sulci narrowly interrupted in middle; transverse sulci of Sternum VI narrowly separated from
submarginal sulcus; Sternum VI with one pair of setae; in male, abdominal Sternum III with median pollinose area, latter
continued onto Sternum II; Sternum IV with small median pollinose area; very small, inconspicuous pairs of tubercles
posterior to transverse sulci at midline on Sterna III, IV; male without ventral tooth on anterior femur; without proximal
tooth on anterior tibia; calcars large, strongly cultrate, curved anteriodorsally, apices recurved.

The divided compound eyes separate this species from all others except the closely related C.
beccarii, described, possibly erroneously, from New Guinea, and Rhyzodiastes ( Rhyzotetrops )
janus of Fiji. C. beccarii has the paramedian grooves much longer, and the calcars are
triangular.

Clinidium ( sensu stricto ) beccarii Grouvelle 1903 NEW COMBINATION

(Fig. 209)

Clinidium beccarii Grouvelle 1903: 140.

Rhyzodiastes beccarii (Grouvelle) Bell and Bell 1978.

Type Material. — HOLOTYPE male, labelled: “Nuovo Guinea: Hatam” (GEN). According to the original
description, collected by Beccarii. In 1978 we erroneously assigned this species to Rhyzodiastes based on the description.
We have not seen this specimen but now have studied detailed sketches of it, kindly supplied by Dr. Poggi.

Description. — Length 8 mm. Head longer than broad; median lobe small, rhomboid; antennal lobe small,
separated from temporal lobe by broad postantennal pollinose area; three temporal setae; eye divided into two ocellus-like
organs, latter only slightly separated.

Pronotum less elongate than in C. argus, length/greatest width about 1.55; median groove broadly dilated, margins
parallel; anterior median pit slightly wider than median groove; discal striole much longer than in C. argus , reaching
almost to anterior median pit.

Male with transverse band of pollinosity connecting transverse sulci in midline on Sternum III; Sternum IV-VI
without pollinosity in midline; paired tubercles near middle of Sterna III, IV; calcars triangular, not cultrate.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

149

ADDITIONS, CORRECTIONS TO PARTS I-III, INCLUDING ADDITIONAL SPECIES

Genus Dhy sores Grouvelle 1 903
Dhysores basilewskyi (Brinck 1965)

Through the courtesy of Dr. Roy Danielsson, we have been able to compare the series of
specimens from the former Belgian Congo (42 mi. N. of Kapona) (CAS) with a specimen
labelled as a paratype in the collection of the University of Lund. The latter, a male, is labelled:
“Tshuapa: Lac Tumba, Mabali, 350 m. (dans humus), N. Leleup 29-IX, 1955”. This specimen
and locality are not mentioned in the original description, and its status as a paratype is
doubtful. Unlike all previous specimens of the genus from the tropical zone in Africa, this one is
definitely from a lowland site, and implies that the genus is not limited to montane forests, as
we previously thought. It also shows that D. basilewskyi has the most extensive range of any
member of the genus.

Dhysores biimpressus new species
(Fig. 214)

Type Material. — HOLOTYPE male, labelled: “Usumbara, Neu Bethel, 10.3, 1905, coll. Jul. Moser” (MNHB).
PARATYPE one female, same data (MNHB). The locality is in Tanzania, formerly German East Africa. It is a small
mountain range, near Lushoto on the Kenyan border.

Description. — Length 6. 2-7. 2 mm. Anterior tentorial pits large, rounded; prefrontal pits entirely absent; frontal
space broader than in other members of genus, nearly as wide as long; frontal grooves scarcely impressed; two pairs of
postlabial setae; basal impressions about 0.6 of length of pronotum, anterior part of impression more abruptly narrowed
than in D. quadriimpressus ; hind angles of pronotum denticulate; one marginal seta on pronotum; elytral humeri not
especially narrowed, resembling those of D. basilewskyi (Brinck) (Bell and Bell, Part II, p. 383); Stria VI represented by
row of fine punctures, its anterior fourth effaced; Stria VII impressed except for anterior 0.15, where it is represented by
row of punctures; elytral setae unusually long.

In our key to species of Dhysores (Part II, p. 382), this species will trace to Couplet 2, where it will not fit either
alternative, since the anterior tentorial pits are large and round, while the prefrontal pits are entirely absent. The absence
of the prefrontal pits and the broader frontal space separate it from the sympatric D. quadriimpressus (Grouvelle). The
most similar species is probably D. thoreyi (Grouvelle), of South Africa, but in the latter species the anterior tentorial pits
are small and oblique, while the prefrontal pits are at least suggested, and the humeri are markedly narrowed.

Dhysores quadriimpressus (Grouvelle))

We have seen 14 additional specimens of this species (all in MNHB), eight from Neu
Bethel, the type locality for D. biimpressus , and six from Ost Usambara, coll. Methner. Dr.
Basilewsky has informed us that we confused the type locality, Usumbara, with Usumbura
(now Bujumbura) in Burundi. It is actually in northeast Tanzania and is the same as that of D.
biimpressus. We are grateful to Dr. Basilewsky for clearing up the confusion.

Genus Kupeus Bell and Bell 1982

This name was substituted for Kupea Bell and Bell 1978, which is preoccupied by Kupea
Philpott 1930.

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Kupeus arcuatus (Chevrolat 1873a) NEW COMBINATION

R. M. Emberson (personal communication) has pointed out that two of the localities listed
by us, Reefton and Springs Junction, are on the South Island of New Zealand. However he has
indicated that the Springs Junction label is an invalid one. If the Reefton record (BMNH) is
correct then this would be the only record of a rhysodine from the South Island.

GENUS KAVEINGA BELL AND BELL 1978
Kaveinga (sensu stricto) occipitalis (Grouvelle 1903)

Rhysodes occipitalis Grouvelle 1903: 105-106.

Type Material — LECTOTYPE male (here designated), labelled: “NUOVA GUINEA, Fly River, L. M.
D’Albertis 1876-1877” (GEN). PARALECTOTYPES two males, two females, same label as lectotype (GEN); one
female, same label as lectotype (MNHN). We erroneously listed the latter specimen as a holotype (Part 11:406).

The hind calcar of the male of K. occipitalis is very small and acute, similar to that of K. strigiceps Bell and Bell.

Kaveinga {sensu stricto) poggii new species
(Fig. 215)

Type Material. — HOLOTYPE male, labelled: “Is. Goodenough:Gennaio 1890, L. Loria” (GEN). This island is
one of the D’Entrecasteaux Group, north of the eastern tip of New Guinea.

Description. — Length 4.8 mm. Antennal Segment I pollinose dorsally; Segments II-V each with narrow pollinose
band; basal setae sparse on Segment VII, more numerous on VIII-X.

Head as long as wide, clypeus broadly separated from median lobe by band of pollinosity; parafrontal boss small,
nearly circular, separated from antennal rim and from median lobe by broad bands of pollinosity, and bordered posteriorly
by pollinose band; sides of median lobe broadly emarginate; orbital groove short, narrowed posteriorly, ending opposite
middle of eye; temporal lobe slightly wider than long; anteriomedial margins oblique, converging posteriorly; medial angle
obtuse, narrowly overlapped by median lobe; temporal setae one or two (right anterior one absent from holotype); anterior
seta in orbital groove; posterior one in round pollinose fovea in temporal lobe; postorbit entirely pollinose; temporal lobe
with distinct overhang in lateral view; suborbital tubercle and gular ridge absent.

Pronotum relatively short, broad, length/greatest width is 1.15; widest anterior to middle; sides strongly curved and
convergent between widest point and apex; sides oblique, slightly convergent from widest point to hind angles, margin not
sinuate anterior to hind angle; latter obtuse; shallow emargination present between hind angle and base; basal knob small,
depressed, pollinose; paramedian grooves deep, pollinose, width at middle nearly equal to that of outer carina; anterior end
of inner carina pollinose, so that glabrous area appears abbreviated anteriorly; posterior tip of inner carina acutely pointed;
marginal grooves broad; marginal seta absent; angular seta present; prosternum with shallow transverse groove between
precoxal carinae; latter not quite reaching anterior margin of pronotum.

Elytra moderately broad, slightly flattened; humeral tubercles not exerted; striae deep, pollinose; intervals convex,
subcarinate; strial punctures coarse, each puncture about 0.5 as wide as interval; Stria II with one basal seta, three setae in
apical fourth; Stria IV with six setae; apical stride without setae; several setae near apex of Stria VII; abdominal Sterna
III- V each with coarsely punctate, pollinose transverse sulcus, latter not interrupted at midline; femora with pollinose
bands; serrulation of middle tibia well developed; hind calcar of male slender, but its extreme tip narrowly truncate; female

unknown.

This species is named for Dr. Roberto Poggi of the Museo Civico di Storia Naturale “G.
Doria” of Genoa, in gratitude for the help he has given us during this study.

In our Key to Kaveinga s. str., this species will trace close to K. abbreviata. Therefore, the
key should be modified to read as follows:

3 (2) Pronotum relatively short, broad; length/greatest width 1.15 or less 3.1

3' Pronotum elongate, length/greatest width 1.2- 1.3 4

3.1 (3) Parafrontal boss small, round, separated from antennal rim and from
median lobe by broad pollinose bands; inner carina of pronotum acutely
pointed posteriorly K. poggii new species

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

151

3.T Parafrontal boss large, triangular, separated from antennal rim and from
median lobe by linear pollinose bands; inner carina of pronotum obtusely

pointed posteriorly K. abbreviata (Lea)

K. poggii clearly belongs to Group I, the species with deep, pollinose striae and subcarinate
intervals. The short pronotum makes it most similar to K. abbreviata , but it differs from the
latter in having the middle tibia strongly serrulate, the inner pronotal carina acutely pointed
posteriorly, and the parafrontal boss small and round.

The remaining members of the Group have the pronotum more elongate. K. fibulata differs
in having the hind angles of the pronotum rounded, while the four species of the K. pignoris
complex lack the parafrontal boss. K. poggii appears to be intermediate between K. abbreviata
and the remaining members of Group I, and makes it appear more likely that the group is
monophyletic.

GENUS GROUVELLINA BELL AND BELL 1978

Grouvellina hexadon new species
(Figs. 216, 220)

Type Material. — HOLOTYPE male, labelled: “COMORES, Mayotte Mamouzou, 13-8-69, s/ecorces, a la
lumiere, Y. Gomy” (GVA)

Description. — Length 4.8 mm. Antennal Segment XI slightly longer than wide; apical stylet short, acute; tufts of
minor setae on Segments V-X; antennal Segment I extensively pollinose in dorsal aspect; Segments II-X with two
transverse pollinose bands, basal band interrupted in Segments VIII-X; head as wide as long, frontal, postantennal grooves
deep, relatively narrow; median lobe rather narrow, broadly rounded posteriorly; parafrontal bosses narrow, rather long,
temporal lobe as wide as long, sinuate anterior to medial angles, latter narrowly separated, obtusely pointed; two temporal
setae; four labral setae; orbital groove complete, broadly pilose; two pairs of postlabial setae; mentum pollinose;
postmentum contrastingly glabrous.

Pronotum moderately long, length/greatest width 1.35; lateral margins nearly parallel; base and apex slightly
narrowed; outer carina not bent outwards at base; three or four marginal setae; prosternum without precoxal carinae
except for trace just anterior to coxa.

Elytral striae deep, very coarsely punctate; intervals broader than striae, not carinate; base of Interval II elevated,
forming small tooth (as in G. edentata)\ humerus prominent, quadrangular, with conspicuous patch of golden pilosity;
Stria I with two setae near apex; Stria II with eight setae; Stria IV with six setae; Stria VII with about nine setae in its
apical 0.2; metasternum entirely coarsely punctate without lateral pollinosity.

Male with ventral tooth on all femora; male with very minute prominent tooth on anterior tibia (Fig. 220); male with
hind calcar truncate at tip.

This species is smaller than any other member of the genus. In our key (Part II: 41 1-413), it
traces to couplet 6. The presence of a ventral tooth on all femora and the small size will
differentiate it from both species at this couplet. The absence of a precoxal carina is an
additional difference from G. tubericeps. It otherwise is almost a miniature of the latter species,
to which it appears to be related.

This species is not Rhysodes planifrons Fairmaire 1893, the only Rhysodine previously
described from the Comoro Islands. We have not been able to locate the type for the latter
species, which we suspect of being a Grouvellina , but the original description indicates that the
parafrontal bosses are united to the median lobe, and the length is given as 8mm.

GENUS YAMATOSA BELL AND BELL 1981

A misprint is present in the description of the genus (Part 11:424). Setae are present in apex
of Striae IV and VII or else are limited to apex of Stria VII.

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152

Bell and Bell

A specimen labelled “Java, J. D. Pasteur 268-94”, (MNHN) is labelled as a type of
“Rhysodes v. longior”. (now Yamatosa longior). It is not conspecific with the remaining types,
but is Omoglymmius ( Hemiglymmius ) rimatus Bell and Bell (Part III, p. 1 39).

The discovery of two additional species makes it necessary for us to revise our summary of
the phylogeny of this genus, and to alter the key to species. Y. kryzhanovskyi is perhaps the
most isolated species in the genus. It differs from all other species in having the prothoracic
pleuron and the disc of the metasternum coarsely punctate. The absence of the “beard” on the
labium links it to the “western” line, while the presence of the antennal stylet is a common
character with the “eastern” line. Y. kabakovi, on the other hand, clearly belongs to the
“eastern” line, and is closest to Y. niponensis.

KEY TO SPECIES

1 Prothoracic pleuron impunctate; punctures of metasternum limited to

margin 2

T Prothoracic pleuron densely punctate; metasternum with numerous

punctures on disc, in addition to row along each lateral margin

Yamatosa kryzhanovskyi new species

2 (1) Segment XI with distinct apical stylet; both eye and marginal groove of

pronotum fully developed 3

2' Segment XI of antenna obtuse, without apical stylet; either eye reduced or

else marginal groove of pronotum reduced 7

3 (2) Prosternum with distinct precoxal carina; discal stride ended at or

posterior to middle of pronotum 4

3' Prosternum without precoxal carinae; discal striole ended at, or anterior to

apical third of pronotum 5

4 (3) Precoxal carina extended more than 0.75 of distance from coxa to anterior

margin of prosternum; discal striole 0.5 of pronotal length

Yamatosa longior (Grouvelle) (Part II, p. 425)

4' Precoxal carina extended about 0.33 of distance from coxa to anterior

margin of pronotum; discal striole 0.33 of pronotal length

Yamatosa peninsularis (Arrow) (Part II, p. 427)

5 (30 Frontal and antennal grooves narrow, equal in width to posterior part of

clypeal grooves; posterior margins of frontal, antennal grooves sharply
defined; discal striole ended at or slightly anterior to middle of pronotum 6

5' Frontal and antennal grooves dilated, much wider than posterior part of
clypeal grooves; posterior margins of frontal and antennal grooves not
sharply defined; discal striole extended nearly to anterior margin of
pronotum Yamatosa arrowi (Grouvelle) (Part II, p. 428)

6 (5) Punctures of elytral striae I-V rounded, hind calcar of male triangular,

pointed Yamatosa niponensis (Lewis) (Part II, p. 427)

6 7 Punctures of elytral striae I-V elongate; hind calcar of male obtuse, with

dorsal “shoulder” Yamatosa kabakovi new species

7 (2') Marginal groove of pronotum absent except in basal fourth of pronotum;

eyes large, much deeper than long

Yamatosa reitteri (Bell) (Part II, p. 429)

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T Marginal groove of pronotum nearly complete, ended short distance from

anterior margin of pronotum; eyes more or less reduced 8

8 (7') Head evenly rounded posteriorly, widest point just posterior to eye; eye only
moderately reduced, deeper than long, with about 100 ommatidia

Yamatosa draco (Bell) (Part II, p. 429)

8' Head broadened posteriorly, widest point far posterior to eye; eye markedly

reduced, longer than deep, with about 50 ommatidia

Yamatosa boysi (Arrow) (Part II, p. 430)

Yamatosa kryzhanovskyi new species
(Figs. 217, 222)

Type Material. — HOLOTYPE male, labelled: “Vietnam, mts. NE Thai, Nguen, 12-1-1964, Kabakov” (LEN).
PARATYPE one female, same data as holotype (LEN).

Description. — Length 5.9-6.3 mm. Antennal Segment XI with slender, acuminate stylet; head cordate; anterior
tentorial pits rather small, punctiform; frontal grooves narrow, well defined; median lobe short, its tip rather broadly
truncate; eye large, deeper than long; mentum with a few punctures near middle, not “bearded” in either sex; one pair of
postlabial setae present.

Pronotum moderately elongate, length/greatest width 1.39; base only slightly narrowed; apex markedly narrowed,
discal stride long, extending about 0.67 length of pronotum; marginal groove complete; propleura sparsely, coarsely,
shallowly punctate (Fig. 222); prosternum with transverse band of punctures anterior to coxae; precoxal carinae absent.

Elytra relatively broad for genus, slightly flattened; strial punctures relatively coarse, close together, separated by less
than diameter of one of them; elytral intervals convex; Striae I-VI scarcely abbreviated at base; basal portions of V, VI
punctate but not impressed; Stria VII effaced in basal third, middle third represented by punctures but not impressed;
apical third impressed, punctate; setae absent from Stria IV; metasternum with punctures on disc as well as margins;
female with small, shallow lateral pit on Sternum IV.

Anterior femur with ventral tooth in both sexes; anterior tibia of male with medial groove, latter bounded both
anteriorly and posteriorly near base by pair of flanges; spurs of middle, hind tibiae nearly equal; hind calcar small, obtuse,
its tip just above level of spurs.

This species is unique within the genus in having the prothoracic pleura and the disc of the
metasternum punctate. It differs in addition from the sympatric Y. kabakovi in having the
elytral striae with coarse, crowded punctures, elytral Striae I-VI not abbreviated at base, the
mentum without a beard, the tip of the median lobe broadly truncate, and the pronotum
broader and less sharply narrowed to apex.

It is a pleasure to name this species for Dr. O. Kryzhanovsky, whose courtesy made it
possible for us to study specimens in the Leningrad collection.

Yamatosa kabakovi new species
(Figs. 218,223)

Type Material. — HOLOTYPE male, labelled: “Vietnam, mountains of Sha-Pa Province, 1600-2000 mm., 5.8,
1962, coll. O. N. Kabakov” (LEN).

Description. — Length 6.0 mm. Antennal Segment XI with apical stylet distinct, though small; head cordate;
anterior tentorial pits small, punctiform; frontal grooves narrow, well defined, median lobe short, its tip narrowly pointed;
eye large, deeper than long; mentum conspicuously punctate and “bearded” in male (female unknown); one pair of
postlabial setae present.

Pronotum elongate, narrow, length/greatest width 1.59; sides nearly parallel except anteriorly; apex distinctly
narrowed; base scarcely narrowed; discal stride ends slightly anterior to middle of pronotum; marginal groove of pronotum
complete; precoxal carina absent; prothoracic pleuron impunctate; prosternum impunctate, including precoxal area.

Elytra very narrow, cylindrical; Striae I-IV impressed, punctate, punctures longer than wide; Striae V-VII not
impressed, represented by fine, widely spaced, round punctures; Stria I with base entire; Striae II, III with base slightly
abbreviated; Striae IV, V effaced in basal 0.20; Stria VI effaced in basal 0.25 and also near apex; Stria VII effaced in
basal 0.33, its extreme apex impressed; setae confined to apex of Stria VII.

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Bell and Bell

Plate 18. Figs. 214-219. Head and pronotum, dorsal aspect. Fig. 214, Dhysores biimpressus new species; Fig. 215,
Kaveinga (s. str.) poggii new species; Fig. 216, Grouvellina hexadon new species; Fig. 217, Yamatosa kryzhanovskyi new
species; Fig. 218, Yamatosa kabakovi new species; Fig. 219, Arrowina punctatolineata (Grouvelle) (redrawn from sketch
by R. Poggi); Fig. 220, Anterior leg (excluding tarsus), male, Grouvellina hexadon new species; Fig. 221, Left elytron,
dorsal aspect, Arrowina punctatolineata (Grouvelle); Fig. 222, Prothorax, left ventrolateral aspect, Yamatosa
kryzhanovskyi new species; Figs. 223-224, Hind tibia, apex, male; Fig. 223, Yamatosa kabakovi new species; Fig. 224, Y.
niponensis (Lewis).

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Bell and Bell

Metasternum with row of punctures along lateral margin but without punctures on disc; metasternum of male
shallowly impressed; abdominal Sternum IV of male with shallow lateral pit (female unknown).

Anterior femur of male with ventral tooth (female unknown); anterior tibia of male with medial groove bordered by
small but distinct flanges near base; spurs of middle and hind tibiae nearly equal; hind calcar of male forming acute angle
above spurs, its proximal margin with distinct “shoulder”.

This slender, nearly cylindrical species has the elytral striation more reduced than in any
other member of the genus. It is closely related to Y. niponensis Lewis (Fig. 224) but differs
from the latter species in being narrower and more cylindrical with the pronotum more
elongate, and with the elytral striae more reduced. In the male, the form of the hind calcar will
separate the two species (Fig. 223).

This species is named for the collector, O. N. Kabakov.

Yamatosa longior (Grouvelle 1903)

In MNHN, there is an additional locality record, a specimen from Mt. Ardjoena, Java, Coll.
Mme. E. Walsh.

GENUS ARROWINA BELL AND BELL 1978

In Parts I and II, we did not include Rhysodes punctatolineatus Grouvelle, as we did not
know the location of the type, and were not able to learn the correct generic placement from the
original description. We have since discovered that it is in the Museo Civico di Storia Naturale
in Genoa. The curator, Dr. Roberto Poggi, has very kindly furnished us with detailed drawings
and notes which make it clear, as he indicated, that it belongs in Arrowina. The range of the
genus, as stated in Part I, p. 71, must be amended to read “Ceylon, southern India, Sumatra
and Japan”. Phylogenetically, A. punctatolineata is most closely related to A. taprobanae and
A. pygmaea. The latter two species are closer together than either is to A. punctatolineata ,
however.

REVISED KEY TO SPECIES (Supersedes that of Part II, pp. 438-439)

1 Head almost twice as long as wide; anterior femur of male with ventral

tooth (female unknown) Arrowina rostrata (Lewis) (Part II, p. 439)

1' Head only slightly longer than wide; anterior femur of male without ventral

tooth (male unknown in A. punctatolineata ) 2

2 (1) Orbital groove absent; lateral margin of inner pronotal carina sloped

gradually into paramedian groove 3

2' Orbital groove complete, somewhat dilated; lateral margin of inner carina

vertical, sharply defined 5

3 (2) Metasternum with a few punctures on anterior margin, otherwise

impunctate; eye reduced; elytral striae not impressed, represented only by
rows of punctures which become obsolete both near base and near apex .

Arrowina punctatolineata (Grouvelle)

3' Metasternum with row of coarse punctures along each lateral margin; eye

not reduced; elytral striae distinctly impressed, punctate from base to apex 4

4 (3') Metasternum with punctures confined to lateral margins; length 5.0 mm or

more Arrowina taprobanae (Fairmaire) (Part II, p. 439)

4' Metasternum with punctures in middle as well as along lateral margins;

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157

length 4.3 mm or less

Arrowina pygmaea Bell and Bell (Part II, p. 441)

5 (2') Eyes large, deeper than long; posteriomedial margin of temporal lobe

sinuate posterior to medial angles

Arrowina nilgiriensis (Arrow) (Part II, p. 441)

5' Eyes reduced, longer than deep; posteriormedial margin of temporal lobe
not emarginate Arrowina anguliceps (Arrow) (Part II, p. 442)

Arrowina punctatolineata (Grouvelle 1903) NEW COMBINATION
(Figs. 219, 221)

Rhysodes punctatolineatus Grouvelle 1903: 116.

Rhysodes punctatostriatus Grouvelle 1903: 97, 143 (error).

Rhysodes punctolineatus Hincks 1950: 11 (error).

Type Material. — HOLOTYPE female, labelled: “SUMATRA: Mte. Singalang” (GEN). (Specimen deformed
on right anterolateral margin of pronotum)

Description. — Length 6.8 mm. Antennal Segment XI slightly longer than wide; stylet minute, scarcely evident;
head slightly longer than wide; rostrum not elongate; median lobe elongate, pointed posteriorly; frontal space scarcely
evident; medial angles obtusely rounded, nearly contiguous; posteriomedial margin of temporal lobe rounded;
posteriolateral margin distinctly emarginate; orbital groove absent; eye reduced, longer than deep, strongly pigmented and
difficult to see; postorbital tubercle present, though very obtuse.

Pronotum elongate, length/greatest width about 1.3; widest at middle, sides evenly curved, apex and base both
markedly narrowed; inner carina slightly wider than outer one; inner carina with lateral margin ill-defined, sloped
gradually into paramedian groove.

Elytra with sides parallel in middle third; humeral region more narrowed than in A. taprobanae ; elytral striae not
impressed, represented only by rows of very fine punctures; strial punctures obsolete in basal 0.15 and apical 0.33; base of
elytron obliquely depressed, forming triangular depression in region of scutellum; elytron with three setae in posterior part
of Stria IV and several setae on lateral face of apical tubercle, and several setae in apex of Stria VII; metasternum with
row of punctures along anterior margin, otherwise impunctate (Fig. 221); middle and hind tibiae with spurs nearly equal.

This distinctive species differs from other known members of the genus in the reduction of
the elytral striation and in the triangular depression at the bases of the elytra. The reduction of
the eye is a feature in common with A. anguliceps , but it differs from the latter species in the
absence of the orbital grooves, as well as in the shape of the temporal lobes, and in the virtual
absence of the frontal space.

SUBGENUS PYXIGLYMMIUS BELL AND BELL 1978

We have found an additional species from Sumatra. In the key it would trace to O.
hesperus. The key can be modified as follows:

7 (6') Postorbital tubercles large, prominent; paramedian grooves relatively

shallow 7.1

7' Postorbital tubercles relatively small, not prominent in dorsal view;

paramedian grooves deep, more sharply defined

O. strabus (Newman)

7.1 (7) Elytral intervals flat; intervals, pronotal carinae, temporal lobes strongly
microsculptured in female, lateral pit of Sternum IV longitudinally striate,
brace weakly developed O. opacus new species

7.1' Elytral intervals convex; intervals, pronotal carinae, temporal lobes shining,
without microsculpture; in female, lateral pit of Sternum IV not striate.

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Bell and Bell

brace strongly developed O. hesperus Bell and Bell

Omoglymmius ( Pyxiglymmius ) opacus new species
(Figs. 225, 234)

Type Material. — HOLOTYPE female, labelled: “Sumatra, Padang, 1890, Modigliani” (GEN).

Description. — Length 9.0 mm. Antennal Segment XI slightly wider than long, tip obtuse; basal setae apparently
absent (though possibly lost from holotype); head slightly longer than wide, large relative to pronotum, as in O. strabus ;
clypeus punctate, pollinose, continuous with median lobe; latter rhomboid, wider than long; posterior angle obtuse,
anteriomedial margin of temporal lobe oblique; first medial angles distinctly, though narrowly separated; second medial
angles contiguous; occipital angles very obtuse though distinct; posteriolateral margin of temporal lobe slightly oblique;
each temporal lobe with two coarse punctures; temporal lobes coarsely microsculptured, opaque; postorbit concave dorsad
to postorbital tubercle, postorbital tubercle large, 0.7 as long, 0.6 as deep as eye; tubercles less divergent than in O.
hesperus, width across tubercles slightly greater than width across eyes.

Pronotum short; length/greatest width 1.26; widest point near middle, apex less strongly narrowed than in O.
hesperus', base moderately narrowed; lateral margin scarcely sinuate anterior to hind angles; paramedian grooves broad,
shallow, punctate; paramedian groove equal in width to inner carina at middle, narrower than outer one at middle; both
carinae strongly microsculptured, coarsely, densely punctate; marginal groove fine, about 0.20 as wide as outer carina at
middle.

Elytra longer, narrower than O. hesperus, but shorter than in O. strabus ; strial punctures relatively coarse; one seta in
apical 0.33 of Stria IV; subapical striole with seta; about four in apical 0.5 of Stria VII; elytral intervals strikingly flat,
densely microsculptured; female with deep lateral impression in posterior 0.67 of metasternum; female with margin of
elytron angulate opposite hind coxa; female with indistinct lateral pits on Sternum I, large lateral pits on Sternum IV,
latter separated medially by about twice width of one of them; lateral pit with slight trace of anteriolateral brace, with
elongate punctures suggesting longitudinal striation.

Male unknown.

The heavy microsculpture separates this species from O. strabus and O. hesperus. The very
flat elytral intervals are also distinctive. In the proportions of the body and size of postorbital
tubercles, it is intermediate between O. strabus and O. hesperus. The lateral pits of Sternum IV
differ from either. O. hesperus has a very strong brace but no trace of longitudinal striation,
while O. strabus has a prominent brace and strong longitudinal striation.

SUBGENUS ORTHOGLYMMIUS BELL AND BELL 1978
Omoglymmius ( Orthoglymmius ) feae (Grouvelle 1895b)

Dr. Poggi has kindly allowed us to study an additional specimen from the Genoa Museum
also labelled as a type. We hereby designate the specimen cited as holotype in Part III (female,
labelled: “Burma. Charin Cheba. 900-1100 m., X-88, coll. L. Fea” (MNHN)) as
LECTOTYPE. The PARALECTOTYPE is a male with same data as lectotype (GEN). It has
a pollinose postorbital tubercle, and is possibly not conspecific with the lectotype. Additional
material is needed in this difficult subgenus.

SUBGENUS OMOGLYMMIUS SENSU STRICTO GANGLBAUER 1892

Three new species O. gressitti, O. craticulus and O. largus are described below.

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Omoglymmius ( sensu stricto) gressitti new species
(Fig. 229)

Type Material — HOLOTYPE male, labelled: “PAPUA NEW GUINEA, Wau, Mt. Missim 1500 m. Feb. 25,
1982, R. T. Bell” (BPBM).

Description. — Length 7.9 mm. Antennal segments I-IV coarsely punctate; outer segments with punctures very
Fine; Segment XI impunctate; head slightly longer than wide; median lobe broad, apex broadly rounded; frontal space
broad, U-shaped, its lateral margins strongly curved; medial angles rounded, well separated; posteriomedial margin
oblique, sightly concave; posteriolateral margin nearly evenly curved; occipital angle obtuse; antennal lobes glabrous;
posteriomedial margin, occipital angle pollinose; orbital groove narrow, very short, ended anterior to middle of eye;
temporal lobe with about 25 rather coarse punctures; one temporal seta; small postorbital tubercle present; eye large,
round.

Pronotum short, length/greatest width 1.07; widest near middle; base slightly narrowed; apex strongly narrowed;
lateral margins strongly curved; margin slightly sinuate anterior to hind angle; inner carina almost twice as broad as outer
carina at middle; outer carina widest at middle, narrowed to apex; narrowed nearly to base, but broadened at extreme
base; outer carina relatively narrow, lateral and medial margins nearly parallel, so carina of nearly even width, strongly
curved; inner carina impunctate; outer carina with about 40 moderately Fine punctures; pronotum without setae;
prosternum without precoxal carinae.

Elytron moderately long, narrow; striae not impressed; strial punctures coarse; base of Stria IV with longitudinal
scarp; transverse basal scarp shining, not pollinose; subapical stride with one seta; Stria VII with one or two setae near
apex; metasternum bluish, opalescent, punctate in midline, along margins, part of disc impunctate; abdominal Sterna
III- V with punctures, Fine, nearly in single line near midline, scattered, coarse near lateral margins; male with rather deep,
semicircular lateral pits on Sternum IV; male with small ventral tooth on anterior femur; middle calcar minute; hind
calcar larger, obtuse.

This is a large species with a minute postorbital tubercle. It is similar to O. follis , also found
near Wau, but differs in having a very narrow, heavily punctate outer carina. This species is
dedicated to the memory of J. L. Gressitt and his wife, Margaret, for their kind hospitality and
assistance on our field trip to the Wau Ecological Institute, Papua New Guinea.

In our world key, this species would trace to couplet 69. The key should be changed as
follows:

69 (68) (unchanged) O. quadraticollis (Arrow)

69' (unchanged) 69.1

69.1 (69') Outer carina narrow, curved, densely punctate . . O. gressitti new species
69.1' Outer carina not conspicuously narrower than inner carina, sparsely

punctate or impunctate 70

In our key to species from New Guinea, this species would trace to couplet 18 and should be

changed as follows:

18 (17) Outer carina relatively narrow, curved O. gressitti new species

18' Outer carina relatively broad, less curved 18.1

(Couplet 18 of the original key is to be renumbered as 18.1)

Omoglymmius {sensu stricto) craticulus new species
(Figs. 227, 232)

Type Material. — HOLOTYPE female, labelled: “N. Guinea, S. E., Moroka 1300 m., Loria, VII-XI, 93”
(GEN). It had been labelled as R. capito Grouvelle.

Description. — Length 7.0 mm. Antennal Segments I-X punctate, outer segments finely so; Segment XI with a
few punctures; head slightly longer than wide; median lobe short, broad, tip rounded; frontal space slightly wider than
long, its margins curved; medial angles rectangular; posteriomedial margin emarginate; occipital angle very obtuse;
posteriolateral margin more strongly, evenly curved, than in O. planiceps, orbital groove fine, extending nearly to posterior

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Bell and Bell

Plate 19. Figs. 225-234. Genus Omoglymmius. Figs. 225-231, Head and pronotum, dorsal aspect; Fig. 225, O.
(Pyxiglymntius) opacus new species; Fig. 226, O. (s. str.) largus new species; Fig. 227, O. (s. str.) tolai new species; Fig.
228, O. (s. str.) craticulus new species; Fig. 229, O. (s. str.) gressitti new species; Fig. 230, O. ( Laminoglymmius )
perplexus new species; Fig. 231, O. (Navitia) peckorum new species; Figs. 232-233, Head, left lateral aspect; Fig. 232, O.
(s. str.) craticulus new species; Fig. 233, O. (Laminoglymmius) perplexus new species; Fig. 234, Metasternum, abdomen
left lateral aspect, O. ( Pyxiglymmius ) opacus new species.

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Bell and Bell

margin of eye; temporal lobe with about 20 very fine punctures; one temporal seta, arising from puncture at margin of
temporal lobe posterior to eye; posterior face of temporal lobe without pollinosity, but with area of strong microsculpture
resembling grid; temporal lobe relatively convex in lateral view; postorbital tubercle short but very deep; eye large, round.

Pronotum very short, length/greatest width 1.05; widest near middle; base slightly narrowed; apex strongly narrowed;
lateral margins slightly curved posteriorly; strongly curved, narrowed anteriorly; margin scarcely sinuate anterior to hind
angle; marginal groove not dilated; in dorsal view, outer carina appears about 0.6 as wide as inner carina at middle; outer
carina convex, directed dorsolaterad so it appears narrower in dorsal than in dorsolateral view; medial margin of outer
carina shallowly sinuate just anterior to base; outer carina widest at base; inner carina strongly narrowed at base; inner
carina impunctate; outer carina with many exceedingly fine punctures, not evident except under high magnification;
pronotum without setae; prosternum without precoxal carinae.

Elytra moderately long; striae not impressed, represented by rows of round, relatively coarse punctures; base of Stria
IV with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical striole with one seta; marginal stria with
apex impressed, with three or four setae; metasternum with broad medial, lateral bands of punctures, surrounding elongate
impunctate space on either side of disc; abdominal Sterna III-VI with many scattered punctures; female with moderately
deep, round lateral pit on Sternum IV; female without ventral tooth on anterior femur; male unknown.

The grid of microsculpture on the posterior face of the temporal lobe separates this species
from all except O. planiceps Bell and Bell. The latter species has much finer, sparser strial
punctures which are elliptical, rather than round. In addition, the temporal lobes are much
more strongly flattened than in O. craticulus and the pronotum is shaped differently.

This species would trace to Couplet 19 in our key to Omoglymmius sensu stricto of New

Guinea. The punctation of the metasternum is equivocal, so the key should be altered as

follows:

19 (17') Posterior face of temporal lobe pilose or scaly; temporal seta not marginal . . . 19.2

19' Posterior face of temporal lobe with microsculpture in grid pattern;

temporal seta marginal 19.1

19.1(190 Strial punctures relatively large, round, separated from neighboring
punctures by about 0.5 of length of one of them; temporal lobe relatively

convex O. craticulus new species

19.1' Strial punctures small, elliptical, separated by more than length of one of
them; temporal lobe strongly flattened O. planiceps Bell and Bell

19.2(19) Metasternum with punctures limited to midline, margins 20

19.2" Metasternum with punctures scattered over entire disc 23

Omoglymmius ( sensu stricto ) largus new species
(Fig. 226)

Type Material. — HOLOTYPE female, labelled: “NOUVA GUINEA, Fly River, L. M. D’Albertis, 1876-77”
(GEN). The specimen also bears a pink label “6880”.

Description. — Length 7.2 mm. Antennal Segments V-X coarsely punctate; Segment XI missing in holotype; head
slightly broader than long; median lobe lance-shaped, broader anteriorly than in O. capito, tip obtuse; frontal space
broader than long, lateral margin shallowly sinuate; medial angles rounded, more widely separated than in O. capito ;
posteriomedial margin curved into posteriolateral margin; occipital angle absent; antennal groove rather narrow, not
expanded laterally; orbital groove shallow, ill-defined; temporal lobe with about 10 rather coarse punctures, not clearly
differentiated into coarse and fine ones, as in O. capito ; one temporal seta; postorbital tubercles about 0.8 as long as eye,
more divergent than in O. capito , width across them much greater than width across eyes; eye large, round; posterior face
of temporal lobe with minute pale scales which are separated from one another.

Pronotum very short, broad; length/greatest width 1.03, widest near middle; base slightly narrowed; apex more
strongly narrowed; anterior part of lateral margins more oblique, less curved than in O. capito; margin slightly sinuate
anterior to hind angle; outer carina about 0.5 as wide as inner carina at middle; outer carina narrow, convex, of nearly even
width; inner carina narrowed just anterior to base, then slightly dilated; paramedian groove broader than in O. capito, apex
of paramedian groove dilated; outer carina with about 30 fine punctures; inner carina impunctate; pronotum without setae;
prosternum without precoxal carina.

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Elytron rather broad, its surface microsculptured, shining, opalescent; striae not impressed, represented by rows of
rather coarse, round punctures; base of Stria IV with longitudinal pollinose scarp; elytral striae without setae;
metasternum nearly completely, finely punctate, but with punctures very sparse on either side of midline; female with oval
lateral pits on abdominal Sternum IV; female with acute ventral tooth on anterior femur; male unknown.

This species has a broad head and divergent postorbital tubercles. It is close to O. capito, but
differs in the shape of the pronotum, especially in having much narrower outer carinae.

O. largus will trace to O. capito in our key to Omoglymmius s. str. of New Guinea. Couplet
24' should be altered to lead to Couplet 25, which will separate the two species as follows:

25 (24') Outer carina slightly narrower than inner carina at middle; outer carina at

anterior 0.33 broader than paramedian groove O. capito (Grouvelle)

25' Outer carina about 0.5 as wide as inner carina at middle; outer carina at
anterior 0.33 narrower than paramedian groove. O. largus new species

Omoglymmius ( sensu stricto ) tolai new species
(Fig. 231)

Type Material. — HOLOTYPE male, labelled: “NEW BRITAIN, Rabaul, 17-VII-79 sur arbre mort, J. D.
Bourne” (GVA). PARATYPE one female, mounted on same pin as male.

Description. — Length 6. 7-7. 2 mm. Antennal Segments I-IV coarsely punctate; Segments V-X more finely
punctate; Segment XI impunctate; head distinctly longer than wide; median lobe short, oval, its tip broadly rounded;
median lobe impunctate; frontal space as long as broad, nearly V-shaped, its anterior medial margin oblique, long; medial
angles nearly rounded, markedly separated; posteriomedial margin curved evenly; posteriolateral margin evenly curved;
occipital angle indistinct; orbital groove narrow, ended posterior to middle of eye; anterior portion of temporal lobe a
convex, pollinose ridge; temporal lobe with 10-28 fine punctures; one temporal seta; postorbital, suborbital tubercles
absent; eye large, round.

Pronotum rather short; length/greatest width 1.14, widest near middle; base slightly narrowed; apex markedly
narrowed, margin evenly curved from middle to apex; margin scarcely sinuate anterior to hind angle; outer carina about
0.67 as wide as inner carina at middle; medial margin of outer carina sinuate just anterior to base; outer carina widest at or
posterior to middle, scarcely narrowed anteriorly except at extreme apex; inner carina narrowed to base; latter truncate;
outer carina with 20-38 fine punctures; inner carina with 21-23 fine punctures; pronotum without setae; prosternum
without precoxal carinae.

Elytron relatively elongate, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria
IV with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with
several setae near apex; metasternum largely punctate, but with small impunctate area near middle of disc; abdominal
Sterna III- VI coarsely punctate; punctures confluent near lateral margin; female with lateral pit on Sternum IV small but
relatively deep, male with similar but shallower pit; both sexes with ventral tubercle on anterior femur, that of female
relatively small; middle calcar small, obtuse; hind calcar triangular, proximal margin slightly concave.

This species, the first of the genus to be described from the Bismarck Archipelago, is close to
several species from New Guinea. In the shape of the pronotum it comes close to O.
puncticornis Bell and Bell. It differs from the latter species in having the punctures of the distal
antennal segments markedly finer than those of the proximal segments.

The punctures of the legs and ventral surface are also notably finer than in O. puncticornis.
O. fringillus Bell and Bell differs in having the temporal lobes more transverse, with the
posteriomedial angles longer and more oblique, so that the most posterior points on the two
lobes are separated by more than 0.5 of the greatest width of the head. The pronotum is more
nearly quadrate, with the lateral margins nearly parallel except near the anterior margin. The
closest species is perhaps O. oroensis Bell and Bell, which resembles O. tolai in the shape of the
temporal lobes, but which has the pronotum similar to O. fringillus.

In our general key, O. tolai will trace to Couplet 20. At this point the key should be altered
as follows:

Quaest. Ent., 1985,21 (1)

164

Bell and Bell

20 (19) Pronotum subquadrate, lateral margins convergent only near apex 20.1

20' Pronotum with lateral margins curved, convergent from middle to apex 21

20.1(20) Most posterior points on temporal lobes separated from one another by

much less than 0.5 of width of head

O. oroensis Bell and Bell (Part III: 240)

20. T Most posterior points on temporal lobes separated from one another by

more than 0.5 of width of head

O.fringillus Bell and Bell (Part III: 240)

(some specimens: see below)

21 (20') Medial angle of temporal lobe obtusely pointed; posteriomedial margin

slightly sinuate; strial punctures elliptical, fine, sparse

O. viduus Bell and Bell (Part III: 226)

21' Medial angle rounded; posteriomedial margin rounded; elytral punctures

coarse 21.1

21.1(21) Antennal Segments V-X as coarsely punctate as Segments I-IV; legs,

ventral surface very coarsely punctate. (New Guinea)

O. puncticornis Bell and Bell (Part III: 241)

21. 1' Antennal Segments V-X more finely punctate than Segments I-IV; legs,

ventral surface more finely punctate (New Britain)

O. tolai new species

The title of the regional key should be altered to read “Key to species from New Guinea, the
Admiralty Islands and the Bismarck Archipelago”. In this key, the species would key to O.
puncticornis (80- The latter species can be separated from O. tolai by using couplet 21.1 of the
general key (above).

O.fringillus Bell and Bell 1982

We have studied a series of four males and two females labelled “XII-78, PNG (Morobe)
umg. Kaiapit” (GVA). These fit the original description of the species except that the lateral
pollinosity of the temporal lobe is interrupted for a short distance posterior to the level of the
eye. This is true also of a few in the type series, so the character should not be used in the keys.
The general key above has been altered to correspond to this inconsistency.

SUBGENUS LAMINOGL YMMIUS BELL AND BELL 1982
We have found an additional species from Sumatra. The key should be modified as follows:

4 (2') Outer carina punctate, at least near margin 4.1

4' Outer carina impunctate 5

4.1(4) Median lobe concave; two medial angles, separated by shallow

emargination O. insularis (Grouvelle)

4.1' Median lobe flat; one medial angle, anteriomedial margin oblique

O. perplexus new species

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

165

Omoglymmius ( Laminoglymmius ) perplexus new species
(Figs. 228, 233)

Type Material. — HOLOTYPE female, labelled: “SUMATRA, dono Grouvelle 1901, philippensis Chev., teste
Grouvelle 1901” (GEN). The specimen was formerly mounted on the same pin as a male of Omoglymmius
( Hemiglymmius ) inermis Bell and Bell.

Description. — Length 7.1 mm. Antennal Segment XI as wide as long, apex rounded; basal setae sparse on
Segments V-VI denser on VII-X; clypeus impunctate, continuous with median lobe; latter impunctate, narrow, tip pointed,
not translucent; anteriomedial margin of temporal lobe almost straight, translucent area semicircular, less sharply
different from remainder of temporal lobe than in O. inaequalis-, one medial angle, latter obtusely rounded, nearly
contiguous with that of opposite temporal lobe; medial angle with very narrow pollinose area; posteriomedial margin nearly
rounded; temporal lobe convex, shining, with two or three coarse punctures, each with minute seta; postorbital tubercle
small, about 0.5 as deep, 0.33 as long as eye, low, opposite lower 0.5 of eye; width across postorbital tubercles less than that
across eye.

Pronotum relatively short, length/greatest width 1.10; widest point slightly anterior to middle; sides curved, strongly
convergent to apex, latter narrow; sides oblique, slightly convergent to base, latter relatively broad; margin scarcely sinuate
anterior to hind angle; paramedian groove deep, strongly narrowed anteriorly, width at middle 0.4 of that of inner carina;
outer carina broad, nearly equal to inner carina at middle; outer carina with five or six punctures near lateral margin in
middle 0.33; inner carina entirely impunctate; marginal groove linear.

Elytra relatively short, broad; striae shallow, intervals slightly convex, faintly microsculptured; strial punctures
relatively coarse, each about 0.33 of width of interval; Stria VII with three to five setae near apex; female with shallow,
semicircular lateral pit on Sternum IV; female with ventral tooth on anterior and posterior femora; male unknown.

The medial translucent area on the temporal lobe is very small and liable to be overlooked.
If so, this species would be traced to Subgenus Boreoglymmius. In the latter species, it would
be keyed to O. lewisi of Japan, because of the presence of a postorbital tubercle. The latter
species differs in the conspicuously oblique posteriolateral margins of the temporal lobes, and in
the much bigger lateral abdominal pits of the female. Also, it lacks a median gular tubercle,
while O. perplexus has one. This appears to be a constant difference between the Subgenera
Laminoglymmius and Boreoglymmius. Another superficially similar species is O. lineatus of
southern India, in Subgenus Indoglymmius. The latter species lacks basal setae on the
antennae, lacks a postorbital tubercle, and has a much more elongate pronotum.

With the Subgenus Laminoglymmius, the most similar species are O. inaequalis of the
Andaman Islands, and O. actae of New Guinea. Both species have much narrower outer
carinae. In addition, O. inaequalis has a much more conspicuous translucent area on the
temporal lobe, and O. actae has two distinct medial angles.

O. inaequalis appears to be the species most closely related to O. perplexus. It appears to us
that the two shared a common ancestor more recent than our hypothetical Species 3 (Part III,
Diagram 2).

SUBGENUS NAVITIA BELL AND BELL 1978
The discovery of an additional species necessitates the substitution of a new species key.

KEY TO SPECIES (Supersedes that in Part III: 164)

1 Frontal grooves pollinose 2

V Frontal grooves glabrous, scarcely evident

O. intrusus (Grouvelle) (Part III: 166)

2 (1) Temporal lobe with eight or more punctures in addition to setiferous

puncture; outer carina of pronotum with three to Five punctures near base

O. stylatus Bell and Bell (Part III: 165)

Quaest. Ent., 1985, 21 (1)

166

Bell and Bell

2' Temporal lobe with one or two punctures in addition to setiferous puncture;
outer carina of pronotum without punctures O. peckorum new species

Omoglymmius ( Navitia ) peckorum new species
(Fig. 230)

Type Material. — HOLOTYPE male, labelled: “FIJI: Viti Levu 1100 m., Nandarivatu Microw, 16-20 VIII -
1978 S & J Peck, Ber. Elfin for litter, rainforest, berlese litter” (BSRI). PARATYPE male, same label as holotype
(BSRI).

Description. — Length 5.5 mm. Antennal stylet more elongate than in related species, about 0.3 of length of
Segment XI; head cordate, slightly broader than long, temporal lobes slightly flattened; frontal grooves pollinose, broader
and deeper than in O. stylatus ; orbital groove distinct, reaching to middle of eye; head broader than in O. stylatus , margins
slightly convergent posteriorly; temporal lobe with three coarse punctures near margin, one or two of them with temporal
seta; temporal lobe otherwise impunctate.

Pronotum elongate, length/greatest width 1.23; widest anterior to middle, lateral margin distinctly sinuate anterior to
hind angle; marginal groove distinct in anterior 0.67-0.75, replaced posterior by group of five widely spaced punctures;
pronotal epipleuron without ventral row of punctures.

Elytra narrow, relatively elongate; last puncture of Stria III enlarged, elongate oval; striae deeper, more coarsely
punctate than in related species; intervals more convex than in related species; Stria IV with complete row of five setae; one
seta at apex of Stria VI; about five setae near apex of Stria VII; punctures of Sternum V not confluent laterally; male with
ventral tooth on anterior femur; middle calcar obsolete; hind calcar obtuse, proximal margin obtusely angulate. Female
unknown.

This species has deeper striae with coarser punctures and more convex intervals than in
other members of the subgenus. The antennal stylet is larger. In other respects it shows a
mixture of the characters of the previously known species. The pollinose frontal grooves give a
superficial resemblance to O. stylatus , while the relatively short marginal groove, and the
absence of punctures on the pronotal epipleuron and the outer carina are similarities to O.
intrusus. The hind wing was checked on the paratype, and was found to be fully developed.

We dedicate this species to the collectors, Drs. S. & J. Peck.

ACKNOWLEDGEMENTS

We wish to thank the numerous curators and collectors whose aid has made this study
possible. Most were mentioned in Parts I-III. Dr. Ivan Lobl, of the Museum d’Histoire
Naturelle, of Geneva, lent us the valuable Rhysodine collection under his care. We wish to
extend special thanks to Dr. George Ball for his editorial help, and for many other types of
assistance at all stages of this project. We are again indebted to Mrs. Joyce Murray for the
typing of the original manuscript. We thank Mrs. Ruth Goodridge and Ms. Gail Porteus for
typing the subsequent revisions. Finally, we acknowledge our debt to the late Dorothy A. Bell
(mother of Ross T. Bell) for many years of support and interest, and for the bequest of funds
which have made it possible to publish this work.

REFERENCES

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present known. The Proceedings of the Royal Entomological Society of London (B) 11:

171-183.

Bell, R.T. 1970. The Rhysodini of North America, Central America and the West Indies.

Miscellaneous Publications of the Entomological Society of America 6(6): 289-324.

Bell, R.T. 1973. A new species of Clinidium from Guatemala. Proceedings of the

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

167

Entomological Society of Washington 75(3): 279-282.

Bell, R.T. and J.R. Bell. 1975. Two new taxa of Clinidium from the eastern United States with
a revised key to U.S. Clinidium. The Coleopterists’ Bulletin 29(2): 65-68.

Bell, R.T. and J.R. Bell. 1978. Rhysodini of the world. Part I. A new classification of the tribe,
and a synopsis of Omoglymmius subgenus Nitiglymmius new subgenus. (Coleoptera;
Carabidae or Rhysodidae). Quaestiones Entomologicae 14(1): 43-88.

Bell, R.T. and J.R. Bell. 1979. Rhysodini of the world. Part II. Revisions of the smaller genera
(Coleoptera: Carabidae or Rhysodidae). Quaestiones Entomologicae 15(4): 377-446.

Bell, R.T. and J.R. Bell. 1981. Coleoptera: Rhysodidae. Insects of Micronesia 1 5(2):5 1—67.

Bell, R.T. and J.R. Bell. 1982. Rhysodini of the World. Part III. Revisions of Omoglymmius
Ganglbauer (Coleoptera: Carabidae or Rhysodidae) and substitutions for preoccupied
generic names. Quaestiones Entomologicae 18(1-4): 127-259.

Blanchard, R. 1889. Observations on some variations of the males in Clinidium. Psyche,
5:165-166.

Brinck, P. 1965. Coleoptera: Rhysodidae XIV, In Hanstrom, B., Brinck, P. and G. Rudebeck.
Results of the Lund University Expedition 1950-1951. Stockholm, Swedish Natural Science
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Broun, T. 1880. Manual of the New Zealand Coleoptera. Part 1: XIX, Wellington. 651 pp.
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Chevrolat, A. 1829, 1844. In Guerin-Meneville. Iconographie du regne animal de G. Cuvier.
Vol. 3. Insects, 1829-38 (1844). 576 pp. Paris.

Chevrolat, A. 1873. Descriptions de Rhysodides nouveaux et enumeration des especes decrites.
Annales de la Societe Entomologique de France 3(5): 207-218.

Chevrolat, A. 1875. Remarques et descriptions. Bulletin de la Societe Entomologique de France
5(5): 182-183.

Costa, O.G. 1839. Clinidium canaliculatum n.sp. Atti della Reale Accademia delle Scienze di
Napoli, 4:104.

Dajoz, R. 1975. Apropos des Coleopteres Rhysodidae de la faune europeenne. L’Entomologiste
31(1 ): 1— 10.

Fairmaire, L. 1868. Notes sur les Coleopteres recueillis par Ch. Coquerel a Madagascar et sur
les Cotes d’Afrique. Rhysodidae. Annales de la Societe Entomologique de France 8(4):
782-783.

Fairmaire, L. 1893. Coleopteres de lies Comores. Annales de la Societe Entomologique de
Belgique 37: 528.

Fairmaire, L. 1895. Descriptions de quelques Coleopteres de Madagascar. Ibid. 39: 10-11.

Ganglbauer, L. 1892. Die Kafer von Mitteleuropa. Familienreihe Caraboidea. Wien. I:
530-534.

Germar, E.F. 1840. Beschreibung der arten von Rhysodes von E. Newman, Esq.;aus dem
Magazine of Natural History, new series, December 1838, S. 663-667. Zeitschrift fur die
Entomologie, 2(2):342-352.

Grouvelle, A. 1895a. Clavicomes des lies de la Sonde et de L’Oceanie recoltes par M. Raffray.
Descriptions d’especes nouvelles de la collection du mseum. Bulletin de Museum d’Histoire
Naturelle, Paris. I: 157-158.

Grouvelle, A. 1895b. Viaggio di L. Fea in Birmania e regioni vicine. LXVI. Rhysodides. Annali
del Museo Civico di Storia Naturale di Genova. 34: 761-762.

Grouvelle, A. 1903. Synopsis des Rhysodides et descriptions d’especes nouvelles. Revue
d’Entomologie 22: 85-148.

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Grouvelle, A. 1910. Description de trois Rhysodides nouveaux (Col.) Bulletin de la Societe
Entomologique de France 1910: 324-327.

Heller, K.M. 1898. Neue Kafer von Celebes. Abhandlungen und Berichte des Koniglichen
Zoologischen und Anthropologisch-Ethnographischen Museums zu Dresden 7(3): p.3.

Hincks, W.D. 1950. Coleopterorum Catalogus, Supplementa, Paris I, Rhysodidae (Editio
seconda): 1-18.

Kirby, W. 1835. The characters of Clinidium, a new genus of insects in the order of Coleoptera,
with a description of Clinidium guildingii. The Zoological Journal 5: 6-9.

Lea, A.M. 1904. Descriptions of new species of Australian Coleoptera. Proceedings of the
Linnean Society of New South Wales 29: 60-107.

LeConte, J.L. 1875. Notes on Rhysodidae of the U.S. Transactions of the American
Entomological Society 5: 162-168.

Lewis, G. 1888. On the Family Rhysodidae. XI. Tha Annals and Magazine of Natural History
2(6): 76-85.

Miwa, Y. 1934. On a new species of Rhysodidae from Formosa. Transactions of the Natural
History Society of Formosa 24(133): 256-257.

Mulsant, E. 1853. Coleopteres, nouveaux on peu connus; Rhysodes sulcipennis. Opuscules
Entomologiques 2: 6.

Newman, E. 1838. A descriptive list of the species of Rhysodes. The Magazine of Natural
History 2(2): 663-667.

Oke, C. 1932. Notes on Australian Coleoptera, with descriptions of new species. II. Proceedings
of the Linnean Society of New South Wales 57(3/4): 148-149.

Reitter, E. 1880. Einige neue Coleopteren. Verhandlungen des Naturforschenden Vereins in
Briinn 18: 29-30.

Reitter, E. 1889. Neue Coleopteren aus Europa, den angrenzenden Landern und Siberien mit
bemerkungen liber bekannte arten. Deutsche Entomologische Zeitschrift 31: p.23.

Sharp, D. 1899. Biologia Centrali- Americana, Insecta; Coleoptera, Colydiidae, Rhysodidae,
Cucujidae. 2(1): 497-560.

Viana, M.J. 1951. Una familia de Coleopteros neuva para la Republica Argentina:
Rhysodidae. Revista de la Sociedad Entomologica Argentina 15(1/3): 141-148.

Vulcano, M.A. and F.S. Pereira. 1975. Os Rhysodidae Sul-americanos (Col). Studia
Entomologia, 18(1/4): 153-188.

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

169

INDEX TO NAMES OF TAXA
(Synonyms in italics)

FAMILY GROUP TAXA
Dhysorini, 149

GENERA AND SUBGENERA
Arctoclinidium Bell, 60, 75, 77
Arrowina Bell and Bell, 156
Boreoglymmius Bell and Bell, 165
Clinidium Kirby, 3, 6, 1 1, 59, 68, 77
Clinidium s. str., 1 1, 48, 60, 74, 77, 93,
118

Grouvellina Bell and Bell, 151
Indoglymmius Bell and Bell, 165
Kaveinga Bell and Bell, 150
Kaveinga 5. str., 150
Kupea Philpott, 149

Laminoglymmius Bell and Bell, 164-165
Mexiclinidium Bell and Bell, 59-60, 75,
77, 137

Navitia Bell and Bell, 165
Omoglymmius s. str., 158, 162-163
Orthoglymmius Bell and Bell, 158
Protainoa Bell and Bell, 59, 69
Rhyzoarca new subgenus, 6, 8
Rhyzodiastes Fairmaire, 3, 6, 99, 119
Rhyzodiastes s. str., 6, 14, 48, 54
Rhyzostrix new subgenus, 6, 1 1, 48, 99,
119

Rhyzotetrops new subgenus, 6, 1 1
Tainoa Bell and Bell, 60, 70, 77
Temoana new subgenus, 6, 1 1-12, 48

SPECIES AND SUBSPECIES
abbreviata (Lea), Kaveinga, 150-151
actae Bell and Bell, Omoglymmius, 165
allegheniense allegheniense Bell and Bell,
Clinidium, 90

allegheniense georgicum Bell and Bell,
Clinidium, 90

alleni new species, Clinidium, 95, 1 18,
122-125

alveus new species, Rhyzodiastes, 14,
46-47

anguliceps (Arrow), Arrowina, 157
apertum allegheniense Bell and Bell,
Clinidium, 90

apertum apertum Reitter, Clinidium,
90-91

apertum Reitter, Clinidium, 78-79, 85,
89-90

argus new species, Clinidium, 93, 98, 146,
148

arrowi (Grouvelle), Yamatosa, 152
baldufi Bell, Clinidium, 78-79, 85, 89-90,
92

balli new species, Clinidium, 60, 62, 68-69
basilewskyi (Brinck), Dhysores, 149
beccarii Grouvelle, Clinidium, 93, 98, 146,
148

bechyneorum new species, Clinidium, 97,
136, 138

bifossulatum Grouvelle, Clinidium, 40
bifossulatus (Grouvelle), Rhyzodiastes, 13,
37-40

biimpressus new species, Dhysores, 149
bipunctatus new species, Rhyzodiastes, 12,
26, 28-29

blomi Bell, Clinidium, 60-61, 63, 66-67,
69

bonsae new species, Rhyzodiastes, 14, 26,
42-43

boroquense Bell, Clinidium, 94, 1 14-115,
117

boysi (Arrow), Yamatosa, 153
burnsi (Oke), Rhyzodiastes, 8-9
burnsi Oke, Rhyzodes , 9
calcaratum LeConte, Clinidium, 77-79,
84-85, 88

canaliculatum (Costa), Clinidium, 77-78,
83-84

canaliculatus Costa, Ips, 83
capito (Grouvelle), Omoglymmius,

162-163

cavicolle Chevrolat, Clinidium, 97,

140-143

centrale Grouvelle, Clinidium, 98, 140,

142, 144

Quaest. Ent., 1985, 21 (1)

170

Bell and Bell

championi new species, Clinidium, 60-62
chevrolati Reitter, Clinidium, 70-71, 74
chiolinoi Bell, Clinidium, 96, 125, 128
convergens new species, Rhyzodiastes, 13,
26, 29,31

corbis Bell, Clinidium, 96, 125-127
costatum (Chevrolat), Clinidium, 55-58
costatus (Chevrolat), Rhyzodes, 58
costatus (Chevrolat), Rhyzodiastes,
54-55, 57-58

costatus Chevrolat, Rhysodes, 58
crassicornis Bell and Bell, Omoglymmius,
148

crater new species, Clinidium, 97, 140,
143, 145

craticulus new species, Omoglymmius,
158-159, 162

curvatum new species, Clinidium, 97,
140-141, 144-145

curvicosta Chevrolat, Clinidium, 70-71,
74

darlingtoni Bell, Clinidium, 70-71, 74
davidsoni new species, Rhyzodiastes,
48-49, 52

denticauda new species, Rhyzodiastes, 13,
36-41

dohertyi (Grouvelle), Shyrodes, 1 14
dormans new species, Clinidium, 96, 123,
129-131

draco (Bell), Yamatosa, 153
dubium Grouvelle, Clinidium, 94, 115
edentata Bell and Bell, Grouvellina, 151
excavatum new species, Clinidium, 97,
136, 138-139

extrarium Bell and Bell, Clinidium, 70
fairmairei (Grouvelle), Clinidium, 45
fairmairei Grouvelle, Rhyzodiastes, 14, 44
feae (Grouvelle), Omoglymmius, 158
fibulata Bell and Bell, Kaveinga, 151
follis Bell and Bell, Omoglymmius, 159
fossatus new species, Rhyzodiastes, 14,
46-47

foveolatum Grouvelle, Clinidium, 97, 140,
142, 145

fraudulentus Bell, Omoglymmius, 38
frater (Grouvelle), Rhyzodiastes, 13,

37-40

frater Grouvelle, Clinidium, 38
fringillus Bell and Bell, Omoglymmius,
163-164

gestroi (Grouvelle), Rhyzodiastes, 13, 42
gestroi Grouvelle, Clinidium, 42
granatense Chevrolat, Clinidium, 74, 94,
99, 113

gressitti new species, Omoglymmius,
158-159

guatemalenum Sharp, Clinidium, 59-63,
66

guildingii Kirby, Clinidium, 93, 96,
133-136

guineensis (Grouvelle), Rhyzodiastes, 12,
25-27

guineensis Grouvelle, Clinidium, 26
haitiense Bell, Clinidium, 95, 125-127
halffteri new species, Clinidium, 59-62, 66
hammondi new species, Clinidium, 94, 99,
113

hesperus Bell and Bell, Omoglymmius,
157-158

hexadon new species, Grouvellina, 151
howdenorum new species, Clinidium, 95,
115-117

humboldti new species, Clinidium, 95,

118- 119, 123

humile new species, Clinidium, 97,

140-143

impressum new species, Clinidium, 93, 99
inaequalis Bell and Bell, Omoglymmius,
165-166

incis Bell, Clinidium, 1 14
incudis Bell, Clinidium, 94, 99, 117
indigens new species, Rhyzodiastes, 12-13,
26, 30-32, 42

inermis Bell and Bell, Omoglymmius, 165
insigne Grouvelle, Clinidium, 94, 115-117,
139

insularis (Grouvelle), Omoglymmius, 164
integrum Grouvelle, Clinidium, 48, 95,

119- 120

iviei new species, Clinidium, 60-61, 69
jamaicense Arrow, Clinidium, 96, 125,

Revisions of Rhyzodiastes Fairmaire and Clinidium Kirby

171

128-129

janus new species, Rhyzodiastes, 6-7
jolyi new species, Clinidium, 95, 1 19-121
kabakovi new species, Yamatosa, 152-153
kochalkai new species, Clinidium, 96, 99,
129, 132

kryzhanovskyi new species, Yamatosa,
152-153

largus new species, Omoglymmius, 158,

162-163

lineatus (Grouvelle), Omoglymmius, 165
liratum (Newman), Clinidium , 57
liratus (Newman), Rhyzodiastes, 54,
57-58

liratus Newman, Rhysodes, 56-57
longior (Grouvelle), Yamatosa, 152, 156
maderiense (Chevrolat), Clinidium , 53
maderiensis Chevrolat, Rhyzodes, 53
maderiensis Chevrolat, Rhyzodiastes ,
48-49, 52-53

marginicolle Reitter, Clinidium, 75,

77-79, 83

maritimus Bell and Bell, Rhyzodiastes, 12,
15,25

mathani Grouvelle, Clinidium, 97,

139-140

menieri new species, Rhyzodiastes, 49, 52
mexicanum Chevrolat, Clinidium, 60, 62,
67-69

microfossatum new species, Clinidium, 96,
133-134

mirabilis (Lea), Rhyzodiastes, 13, 26,
30-32

mirabilis Lea, Rhysodes , 30
mishmicum Arrow, Clinidium , 32
mishmicus (Arrow), Rhyzodiastes, 12,

32- 33, 37-38

moldenkei new species, Clinidium, 98,
146-147

montrouzieri (Chevrolat), Rhyzodiastes, 8
montrouzieri Chevrolat, Rhyzodes , 8
myopicum Arrow, Clinidium , 37
myopicus (Arrow), Rhyzodiastes, 13,

33- 34, 36-37

newtoni new species, Clinidium, 60-61, 63
nilgiriensis (Arrow), Arrowina, 157

niponensis (Lewis), Yamatosa, 152, 156
nitidus new species, Rhyzodiastes, 48, 52
oberthueri Grouvelle, Clinidium, 95,
119-122

occipitalis (Grouvelle), Kaveinga, 150
occipitalis Grouvelle, Rhysodes , 1 50
opacus new species, Omoglymmius,
157-158

oroensis Bell and Bell, Omoglymmius,

163- 164

pala new species, Clinidium, 97, 136, 139
parum-costatus Fairmaire, Rhyzodes,
54-56

parumcostatus (Fairmaire), Clinidium , 55
parumcostatus (Fairmaire), Rhyzodiastes,
6, 54-55

patruus new species, Rhyzodiastes, 13,
37-39

peckorum new species, Omoglymmius, 166
penicillatum new species, Clinidium, 96,
129, 131

peninsularis (Arrow), Yamatosa, 152
pentacyclus new species, Rhyzodiastes,
54-55

perplexus new species, Omoglymmius,

164- 165

pilosum Grouvelle, Clinidium, 95,

119-121

planiceps Bell and Bell, Omoglymmius,
159, 162

planifrons Fairmaire, Rhysodes, 1 5 1
planum (Chevrolat), Clinidium, 97,
133-136

planus Chevrolat, Rhyzodes, 135
poggii new species, Kaveinga, 150-151
pollinosus Bell and Bell, Rhyzodiastes,
11-12, 14-15

preobitalis new species, Rhyzodiastes, 12,
32, 34, 42, 54

probius Lewis, Rhysodes, 9
propinquus new species, Rhyzodiastes, 14,
43-44

propium (Broun), Clinidium, 9
proprius (Broun), Rhyzodiastes, 8-9
proprius Broun, Rhysodes, 9
punctatolineata (Grovelle), Arrowina, 156

Quaest. Ent., 1985, 21 (1)

172

Bell and Bell

punctatolineatus Grouvelle, Rhysodes,

156- 157

puncticornis Bell and Bell, Omoglymmius,
163-164

pygmaea Bell and Bell, Arrowina, 157
quadriimpressus (Grouvelle), Dhysores,
149

quadristriatum (Chevrolat), Clinidium,
48-49

quadristriatus (Chevrolat), Rhyzodiastes,
49

quadristriatus Chevrolat, Rhyzodes,
48-49

raffrayi (Grouvelle), Clinidium , 15
raffrayi Grouvelle, Rhyzodiastes, 12, 15
reitteri (Bell), Yamatosa, 152
rimatus Bell and Bell, Omoglymmius, 152
rimoganense (Miwa), Clinidium, 28
rimoganensis (Miwa), Rhyzodiastes, 13,
26, 28-30

rojasi Chevrolat, Clinidium, 97, 136-139
rosenbergi Bell, Clinidium, 78-79, 91-92
rossi Bell, Clinidium, 96, 118, 129
rostrata (Lewis), Arrowina, 156
sculptile (Newman), Clinidium, 75,
78-79,85,91-92

sculptilis Newman, Rhysodes, 91-92
segne new species, Clinidium, 96, 129,
131-132

simplex Chevrolat, Clinidium, 137
singulare Heller, Clinidium, 27
singularis (Heller), Rhyzodiastes, 12, 25,
27

smithsonianum new species, Clinidium,

97, 133-134, 136

spatulatum new species, Clinidium, 98,
142, 145

spissicorne (Fairmaire), Clinidium, 45
spissicornis Fairmaire, Rhyzodiastes, 14,
44-46

strabus (Newman), Omoglymmius,

157- 158

strigiceps Bell and Bell, Kaveinga, 150
stylatus Bell and Bell, Omoglymmius,
165-166

sulcicollis (Grouvelle), Rhyzodiastes,

11-12, 15,24-25,48
sulcicollis Grouvelle, Clinidium, 24
sulcigaster Bell, Clinidium, 98, 146-147
sulcipennis Mulsant, Rhysodes, 83
suturalis new species, Rhyzodiastes, 54, 59
taprobanae (Fairmaire), Arrowina,
156-157

tolai new species, Omoglymmius, 163-164
trionyx new species, Clinidium, 95,
124-125

triplehorni new species, Clinidium, 60, 62,
68

trisulcatum (Germar), Clinidium, 83
trisulcatus Germar, Rhysodes, 83
tubericeps (Fairmaire), Grouvellina, 151
turquinense Bell, Clinidium, 74
vadiceps new species, Rhyzodiastes, 13,

34, 36-38

valentinei Bell, Clinidium, 77-79, 85, 90
validum Grouvelle, Clinidium, 98, 140
143-146

veneficum Lewis, Clinidium, 52, 75,
77-79, 84, 88

viduus Bell and Bell, Omoglymmius, 164
waterhousei (Grouvelle), Rhyzodiastes,

12, 32-34

waterhousei Grouvelle, Clinidium, 33
whiteheadi new species, Clinidium, 95,

118, 122, 124

xenopodium Bell, Clinidium, 70-71, 75

BOOK REVIEW

Belton, Peter. 1983. Review of mosquitoes of British Columbia. British Columbia
Provincial Museum Handbook 41. 189 pp. $5.00 Canadian.

This excellent little book describes, in simple terms, the mosquito fauna of B.C.
Important information such as the biology and morphology of mosquitoes is dealt
with adequately, both descriptively and diagramatically. Simple and workable
couplet keys for species identification of adult females and fourth instar larvae, are
greatly aided by line drawings, habitat and species descriptions. With a little effort,
the most amateur of amateurs could identify all of British Columbia’s mosquitoes.
The book will be an asset to any mosquito control program, where identification by
inexperienced personnel is required.

Introductory sections add interest to the book. The description of the Life Zones of
B.C. shows the great diversity of British Columbia’s terrain - from rain forest, tidal
pools and salt marshes in the west to arid regions in the east, and from arctic and
subalpine zones in the north to more temperate ones in the south. With this great
climatic and geographic diversity goes a concomitant diversity in the mosquito fauna.
Sections about mosquitoes in the culture of The Northwest Coast Indians, the
collection and preservation of mosquitoes, and the history of mosquito study and
management in B.C. are included, as well as a useful glossary and reference section.

Because the species composition of B.C. is very similar to that of Alberta, the key
could be used in Alberta. A total of 46 species in 5 genera occur in B.C. - 33 Aedes , 3
Anopheles , 3 Culex, 6 Culiseta and 1 Mansonia. In Alberta, 42 species occur in the
same 5 genera. Five species occur in Alberta but are absent from the species record of
B.C., namely, Aedes churchillensis, Ae. decticus, Ae. nigromaculis, Ae. schizopinax
and Culex restuans. Of these five Dr. Belton has included Ae. nigromaculis and Ae.
schizopinax in the couplet key. The other three species are rare in Alberta and should
not prove a problem. The existance of regional morphological variations should be
kept in mind, however, when using the key.

I have only two criticisms, firstly the book binding will not survive the great deal
of use that such a key will receive; a ring binding would have been more serviceable.
And secondly, there is no key to male identification. Although the author does point
out that males can be keyed using Carpenter and La Casse, this key is now somewhat
out of date. Male identification is somewhat easier, but requires slide preparation in
most cases and the use of a compound microscope.

The publication of this book by the British Columbia Provincial Museum is the
fortyfirst in what is already an excellent series. I anticipate that Peter Belton has
started something that will result in others writing similar pocket books on their local
mosquito fauna.

P.J. Scholefield

Alberta Environment, Pollution Control Division
Pesticide Chemicals Branch
2938- 11 St. N.E.
Calgary, Alberta, T2E 7L7

Quaest. Ent., 1985,21 (1)

174

BOOK NOTICE

Griffiths, G.C.D. (Editor). Flies of the Nearctic Region. E. Schweizerbart’sche
Verlagsbuchhandlung (Nagele u. Obermiller) Stuttgart. Volume VIII. Cyclorrhapha
II (Schizophora: Calyptratae) Part 2, Number 2. Anthomyiidae, by G.C. Griffiths,
pp. 161-288 (1983). $56.76 US. Part 2, Number 3. Anthomyiidae, same author, pp.
289-408 (1984). $56.76 US.

G.E. Ball reviewed earlier issues of this series in Quaestiones Entomologicae
(1980, 16(3/4): 676-678; 1983, 19(3/4): 489-490), including Part 1 of Griffiths’
treatment of the nearctic species of Anthomyiidae. That part contained a brief
introduction to the family, a thorough taxonomic treatment of the genus Pegomya
Robineau-Desvoidy subgenus Pegomya and ended mid sentence in his description of
P. magdalensis new species. Part 2 concludes his consideration of the subgenus
Pegomya , begins that of the subgenus Phorea and ends mid sentence in a description
of P. valmariensis new species.

In Part 3, Griffiths finishes with Pegomya and provides separate keys for the
identification of males (pp. 337-346) and females (pp. 346-351), thus rectifying one
of the weaknesses of Part 1 noted by Ball ( Quaest . Ent. 19: 490). In addition, this
fasicle contains full taxonomic consideration of the species of Emmesomyia Malloch
and Parapegomyia new genus, including keys to species and a “first reference list” of
papers cited in Parts 1 - 3 and expected to be cited in forthcoming parts. Thus,
Griffiths’ finished treatment of the family will be unusual in having its list of
references in the middle rather than at the end. All Ball’s critical comments about
Part 1 apply equally to Parts 2 and 3.

B.S. Heming
Department of Entomology
University of Alberta

Q.Va<V
<o ?LO0

Quaest

lones

Entomologicae

M(jc>

i ?
in &^y°°L
JUftl0K

A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.

VOLUME 21

NUMBER 2

1985

Abstracts are required one in English, and one in another language,
preferably in French.

Communications regarding subscriptions and exchanges should be
addressed to the Subscription Manager, and regarding manuscripts to the
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QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.

Volume 21 Number 2 1985

CONTENTS

Fredeen-Some Economic Effects of Outbreaks of Black Flies ( Simulium luggeri

Nicholson and Mickel) in Ssakatchewan 175

Ward-The Nearctic species of the genus Pseudomyrmex (Hymenoptera:
Formicidae) 209

SOME ECONOMIC EFFECTS OF OUTBREAKS OF BLACK FLIES ( SIMULIUM
LUGGERI NICHOLSON AND MICKEL) IN SASKATCHEWAN1

F.J.H. Fredeen

Agriculture Canada Research Station
107 Science Crescent
SASKATOON, Saskatchewan, Canada
S7N0X2

Quaestiones Entomologicae
21:175-208 1985

ABSTRACT

Larvae of Simulium luggeri Nicholson and Michel were first detected in the South
Saskatchewan River in 1968, coincidental with appearances of aquatic weeds. They became
abundant in both the South and North branches in Saskatchewan by 1971. Damaging
outbreaks occurred for the following reasons: larvae readily drifted downriver and colonized
extensive beds of weeds, S. luggeri is multivoltine, adults dispersed widely and attacked most
mammals including people, swarming about heads causing stress and hyperactivity. During
outbreaks, grazing and breeding activities of livestock were interrupted and lactation reduced.
Increased incidence of stress-related injuries and diseases including broken limbs, keratitis,
mastitis, calfhood scours and pneumonia occurred.

Chronic outbreaks of S. luggeri have occurred every summer since 1975 in east-central
Saskatchewan. Use of methoxychlor larvicide reduced potential severity of many outbreaks.
The most destructive outbreaks occurred in 1978 when black flies spread onto about 38,000
km 2 of east-central Saskatchewan and caused measurable economic losses in about 5,700 km2.
Losses to beef producers in 1978 were estimated to have exceeded $2.9 million and included
unrealized weight gains, delayed conceptions, fatalities, replacement of debilitated bulls and
increased costs for labour, veterinarians’ services, fence repairs and supplementary feeding.
Losses to dairy producers were estimated to have exceeded $57,000. Milk production from
severely affected cows did not return to normal until after new lactation cycles commenced,
sometimes several months after outbreaks ceased in the fall.

Producers responded in several ways, for example by changing management practices, by
reducing or eliminating herds, by converting pasture lands to less productive uses, and by
submitting petitions to governments for improved control of larvae.

RESUME

En 1968, la mouche noire, Simulium luggeri Nicholson & Mickel, commenqait a se reproduire dans la riviere
Saskatchewan. L’apparition de S. luggeri a coincide avec I’envahissement de la riviere par la f lore aquatique. Selon
■I’auteur, des attaques severes de mouche noire se sont produites pour les raisons suivantes: les larves sont aisement
entrainees par le courant et s’attachent aux vastes bancs de plantes aquatiques; la mouche noire est plurivoltine; les
insectes adultes sont capables de coloniser de vastes superficies et ils s’attaquent h la plupart des mammi feres, y compris
1‘homme; les insectes ont tendance a s’attaquer d la fete des animaux, causant chez ceux-ci des signes de stress et
d’hyperactivite. Au cours des attaques de mouche noire, on a observe que les animaux cessaient de brouter et de se
reproduire et que les vaches en lactation produisaient moins de lait. On a aussi note un nombre accru de blessures et de
maladies causees par le stress, telles que membres rompus, keratite, mastite, diarrhee du veau et pneumonie.

‘Contribution No. 866 of the Research Station, Saskatoon

176

Fredeen

Les attaques de mouche noire se sont produites de faqon chronique chaque ete depuis 1975 dans la region centre-est
de la Saskatchewan. L'emploi du larvicide methoxychlor a contribue d reduire la severite de plusieurs attaques. Les
attaques les plus severes se sont produites en 1978, les mouches noires infestant pres de 38 000 km 2 et causant des
dommages mesurables sur pres de 5 700 km2. En 1978, les pertes subies par les producteurs de boeuf de boucherie ont ete
estimees d plus de 2.9 millions de dollars. Ces pertes ont resulte des fails suivants: gains de poids plus lents, conceptions
retardees, pertes d’animaux, ainsi que couts de remplacement des taureaux malades, de la main d’oeuvre
supplemental , des soins veterinaires, des reparations aux clotures et des supplements alimentaires. Par ailleurs, les
producteurs laitiers ont subi des pertes estimees d plus de 57 000 $. II fut observe que le rendement des vaches laitieres
serieusement atteintes ne redevenait normal qu’apres le debut d'un nouveau cycle de lactation, qui ne survenait parfois
que plusieurs mois apr'es la disparition des mouches noires, en automne. Les producteurs ont combattu la mouche noire
de differentes faqons: en modifiant leurs methodes de gestion, en reduisant ou en eliminant completement les troupeaux,
en utilisant les paturages d des fins moins productives, ou en petitionnant pour l’ amelioration du controle des larves.

INTRODUCTION

Every summer since widespread outbreaks of the black fly Simulium luggeri Nicholson and
Mickel commenced in 1976 in east-central Saskatchewan, residents have demanded
government assistance with abatement. Methoxychlor larvicide is effective (Fredeen, 1974,
1975) and its use in the Saskatchewan River apparently is not permanently harmful to
non-simuliid fauna (Fredeen, 1983). However, use of larvicide cannot be condoned without
unequivocal proof of need. The purpose of this paper is to compile and assess evidence of losses
to beef and dairy cattle producers in Saskatchewan resulting from outbreaks of S. luggeri.

Populations of black fly larvae in both branches of the Saskatchewan River in Saskatchewan
were dominated by S', arcticum until the mid- 1970’s. From this river sporadic outbreaks of S.
arcticum spread widely into surrounding farmlands killing numerous animals (Rempel and
Arnason, 1947; Fredeen, 1958). Major outbreaks of S. arcticum ceased in 1948 with the advent
of chemical larviciding (Fredeen, 1953, 1977(a)) but minor outbreaks continued because of
downstream drift of eggs and larvae from untreated sections.

In early years, the Saskatchewan river usually was deep and turbid throughout much of the
ice-free season with beds of sand and rocks completely free of vegetation. In 1968, completion
of a hydroelectric dam on the South Saskatchewan River, 1 1 5 km south of Saskatoon (about
350 km above its confluence with the North Saskatchewan River) (Fig. 1) created a reservoir
250 km long, a complete barrier to further migration of larvae down that river (Fredeen,
1977(b)). By 1971 the South Saskatchewan River below the reservoir had become relatively
shallow during ice-free months due to storage of water for wintertime generation of hydropower
(Fig. 2, S.S.R.). The reservoir served as a sink for suspended solids, and the combination of
shallow, clear water in the river below the dam allowed sufficient insolation to encourage, for
the first time, growth of massive beds of algae1 and broad-leaved plants2 on the river bed.
Growths of aquatic plants undoubtedly also were enhanced by nutrients released from urban
and rural communities. This is evident today when comparing growths above and below large
cities on the Saskatchewan River.

In 1975 similar trends became evident in the North Saskatchewan River (Fig. 2, N.S.R.).
Relatively shallow, clear water replaced the large, turbid summertime volumes of previous
years. This was due in part to drought conditions which greatly reduced runoff in a major

'Mainly Cladophora glomerata (L.) Kutzing

The four most common species are Ceratophyllum demersum L.,
Myriophyllum exalbescens Fernald, Potamogeton crispus L., P. pectinatus
L.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

177

Fig. 1 . Map of Alberta, Saskatchewan and Manitoba indicating sites where immature stages of Simulium luggeri were
collected from the Saskatchewan River in southern Saskatchewan and Alberta, and from other river systems. The
boundaries of Crop Districts 8 and 9 outline regions in Saskatchewan where most outbreaks occurred in recent years.

portion of that watershed, and in part to completion of two hydroelectric reservoirs in the
foothills, Brazeau in 1962 and Abraham Lake in 1972. Together they controlled about half of
the volume of water reaching the lower end of the North Saskatchewan River. Extensive
summertime beds of algae and aquatic broad-leaved plants appeared, especially between
Edmonton, Alberta and North Battleford, Saskatchewan. Paterson and Nursall (1975)
suggested that the presence of algae below Edmonton was the result of increased nutrient
content (chiefly nitrates) in that portion of the river. Weed beds above North Battleford are
occupied mainly with larvae of S. vittatum Zetterstedt, a species relatively tolerant of organic
pollution.

These environmental changes in both branches of the Saskatchewan River discouraged
accumulation and development of larvae of S. arcticum which prefer to attach to clean boulders
in fast-flowing water, but encouraged invasions of S. luggeri , S. vittatum , S. meridionale Riley,

Quaest. Ent., 1985,21 (2)

VOLUME (M3/Sec) VOLUME (M3/Sec)

178

Fredeen

MONTH

Fig. 2. Average monthly volume flows in the Saskatchewan River: S.S.R. = South Saskatchewan River at Saskatoon,
1911 through 1970, and 1971 through 1981;N.S.R. = North Saskatchewan River 1911 through 1974, and 1975 through
1981 (Environment Canada 1980 (a), 1981, 1982).

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

179

TABLE 1. MAXIMUM DENSITIES OF LARVAE AND PUPAE OF FOUR SPECIES OF
BLACK FLIES OBSERVED ON NATURAL AND ARTIFICIAL SUBSTRATES IN
THE NORTH AND SOUTH SASKATCHEWAN RIVERS IN SASKATCHEWAN

(NUMBER/CM2)

S. arcticum

North Saskatchewan River
near Prince Albert, Sask.

-S!

| £
bo a

-3 5

S', arcticum

South Saskatchewan River
near Birch Hills, Sask.

§ 5

jts ^ 5

bo -j- a

-3 £ >

Co oj

1947 to
1968

100 +

<1*

1969

1970

1971

1972

1973

1974

1975

1976

1977

128

1978

1979

316

1980

370

1981

469

*In the South Saskatchewan River larvae of S. luggeri were first detected in 1968.

Quaest. Ent.. 1985,21 (2)

180

Fredeen

and other species which prefer to attach to leaves of aquatic plants (Fredeen, 1981).

RECENT TRENDS IN SPECIES OF BLACK FLIES INHABITING THE
SASKATCHEWAN RIVER IN SASKATCHEWAN

Methods

Annual trends in maximum densities of larvae of the four main species of black flies
inhabiting the north and south branches of the Saskatchewan River in Saskatchewan are shown
in Table 1. Until 1969, populations of larvae were estimated by counting numbers attached to
rocks collected from rapids. In 1969 accuracy of estimates presumably was improved when we
began to anchor artificial substrates for larvae to attach to (metre-length pieces of rope)
(Fredeen and Spurr, 1978). Between 1969 and 1976 larvae were counted at weekly intervals for
only a few weeks in late spring each year or until chances of outbreaks of S. arcticum were
considered to have ended for the year ( S . arcticum generally peaked in May or June). But
beginning in 1977, weekly samples were collected throughout each summer from both branches
of the Saskatchewan River because the newly-established S. luggeri was multivoltine and
larvae were abundant and had to be monitored throughout much of the ice-free season.

South Saskatchewan River

Populations of larvae in the South Saskatchewan River were dominated by S. arcticum each
spring until about 1977 (Table 1). The final major outbreak of S. arcticum believed to have
originated at least in part from the South Saskatchewan River, occurred June 13 to 18, 1967.
In that outbreak 43 animals were known to have been killed in communities extending
southeastwards more than 100 km from Prince Albert.

Larvae of S. luggeri (indicating a breeding population) were first collected from the South
Saskatchewan River in July and August, 1968. But it was not until 1977 that we commenced
regular weekly collections from that river and these showed that S. luggeri had replaced S.
arcticum as the dominant species. Previously, I had found it breeding only in small, weedy
rivers across Manitoba, Saskatchewan and Alberta (Fig. 1). Shewell (1958) reported it from
the Churchill River on Hudson Bay and from the Mackenzie River and tributaries as far north
as Norman Wells, N.W.T. The earliest observed outbreak of S. luggeri , believed to have
originated from the South Saskatchewan River, occurred on August 22, 1972 when this species
was identified in swarms causing cattle to run. Since then outbreaks of varying severity have
originated from the final 150 km of this river every year.

North Saskatchewan River

The final recorded outbreak of S. arcticum of economic proportions from the North
Saskatchewan River occurred in mid-June 1972. During that outbreak at least 19 cattle were
killed near a section of that river upstream from Prince Albert.

In 1971 and 1972, during tests of artificial substrates, there were significant increases in
numbers of larvae of S. luggeri in samples collected in late summer from several sites in the
North Saskatchewan River indicating establishment of a breeding population in that river.
Previously, larvae had been collected from the North Saskatchewan River only from restricted
sites below the mouths of small tributaries.

In May through October 1975, the volume of the North Saskatchewan was greatly reduced
(to about 50 percent of long-term means) and coincidentally by early June, larvae of S. luggeri
became unusually abundant. In May, 1976 its larvae outnumbered those of S. arcticum in the

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

181

North Saskatchewan for the first time (Table 1) and within a month the first widespread
outbreaks of S. luggeri began from that river. Since then, larvae of S. luggeri have remained
relatively abundant, especially in the final 300 km of the North Saskatchewan River before its
confluence with the south branch, and in the entire 130 km of the main Saskatchewan River
between the confluence and Tobin Lake at Nipawin (Fig. 1). Although numbers of larvae of S.
vittatum began to surpass those of S. luggeri by 1979 (Table 1), significant outbreaks of that
species have not been reported yet, perhaps because livestock appear to be relatively tolerant of
it. Larvae of S. arcticum still occur regularly although in small numbers every year in both
branches of the Saskatchewan River, indicating potential for staging a comeback should
conditions change in its favor.

CHRONOLOGY OF OUTBREAKS OF S. LUGGERI FROM THE SASKATCHEWAN

RIVER

1976

In 1976, larvae of S. luggeri accumulated on artificial substrates (rope pieces) anchored in
the North Saskatchewan River in rapids 22 km below Prince Albert, Saskatchewan at an
average rate of 1550 larvae per metre of rope per week during May and June. A maximum
density of 32 larvae per cm2 of substrate surface (Table 1) was observed on June 22.
Populations were not measured in the South Saskatchewan River that year. Although the
maximum density observed in the North Saskatchewan was three times larger than that
observed in 1975 there were no concerns about possibilities of damaging outbreaks because S.
luggeri had not been known to lethally poison animals as did its predecessor, S. arcticum. Also
there had been no complaints of black fly attacks on livestock in 1975, although a year or more
later some producers did recall that their cattle had been noticeably bothered by black flies in
1975.

Environmental changes in 1976 that may have accounted, at least in part, for observed
increases in numbers of larvae included greatly reduced river volumes and higher water
temperatures. Ice on the North Saskatchewan River broke up a week earlier than normal that
spring, and water volumes in both branches remained much below normal throughout May,
June and July. Also mean daily air temperatures in May were 17 percent above normal and
hours of sunshine 32 percent above normal. For all of these reasons, river water in both
branches of the Saskatchewan River warmed up more rapidly than normal that spring,
attaining daily maxima of 20°C 3 to 4 weeks earlier than in previous years. Also, because
relatively low water levels and low water turbidities had allowed greatly increased insolation,
growths of aquatic weeds (favored attachment sites for larvae of S. luggeri) and planktonic
algae (which served as food) were increased.

During ice-free months in earlier years, water turbidities as high as 7270 mg/L were
reported for the South Saskatchewan River and 3050 mg/L for the North Saskatchewan
(Environment Canada, 1980 (b)). Microscopic examination of turbid water showed
phytoplankters to be very scarce but numbers were not recorded.

In 1976, weekly measurements of turbidity indicated summertime maxima of only 52 mg/L
for the south branch at Birch Hills and 99 mg/L for the north branch near Prince Albert.
Phytoplankters were so abundant that the water was greenish, but numbers were not counted.
In 1978, under similar turbidity conditions (maxima of only 97 mg/L for the south branch and
211 for the north), maxima of 550 and 1 100 phytoplankters per 0.001 mL of river water were
recorded for these two branches.

Quaest. Ent., 1985, 21 (2)

182

Fredeen

Weather conditions, especially air temperatures, also generally favored black fly attacks on
animals. Attacks by S. luggeri are most vigorous between about 20 and 25° C, and daily
maximum temperatures above 20° C were recorded on 21 days in May, 19 in June, 28 in July
and 28 in August.

In 1976, pupae of S. luggeri were first collected on May 25. The first report of outbreaks in

1976 came from Mr. Glen Love, who had a mixed farm between the north and south branches
of the Saskatchewan River about 30 km east of Prince Albert. He first observed black flies
around his cattle on the afternoon of May 29. From then on until June 8 his livestock were so
severely attacked when outside during daylight hours that they had to be fed and watered
indoors. Mr. Love marketed most of his cattle on June 8 to avoid further costs of supplementary
food and labour. Black fly swarms at that time consisted of 98 percent S. luggeri with about 2
percent S. arcticum, S. meridionale and S. vittatum.

During that first week of June, livestock throughout a 7,000 km2 area, extending some 30
km on either side of the entire Saskatchewan River between Prince Albert and Nipawin (Fig.
3), were severely attacked by S. luggeri every day. Producers claimed that their cattle were
continually surrounded by clouds of black flies during daylight hours and that attacks were
especially severe before thunderstorms on June 3 and 4. Smudges were kept burning
throughout the region, even in the largest community pastures. There were numerous
complaints from people who claimed they also were bitten. Even with repellents, black flies
swarmed around them so thickly that they could not avoid breathing them in.

Cooler weather (daily maxima of 13 to 20°C) on 11 days between June 8 and 26 was
believed responsible for several lulls in outbreaks. There was also a decline in numbers of larvae
in the river until after second generation larvae appeared in June. However, between June 26
and July 15 swarms of S. luggeri again severely affected livestock and people in widespread
areas, including an additional 10,000 km2 of rural and urban lands to the south, and 6,000 km2
of recreational lands to the north, extending almost as far north as La Ronge (Fig. 3). This
second series of outbreaks declined noticeably by mid-July, apparently due in part to a single
injection of methoxychlor larvicide into the North Saskatchewan River on July 7.

A third series of widespread outbreaks in 1976 occurred throughout most of August and
September and lasted until warm weather ended in the fall.

In summary, residents in some 23,000 km2 of agricultural and recreational areas in
Saskatchewan complained in 1976 of black fly outbreaks, by telephone calls and letters to
federal and provincial offices in at least ten centres. Complaints of damage to livestock came
from producers from diverse regions totalling about 2,500 km2. There were three major periods
of attack, probably related in part to production of at least three generations of S. luggeri.
Ninety-five to 99 percent of the black flies in attacking swarms were S', luggeri. The remainder
were S. arcticum , S. meridionale and S. vittatum as indicated by sweep net collections.

1977

The larval monitoring program was greatly enlarged in 1977 with artificial substrates (rope
pieces) anchored and exchanged weekly in six sites, May through August. Comparisons of rates
of accumulation of larvae with rates in 1976 were possible only for May and June, and only for
one site located in rapids 22 km below Prince Albert on the North Saskatchewan River. There,
larvae accumulated at an average rate of 7,000 larvae per metre of rope per week through May
and June and 7,180 for May through August. This indicated an unusually large, persistent drift
of larvae downstream into the larvicide-treated section of the river throughout the summer. The

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

183

Quaest. Ent., 1985,21 (2)

Fig. 3. Maps of central Saskatchewan showing approximate areas subjected to outbreaks of Simulium luggeri in 1976 and 1977, and specific localities from which

184

Fredeen

May-June rate was about 4.5 times larger than for the same period in 1976. A maximum
density of 128 larvae/cm2 of substrate surface occurred on May 10 (Table 1).

Each of the north and south branches of the Saskatchewan River was injected at one site
with methoxychlor larvicide on each of three dates in 1977 (Fredeen, 1983). Treated sites were
rapidly repopulated that summer. For example, following injections on July 4, numbers of
larvae on artificial substrates 42 km downstream in the south branch initially declined by 78
percent but returned to pre-treatment levels within seven days. In the north branch 61 km
downstream from the injection point in that river, numbers of larvae initially declined by 85
percent but a week later were 2.4 times larger than before treatment! In the main
Saskatchewan River, an additional 71 km downstream from those monitored sites, numbers of
larvae on artificial substrates declined by 75 to 95 percent following each injection, but within
three weeks exceeded pre-treatment numbers.

Attacks on livestock commenced during the last week of May, but practically ceased by
June 3. They recommenced June 29 and continued sporadically throughout the remainder of
the summer whenever the weather was suitably warm. The total area where black flies were
seen included some 6,000 km2, generally within 10 to 15 km of either branch of the
Saskatchewan River (Fig. 3). Severe outbreaks were reported from about 400 km2 of
farmlands.

1978

The worst outbreaks of S. luggeri on record occurred in 1978. In May through August, a
mean density of 3180 larvae attaching weekly to metre-length rope-piece substrates anchored
in the North Saskatchewan River indicated a high rate of drift of larvae downstream into
treated sections of that river. Larvicide was injected three times into the North Saskatchewan
River, all at Prince Albert, 25 km above the monitoring site. (This was unlike the previous year
when only one of the injections was made above the monitoring site.) Four days after the first
injection on May 26, numbers of larvae attached to artificial substrates had declined by 77
percent. Seven days later numbers were still 48 percent lower than before the injection.
However, an unusually high rate of downriver drift of larvae following the second injection on
June 20 caused a 2.9 fold increase in density, rather than the expected decrease within one
week. Two weeks after that injection, density had increased even more, to 3.8 fold that seen
before the injection. One week after the third and final injection on August 8, numbers of
larvae had declined by 62 percent and two weeks afterwards, were still 48 percent lower than
before the injection.

In the South Saskatchewan River mean weekly numbers, May through August, increased
from 1,400 in 1977 to 3,340 in 1978. This occurred despite four larvicide injections 32 km
above the monitoring site. The first injection, May 26, was considered successful and caused an
84 percent reduction in numbers of larvae during the first week and a further decline to 95
percent by the end of the second week. The second injection, June 20, was not successful
because within one week, numbers of larvae had increased by 1 .4 fold pre-treatment values and
by the end of the second week, by 3.1 fold. One week after the third injection, on July 21,
numbers of larvae had declined by 70 percent but, by the end of the second week, numbers had
increased by a factor of 5.3 over those seen before the injection. Reductions of 69 and 76
percent were observed after the fourth injection on August 8, perhaps not so much because of
that injection, but because of the normal seasonal decline at that time of the year.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

185

In the main Saskatchewan River 70 km downriver from the confluence of the two branches,
mean densities of larvae attaching weekly to artificial substrates May through August
increased from 275 per week in 1977 to 1,600 per week in 1978. Drift of larvae out of the
branches following larvicide injections was responsible for much of this increase. Although
numbers declined by 51 percent following the first injections into both branches on May 26,
they increased six-fold following the second pair of injections on June 20 and 65-fold after the
third injections (south branch only) on July 21! After the fourth and final pair of injections on
August 8, numbers declined by 99 percent but this decline may have been due partly to a
normal seasonal trend not related to larviciding. Results in general indicated, as in previous
years, that injections of larvicide into the two branches of the river could not guarantee
reductions in the main river below the confluence, but that dangerous increases might occur
instead, especially during June and July.

Casual observations at sites additional to the three regular monitoring sites in 1978
indicated that each larvicide injection did reduce numbers of larvae in at least a portion of the
river. However, in general, these treatments failed to prevent massive outbreaks for at least
three reasons: 1) The first injections were made at least 1 week too late, delayed until May 26
because of a lengthy hatching period of overwintered eggs that year. This was a mistake,
because, by that date, adults of S. luggeri had already commenced emerging. These eventually
laid sufficient eggs to allow production of massive numbers of second-generation larvae. 2) The
weedy South Saskatchewan River should have been injected at several sites on each date
instead of at only one. 3) The main Saskatchewan River downstream from the confluence of the
two branches also should have been injected in June, July and August. By May 26 adults of S.
luggeri already were causing cattle to stampede in pastures eastward from Prince Albert,
mainly along the South Saskatchewan River. By May 31 many herds in more than 5,000 km2 of
farmlands between Prince Albert and Nipawin (Fig. 4) were reported to be under very severe
attack, and by June 3 black flies had moved north, against prevailing winds, through some 20
km of dense woodlands in the uninhabited Nisbet Forest from the main Saskatchewan River, to
commence what was to be about 16 weeks of continuous harassment of people and livestock in
the Smeaton, Choiceland, and White Fox areas. Six days of northerly winds in late May and
early June also carried dense swarms southward, and by June 3 reports of severe black fly
problems were coming from as far south as Wynyard, some 170 km south of the nearest point
of emergence on the Saskatchewan River. Strong easterly winds on June 5, 6, 9, 10, 12 and 13
then carried many of these black flies even further, to more than 40 km southwest of Saskatoon
(Fig. 4). These black flies apparently had travelled more than 300 km from their origin.

In summary, livestock and people in an area probably exceeding 38,000 km2 were affected
by black flies in 1978. Almost one-seventh of this area, about 5,700 km2, was subjected
repeatedly to very severe attacks until after a second set of larvicide treatments on June 20, and
sporadically thereafter until late September. Outbreaks in 1978 were more severe and
widespread than in any year since outbreaks commenced in 1972.

1979

Minor outbreaks of relatively short duration occurred throughout an 18,000 km2 area in
central Saskatchewan in 1979 (Fig. 4). Black flies were reported mainly from areas adjacent to,
and between, the two branches of the Saskatchewan River. Severe outbreaks were reported
from areas totalling only about 750 km2, immediately adjacent to the Saskatchewan River
between Prince Albert and Nipawin.

Quaest. Ent., 1985, 21 (2)

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Fredeen

Fig. 4. Maps of outbreaks in 1978 and 1979.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

187

Numbers of larvae accumulating weekly May through August on metre-length rope pieces
anchored in the North Saskatchewan River averaged 3,340, numbers similar to those seen in
1978. However, few larvae were allowed to pupate because of five timely injections (one in two
locations) of methoxychlor larvicide. Mean densities of larvae in the South Saskatchewan River
were only about one-tenth those seen in 1978 due not only to three timely injections of larvicide
but also to the use of several injection sites in this weedy river.

For the first time, the main Saskatchewan River below the confluence was injected. Four
consecutive injections (one in two sites) prevented most larvae which may have drifted
downriver from the two branches from maturing.

Livestock producers were particularly satisfied with results of the larval abatement program
in 1979 and petitioned federal and provincial governments that the program be continued in
future years.

1980

In 1980 outbreaks again were relatively minor and although black flies were reported at one
time or another from about 20,000 km2, severe outbreaks occurred in only about 400 km2,
mainly near the rivers as in 1979 (Fig. 5).

Densities of larvae in both branches of the river were unusually high in May but timely
treatments with larvicide prevented many from maturing. Mean numbers from May through
August were similar to those observed in 1979: 3,310 larvae per metre of rope in the north
branch and 540 in the south. Larvicide was injected only once, on May 16 into the north branch
and twice, at two locations each time into the south branch. The main river below the
confluence was not injected because populations of larvae remained very low there all summer.

1981

Outbreaks were less troublesome in 1981 than in 1980 with black flies being reported from
only about 7,400 km2 (Fig. 5).

Mean weekly numbers of larvae of S. luggeri declined from those seen in 1980 to 1,570 per
metre of rope in the north branch, but increased slightly to 670 in the south branch. Larvicide
was injected four times into the north branch, three times into the south branch and once below
the confluence of the branches.

FACTORS CONTRIBUTING TO DAMAGING OUTBREAKS OF S. LUGGERI

My first experience with an outbreak of S. luggeri occurred on August 6 and 7, 1951 when I
observed horses stampeding under attack from black flies that were emerging from the
relatively small Battle River about 20 km west of North Battleford, Saskatchewan. Numbers of
black flies (all S. luggeri) were relatively small and effects were negligible. This species had
been named only the previous year by Nicholson and Mickel (1950) from specimens collected
in Minnesota and Wisconsin. The authors believed at that time that S. luggeri was occasionally
bothersome to horses but not to people.

As long as S. luggeri was restricted to breeding in small prairie rivers, potential for creating
damaging outbreaks was limited. But when it commenced breeding in the Saskatchewan River,
potential with regard to intensity, duration and areas affected increased manyfold. At first
there seemed no reason for concern because larvae became abundant only in late summer, and
numbers waned quickly with advent of cooler temperatures in August and September.

Quaest. Ent., 1985, 21 (2)

1980 M > 1981

188

Fredeen

Approximate margin of outbrea
Region of severe outbreaks

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

189

Furthermore, during occasional brief outbreaks no animals were killed as had happened
frequently with S. arcticum.

It was not until 1976 that the first summer-long outbreaks of S. luggeri from the
Saskatchewan River occurred. Those outbreaks became so severe by mid-summer that
larviciding was initiated using strategies initially developed to prevent outbreaks of S. arcticum
(Fredeen, 1974, 1975). But whereas an outbreak of S. arcticum usually could be prevented
with a single injection of methoxychlor larvicide into the Saskatchewan River, such a treatment
proved relatively ineffective against S. luggeri. Research in subsequent years indicated several
reasons why outbreaks of S. luggeri of economic proportions developed, and why major failures
occurred in a larviciding program that had been used successfully against S. arcticum.

Major outbreaks occurred because:

1. Immense numbers of larvae of S. luggeri were capable of inhabiting weed beds in the
Saskatchewan River, and since the species was multivoltine,numbers tended to increase during
succeeding generations May through August each summer, sometimes in spite of larvicide
treatments. Enormous numbers of larvae frequently drifted downriver from untreated sections
for natural reasons still under investigation.

2. Adults were capable of dispersing great distances on the wind. The Saskatchewan River
had the potential for producing such enormous numbers of black flies that even long-distance
movements did not disperse the black flies too thinly to prevent them from disturbing livestock.
In 1978 S. luggeri drifted on northerly winds and created problems as far away as Wynyard,
some 170 km south of the Saskatchewan River. Then, when winds changed to easterly for a few
days, some of these same black flies were redirected toward Saskatoon and beyond, apparently
travelling a total distance of more than 300 km.

S. luggeri also regularly infiltrated forested areas against prevailing winds. Pastures along
the northern edge of the Nisbet Forest reserve near Choiceland and Smeaton were chronically
affected by dense swarms of S. luggeri regardless of wind directions. Every summer swarms
moved through trees, often against prevailing winds, at least 30 km from breeding sites in the
Saskatchewan River.

3. S. luggeri adults caused hyperactivity, panic and stampeding in livestock by swarming
densely around the animals’ heads. Animals ceased grazing and breeding when under severe
attack. Calves could not nurse properly when herds were tightly packed and in constant turmoil
and with peripheral animals forcing themselves into the herds. Calves also suffered
malnutrition when milk flow from cows was reduced. Also, since S. luggeri adults bit animals
around their eyes and udders, they may well have been instrumental in carrying disease
organisms responsible for bovine keratitis and mastitis. Increased incidences of these diseases as
well as stress-related diseases such as hoof rot in mature animals, and pneumonia and scours in
calves always accompanied severe outbreaks of S. luggeri. In comparison, S. arcticum adults
did not swarm intensively around animals’ heads but concentrated their attacks along
underlines, often unnoticed by victims. No animals died suddenly after being bitten by S.
luggeri as had happened frequently with S. arcticum (Fredeen, 1981). However, during
outbreaks of S. luggeri some animals died during stampedes, suffered broken limbs and had to
be destroyed, or died from stress-related diseases. Pastures were unevenly grazed when cattle
refused to stay in rich lowlands. Supplementary feeding was required, especially for bulls, as
well as additional veterinary and labour services.

4. Unlike those of S. arcticum , adults of S. luggeri actively attacked people as well as other
warm-blooded animals, except birds.

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Fredeen

5. A major cause for ineffectiveness of larviciding against S. luggeri became apparent in
later years when tests showed that a single injection of methoxychlor into the shallow, weedy
Saskatchewan River often was not effective beyond 20 to 30 km downstream. In comparison, a
single injection into the North Saskatchewan River in 1973 (fairly weed-free at that time) had
been reasonably effective against S. arcticum larvae throughout at least 160 km (Fredeen,
1975).

6. An unexpected effect of larvicide treatments was that certain larvae removed from the
Saskatchewan River (south branch in particular) apparently drifted downstream to reattach in
suitable sites, often in the main Saskatchewan River downstream from the confluence of the
south and north branches (Fredeen, 1983). Major outbreaks in 1977, 1978 and 1982 (not
discussed in this paper) were believed to have resulted, at least in part, from downstream
accumulations of reattached larvae following larvicide treatments further upstream.

For these reasons, sporadic outbreaks of S. luggeri continued to plague portions of
east-central Saskatchewan in 1976 through 1982 during development of abatement strategies
and studies of environmental effects of larvicide injections into the Saskatchewan River.

COMPLAINTS FROM RESIDENTS IN REGIONS AFFECTED BY OUTBREAKS IN

1978

Outbreaks of S. luggeri in 1978 were more widespread and damaging than in any year since
outbreaks commenced in 1972. In 1978 I received more than 100 written complaints and
petitions from at least 46 mailing centers, and many telephone calls from these and other
districts. Most complaints of sustained severe attacks came from within about 100 km of the
river, but others came from up to or even beyond 200 km (Fig. 4). In many districts there were
few remissions from outbreaks from June through August.

A selection of quotations from people personally attacked in 1978 included: “dense swarms
of black flies attacked like angry bees all day; many people wore netting when working outside,
others carried portable smudges; repellents did not provide sufficient protection; fence and
machinery repairing, construction work, telephone line repairs and other outside jobs were at
times impossible, even with use of repellents; repellents did not prevent densely-swarming black
flies from entering nose, mouth, eyes and ears; some people required medical attention for bites;
rural and urban people alike lost many days of out-of-doors work and recreation, children and
older people were unable to work or play out-of-doors; farm work was neglected when cattle
required so much additional attention; air cleaners on tractors required daily servicing to
remove black flies.”

Remarks from people concerned about welfare of livestock in 1978 (owners, veterinarians
and government employees) included: “milk production greatly reduced when animals unable
to graze; increased incidence of mastitis believed caused by black flies which sometimes caused
udders to drip with blood; cows with irritated udders produced less milk and could not tolerate
nursing calves; cattle were in constant motion and stampeded frequently because of black flies
swarming densely around their heads; a stampeding herd tore down a fence; cattle refused to
graze outside until after sunset and thus required supplementary food; cattle bunched on
hilltops or near farmyards; where there was access to mud or manure beds, cattle stood or lay in
them; cattle crowding into a shed pushed out rear wall; cattle became noticeably thinner in
pastures instead of showing expected weight gains; cattle unable to breed; bulls became
impotent because of black fly attacks; calves burnt when pushed by herds into smudges; cattle

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

191

broke shoulders and limbs during stampedes; cattle in feed lots sold prematurely to avoid
further weight losses; greatly increased incidences of bovine keratitis, mastitis, foot rot,
calfhood scours and pneumonia blamed on extraordinary physical and nutritional stresses
imposed by severe black fly attacks; livestock owners found it difficult to approach or herd
animals in pastures; extra riders hired to cope with hyperactive herds; horses difficult to
manage when under attack; horses, cattle, sheep and hogs had to be fed indoors; dogs severely
bitten; moose and elk submerged themselves in water and were oblivious to human presence.”

COMPLAINTS FROM PATRONS AND MANAGEMENT OF ONE LARGE PASTURE

Following the severe outbreaks of 1978, Mr. Stephen Burkell, Director of Pastures for the
Prairie Farm Rehabilitation Administration for northeastern Saskatchewan, and I were invited
to attend an annual meeting of directors and management of the James Smith Pasture to
discuss complaints. This pasture, with an area of 3,230 ha, is located north of Kinistino (Fig. 4)
and has, as its northern boundary, the main Saskatchewan River from whence repeated
outbreaks of S. luggeri had emerged in 1978 and in two previous years.

Eight hundred and forty cows, many accompanied by suckling calves had been received into
this pasture during the last week of May in 1978 and discharged 140 days later in mid-October.
Patrons had expected cows and suckling calves to gain significantly in weight and had expected
timely conceptions of cows from use of 35 high quality, purebred bulls. Management had
expected minimal handling problems and uniform grazing of grasslands. Due to severe and
prolonged outbreaks of S. luggeri these expectations were not realized.

Approximately 260 ha of lowlands in this pasture, including some of the richest grasslands,
were not grazed in 1978. Although cattle were herded into those lowlands several times that
summer, they would not stay because of continuous presence of black flies. Instead, cattle often
congregated on overgrazed hilltops, apparently to take advantage of winds.

Management reported that five purebred bulls out of the 35 in service that summer had to
be replaced when they became impotent due to severe debilitation and infections of the sheath
and scrotum from black fly bites. Replacement costs totalled $11,250.00 among bulls alone,
despite daily supplemental feeding and adequate veterinarian services (Table 2).

Of 840 breeding cows in that pasture in 1978, there were only two fatalities attributed
directly to black fly attacks (Table 2). Both animals suffered broken limbs during stampedes
and had to be destroyed. Prompt attention saved many other sick animals, especially those with
bovine keratitis (pinkeye), mastitis and foot rot.

Another loss, more difficult to assess, was an increase in proportions of cows showing
delayed conceptions (Table 2). Our observations during outbreaks in 1978 confirmed
producers’ complaints that breeding was completely interrupted whenever animals came under
severe attack. Mr. Allan Blair, Livestock Specialist for Saskatchewan Agriculture in
east-central Saskatchewan, estimated that more than 20 percent of all cows in outbreak areas
conceived at least one month later than expected in 1978. A calf born one month late the
following year would have been about 30 kg lighter than expected at weaned-calf sales in the
fall. The average price for weaned calves in Saskatchewan in the fall of 1979 was $2. 20/kg,
indicating losses of $66.00 for each weaned calf born one month later than expected. Thus if 20
percent of the cows in this pasture were affected in 1978, the resultant 168 conceptions, late by
only one month each would have caused a loss of more than $1 1,000.00 in the following year
(Table 2). Losses from conceptions delayed for more than one month were not estimated. Allan

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192

Fredeen

Blair estimated that the number of cows not conceiving at all in 1978 increased by at least two
percent over that observed in 1977. However, conception rates also vary between herds
according to culling and other management practices.

Ryan and Hilchie (1980) reported that, in areas of central Alberta affected by severe
outbreaks of S. arcticum from the Athabasca River in 1978, 44 percent of calves born the
following year were later than expected. In areas less affected by outbreaks, 21 percent of all
calves were born later than expected. They also reported that non-conception rates varied from
8.41 to 12.17 percent in severely affected regions, but only 1.26 to 6.35 percent in areas less
affected.

The major loss among beef cows in the James Smith pasture undoubtedly consisted of
unrealized weight gains. Livestock owners and pasture managers unanimously claimed that
their animals had lost weight during the 140-day grazing season in 1978 whereas past
experience had led them to expect gains of about one kg per day. Photographs taken of herds in
that pasture indicated that animals actually appeared thin when compared with unaffected
animals from pastures further south. However, animals were not weighed in or out of the James
Smith pasture and thus owners’ claims could not be verified. To estimate values of unrealized
weight gains I have used data obtained in 1982 from two herds of purebred hereford cattle
pastured near Choiceland, Saskatchewan. In one of those herds, partly protected from attacks
by S. luggeri with use of fenvalerate-impregnated ear tags (BovaidR, cows with suckling calves
showed average individual weight gains of 0.967 ± 0.342 kg per day, whereas cows from an
adjacent “unprotected” herd showed average gains of only 0.508 ± 0.270 kg per day, a
difference of about 0.46 kg per day. BovaidR-“protected” dry yealings without calves showed an
average advantage of about 0.57 kg per day over “unprotected” dry yearlings. Thus for the
James Smith herd, of 838 surviving cows I have assigned weight gains of 0.5 kg per day per cow
in 1978 instead of 1.0 kg that might have been realized had there been no black flies. Even this
partial loss would have cost producers $102,655.00 (Table 2).

There were 700 suckling calves in the James Smith pasture in 1978 and deaths of 12, valued
at $4,230.00, could be attributed directly to black fly attacks. Some of these calves had suffered
fatal trampling injuries; others had died from nutritional and physical stress-related diseases
such as scours and pneumonia. But again, main losses were unrealized weight gains due in part
to delayed conceptions the previous year as discussed above and in part to hyperactivity and
inability to suckle or graze properly in 1978. Milk production from beef cows in 1978 would
have suffered long-term reductions similar to those for dairy cows, discussed later. Owners
claimed that the average weight of a weaned calf from this pasture at fall sales of weaned
calves was about 135 kg as compared to about 180 kg for weaned calves from pastures further
south and less affected by black flies. This indicated that unrealized weight gains for suckling
calves in the James Smith pasture averaged 0.32 kg per day per calf. Producers did not provide
proof of these claims. Thus I have calculated losses on the basis of data obtained from test herds
near Choiceland in 1982 and Prince Albert in 1979. At Choiceland, suckling calves from a herd
partly protected from black flies for 100 days with the use of BovaidR ear tags, gained an
average of 0.121 kg more per day than similar calves in a nearby “unprotected” herd. Similar
tests with Aberdeen Angus near Prince Albert in 1979 showed that calves from a herd
protected for about 18 days with permethrin spray gained, on the average, 0.092 kg more per
day than calves from a nearby “unprotected” herd.

Thus, while there were no reasons to doubt producers’ claims of unrealized weight gains of
0.32 kg per calf per day for animals severely affected in the James Smith pasture in 1978, I

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

193

have assigned losses of 0.1 kg per calf per day. This would have amounted to an average loss for
the 140-day grazing season of $28.00 per calf for a herd total of $19,260.00 (Table 2). This
would have been additional to losses caused by delayed conceptions calculated earlier.

While compensatory weight gains could have occurred after black fly outbreaks ceased in
the fall, losses calculated from unrealized weight gains described above for pastured cattle
should be considered real losses for producers. Many animals, especially suckling calves and
yearlings, were sold in the fall soon after leaving summer pastures, before compensatory gains
could occur. Other animals, overwintered on home farms, could have achieved compensatory
gains only at the expense of forages grown in home pastures or of forages especially harvested
for use in winter.

Thus, total losses, conservatively estimated for the 140-day grazing season in 1978, in this
one community pasture, apparently exceeded $150,000.00 (Table 2). Costs not included in
these calculations were veterinarians’ fees and medications required to treat 260 cases of bovine
keratitis, mastitis, foot rot, scours and pneumonia in excess of those treated the previous year,
as well as costs of supplementary feeding, fence repairs, daily smudge building, increased
animal insurance costs and hirings of extra riders to handle hyperactive herds and to monitor
herds for sick animals.

ESTIMATED FINANCIAL LOSSES TO BEEF PRODUCERS IN AREAS OF
EAST-CENTRAL SASKATCHEWAN AFFECTED BY SEVERE OUTBREAKS OF S.

LUGGERI IN 1978

Regions where repeated complaints of damage to livestock originated during outbreaks in
1978 (Fig. 4) included about 5,700 km2 (approximately 0.5 percent of Saskatchewan’s Crop
District 6, 13.0 percent of District 8 and 1.5 percent of District 9). Cattle populations in those
portions of the three districts included at least 650 bulls, 11,000 beef cows, 9,500 suckling
calves, 1,100 dairy cattle and 7,000 other cattle (mainly 1- to 2-year old steers, heifers and
bulls). Practically all would have been either purebred or high-quality crossbred animals. Some
bulls and other cattle would have had access to indoor feeding during outbreaks.

In September, 1978, 39 livestock owners in six municipalities (Numbers 399, 400, 429, 430,
459 and 460, Fig. 6) were asked by the local Agricultural Representative, Eugene Bendig, to
complete a form indicating effects of black fly outbreaks on their farms that year. Eleven
reported that animals had to be housed much of the summer, three reported premature sales of
animals including some animals blinded by bovine keratitis believed transmitted by infected
black flies, and 10 reported fatalities caused by black flies of 14 calves, nine yearlings, three
cows and three bulls. All claimed that pastured animals became thinner during the summer and
also that outbreaks caused prolonged and severe disruptions of all outdoor activities.

George O’Bertos, Director, Saskatchewan Lands Branch, Tisdale, reported that in every
provincial community in his jurisdiction, 12 to 20 calves in each population of 800 to 1,000
calves per pasture were killed by black flies that summer. These calf fatalities of 1.5 to 2.0
percent were comparable to the loss of 1.72 percent reported that same year for the James
Smith pasture.

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194

Fredeen

107° 106° 105° 104°

I I I I

4*4 Forest reserve

0 5 0 1 0 0 K m

1 I 1

Fig. 6. Map of central Saskatchewan showing locations of 6 provincial community pastures in which cattle were reported
to have been affected by black flies in recent years, and 1 2 municipalities affected by different intensities of outbreaks of
Simulium luggeri in 1976 through 1981.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

195

TABLE 2. ESTIMATED VALUES OF BEEF CATTLE LOSSES IN THE JAMES SMITH
COMMUNITY PASTURE NORTH OF KINISTINO, SASKATCHEWAN DUE TO
REPEATED OUTBREAKS OF BLACK FLIES, SIMULIUM LUGGERI IN 1978.

Number of

Net loss per

animals in

Animals affected

affected

Total value

Category

pasture

Type of loss

(no.)

(%)

animal ($)

of losses ($)

Bulls

Fatalities

0.00

Replacements

14.30

2,250.00(1)

11,250.00

Cows

840

Fatalities

0.24

787.50(2)

1,575.00

Delayed

conceptions

168

20.00

66.00(3)

11,088.00

Unrealized

weight gains

838

99.76

122.50(4)

102,655.00

Calves

700

Fatalities

1.72

360.00(5)

4,320.00

Unrealized

weight gains

688

98.28

28.00(6)

19,260.00

TOTAL

LOSSES

150,148.00

(,)Replacement cost ($3,000.00) minus commercial sale value ($750.00).

(2) Estimated 450/kg animal at $1. 75/kg.

(3) Conception estimated to be at least 1 month late, with the weaned calf weight about 30 kg
lighter than expected the following year with an average value of $2. 20/kg.

(4) Estimated 70 kg unrealized weight gain/cow at the end of the 140-day grazing season, with
an average value of $ 1.75/kg.

(5) Estimated 180 kg/weaned calf at $2.00/kg in the fall of 1978.

(6) Estimated 14 kg unrealized weight gain/calf at the end of the 140-day suckling season at
$2.00/kg.

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Fredeen

These data suggest that effects of outbreaks in 1978 on livestock outside the James Smith
pasture were similar to those within. On that basis the following losses were calculated: If
one-seventh of the 650 bulls died or required replacement at an average net loss of $2,250 per
bull, losses from this source would have totalled $209,000. An estimated 26 beef cows and 160
suckling calves, valued at more than $78,000, may have been killed. If 20 percent of the 1 1,000
beef cows suffered delayed conceptions equivalent to single oestrus cycles with an average net
loss of $66 per calf at weaned calf sales the following year, losses from this source alone would
have totalled $145,200. But the largest financial losses would have resulted from unrealized
weight gains for 11,000 beef cows, 9,500 suckling calves and 7,000 weaned cattle. Despite
claims by most producers that their animals had actually lost weight during the summer-long
outbreaks in 1978, I estimated from our tests reported earlier that suckling calves may have
gained an average 0.1 kg per day (14 kg for the grazing season) and cows and weaned cattle 0.5
kg per day (70 kg for the season) instead of anticipated seasonal gains of 28 kg and 140 kg
under black fly-free conditions. On this basis, unrealized weight gains for some 9,500 calves in
severely affected areas in 1978 may have cost producers at least $266,000 and for some 18,000
cows and immature cattle, $2,205,000. Accumulated losses for all classes of beef cattle in
severely affected areas thus were estimated to have exceeded $2,903,000. These losses were
calculated for only those areas totalling about 5,700 km2 considered to be severely affected by
black fly outbreaks in 1978. Losses in an additional 32,000 km2 less severely affected (Fig. 4)
were not included in these calculations. Also excluded were estimates of costs of supplementary
feeding, especially for cattle kept indoors, supplementary feeding, especially for cattle kept
indoors, supplementary labour, veterinary services, repairs to fences and barns, and increased
insurance against future losses. Thus, actual losses to the entire beef cattle industry in
east-central Saskatchewan in 1978 were believed to have been much larger than the $2,903,000
shown in these calculations.

TRENDS IN CATTLE POPULATIONS IN PROVINCIAL COMMUNITY PASTURES
AFFECTED BY BLACK FLY OUTBREAKS

Livestock producers anticipate advantages in committing cattle to community pastures.
These pastures offer expansion of production beyond home pasture capacities as well as
opportunities for quality grazing and quality breeding from registered herd sires or artificial
insemination (A.I.). However, in large pastures animals cannot be given the same kind of
individual attention available in small home pastures. Producers claimed that during black fly
outbreaks their animals were subjected to severely debilitating stresses causing weight losses,
sickness and missed oestrus cycles whether in natural or A. I. breeding programs. Because of
this some producers ceased committing cattle to large pastures in regions prone to outbreaks of
S. arcticum until 1973 and of S. luggeri after 1975. However, their places in pasture quotas
generally were filled by other producers hopeful of improvements in black fly control measures.

There are three types of community pasture organizations in Saskatchewan: provincial,
federal and co-operative. To investigate complaints from producers and pasture managers,
numbers of cattle committed annually to six provincial community pastures in municipalities
frequently affected by black flies (those bordering the Saskatchewan River between Wingard
and Nipawin) (Fig. 6) were compared with numbers in all other 48 provincial community
pastures in the province, 1969 to 1981, inclusive (Saskatchewan Agriculture,, 1970-1982).

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

197

On average, less than 85 percent of official carrying capacities of “affected pastures” was
used 1969 to 1981, inclusive, as compared with more than 89 percent for pastures in the
remainder of the province (Fig. 7 A). The difference was significant (P = 0.05). Pasture
managers reported that some of the richest lowland areas often were undergrazed because
cattle congregated on windy hilltops to avoid black fly attacks. Reduced usage of “affected”
pastures was particularly noticeable before 1973 and after 1977.

Percentages of breeding cows among populations of mature cattle committed to “affected”
pastures each year were consistently lower (some by as much as 14 percentage points) than
populations in pastures in the remainder of the province (P = 0.01) (Fig. 7B). This supports
claims by some owners that they withheld breeding cows with expectations of obtaining
improved conception rates in home pastures.

Percentages of suckling calves among breeding cows committed to “affected” pastures were
lower than in other provincial pastures in nine of the 13 years (Fig. 7C) but means for the 13
consecutive years did not differ significantly. In particular, conception rates appear to have
been reduced during black fly outbreaks 1969 to 1973 ( S . arcticuml ) and 1976 to 1981 ( S .
luggeri ) as claimed by livestock owners. In 1979, 6.5 percent fewer cows were accompanied by
calves in “affected” pastures than in other provincial pastures, suggesting that average
conception rates had been reduced by that amount during the severe outbreaks of 1978.

TRENDS IN CATTLE POPULATIONS AND LAND-USE PATTERNS IN DISTRICTS

SUBJECTED TO CHRONIC OUTBREAKS OF S. LUGGERI , 1976 THROUGH 1981

Many livestock producers in outbreak areas stated that they reduced or even eliminated
herds because black flies rendered their operations less profitable than expected. While
numbers of dairy and beef cattle did decline in the entire province, 1975 to 1981, presumably
for economic reasons, reductions were greatest in Crop District 8 (Figs. 1, 8) (Saskatchewan
Agriculture,, 1976, 1982). During those seven years, numbers of beef cattle declined by 27
percent in Crop District 8, 19 percent in District 9, and 21 percent in the remainder of the
province. Numbers of dairy cattle declined by 32, 20 and 6.5 percent in those same three
districts.

On a finer scale, largest declines occurred in municipalities bordering the Saskatchewan
River (Fig. 6, Table 3). In three such municipalities where average distance from farm to river
was about 13 km, dairy cattle numbers declined by 70.3 percent between 1971 and 1981
(Statistics Canada, 1973, 1983). Numbers of “other” cattle declined by 29.2 percent. At an
average distance of 43 km, numbers declined by 52.5 and 15.1 percent, at 74 km by 47.3 and
1.7 percent, and at 101 km by 54.0 and 1 1.7 percent for dairy and “other” cattle, respectively.

Amounts of land devoted to improved pasture declined by 32.4 percent between 1971 and
1981 in municipalities averaging 13 km from the river, by 46 percent at 43 km and by 6.9
percent at 74 km, but increased by 10.4 percent at 101 km (Table 3). Amounts of land devoted
to cultivated crops did not change in inverse proportions as expected. However, much of the
land adjacent to the valley of the Saskatchewan River is classified as unsuitable for cultivation
on account of steep contours, stoniness, light soil textures subject to wind erosion, and even high
water tables. Thus it appears that some land withdrawn from use as cultivated pastures may
have been abandoned when it became uneconomical to produce livestock. For instance, on
farms averaging 13 km from the river in the three municipalities studied, 1,237 ha of land
formerly classified as pastures, were unaccounted for in the Canadian Census of 1981,

Quaest. Ent., 1985, 21 (2)

198

Fredeen

Fig. 7. Trends in populations of mature cattle, breeding cows and suckling calves, 1969 through 1981, committed to six
provincial community pastures (see Fig. 6 for locations), in municipalities adjacent to the Saskatchewan River between
Wingard and Nipawin in Saskatchewan (a region affected by chronic outbreaks of black flies), compared with numbers
committed to community pastures in the remainder of the province: A - populations of mature cattle expressed as
percentages of carrying capacities of those pastures; B - populations of breeding cows - expressed as percentages of total
adult cattle assigned to those pastures; C - populations of suckling calves expressed as percentages of total breeding cows
assigned.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

199

TABLE 3. TRENDS IN CATTLE POPULATIONS, AND IN LAND AREAS DEVOTED
TO IMPROVED PASTURES AND TO CULTIVATED CROPS, BETWEEN 1971 AND
1981, IN FOUR SETS OF MUNICIPALITIES LOCATED AT DIFFERENT
DISTANCES FROM THE SASKATCHEWAN RIVER

Trends, 1971 to 1981 (%)<2>

Average

Cattle

from river

(excluding

Improved

Cultivated

Municipalities0*

(km)

dairy cows)

Dairy cows

pasture

crops

368, 369, 370

101

-11.7

-54.0

+ 10.4

+ 6.3

398, 399, 400

- 1.7

-47.3

-6.9

+ 2.3

428, 429, 430

-15.1

-52.5

-46.0

+ 7.1

459, 460, 487

-29.2

-70.3

-32.4

+ 0.1

(1) See Figure 6 for locations of municipalities in relation to the Saskatchewan River.

(2) Calculated from census data (Statistics Canada, 1973, 1983).

suggesting abandonment.

LOSSES SUSTAINED BY MILK PRODUCERS DURING OUTBREAKS OF S.

LUGGERI

Many dairy herds in central Saskatchewan are either kept indoors or at least allowed free
access to barns. By 1982, in those areas around Prince Albert chronically affected by black fly
outbreaks, only five of 18 producers still pastured their milking cows outside all summer. A
number of other dairies ceased operations in recent years because they were unwilling to
convert, in the face of chronic black fly outbreaks, to housing that would have been both capital
and labour intensive.

Data about milk-shipments were obtained from three dairies for five consecutive years, 1977
to 1981, to determine whether outbreaks of S. luggeri affected productivity. None of 15 other
dairies visited near Prince Albert were able to provide uninterrupted data for this 5-year period
or even for the one year especially investigated, 1978. Dairies “A” and “B” (Fig. 4, 1978) were
located in a region subjected to relatively severe outbreaks of S. luggeri at least during the first
three summers of this study. Outbreaks at “c” were less severe. I did not obtain data from
regions completely free of black fly outbreaks.

Quaest. Ent., 1985, 21 (2)

PERCENTAGE OF 1975 POPULATION =100

200

Fredeen

Fig. 8. Annual trends in populations of cattle on Saskatchewan farms expressed as percentages of 1975 populations in Crop
Districts 8 and 9 (see Fig. 1 for locations) near the Saskatchewan River, and in the remainder of Saskatchewan: A - beef
cows; B - milking dairy cows.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

201

4000

</>

3000 -

****••... Producer

i i

F M

MONTH

i i

N D

Fig. 9. Milk shipments (2-day accumulations) throughout 1978 from three dairies in areas of central Saskatchewan
affected by chronic outbreaks of black flies ( Simulium luggeri) (see Fig. 4, 1978, for locations of dairies). Arrows indicate
commencement of outbreaks on May 26 and approximate ending 74 days later on September 7, 1978.

Producer “A” always pastured his milking cows outside all summer because of an
abundance of rough, wet pasture lands that could not be used for grain or hay crops. He always
provided supplementary cut fodder, outside, as required, in addition to the usual grain
concentrates at milking time.

Milking cows of producer “B”, located five to six km southeast of “A” were confined to the
barn at all times. Producer “C” was located about 75 km southwest of “A” and “B”, and
although his cattle spent much time out-of-doors, they were allowed free access indoors at all
times. He reported that his animals voluntarily remained indoors during fly outbreaks and were
fed there.

These dairymen attempted to maintain uniformly high production to fulfill official milk
quotas by providing high quality rations and by having cows freshen at uniform intervals. They
reported that milk production increased significantly after cattle commenced grazing new grass

Quaest. Ent., 1985, 21 (2)

202

Fredeen

(%) Nononaodd

Fig. 10. Trends in volumes of milk shipments from three dairies in central Saskatchewan (see Fig. 4 for locations) in each of 5 years, 1977 through 1981. The “100-percent” starting point
each year indicates average daily production throughout 10 days immediately before the beginning of outbreaks. Successive points thereafter at intervals of 100, 200 and 300 days after
commencement of outbreaks represent percentages of that “pre-outbreak” production each year, also calculated from 10-day means.

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

203

TABLE 4. MILK PRODUCTION FROM THREE DAIRIES IN CENTRAL
SASKATCHEWAN BEFORE AND AFTER COMMENCEMENT OF BLACK FLY
OUTBREAKS (S. LUGGERI N. AND M.) ON MAY 26, 1978.

Average daily production during two
175-day periods, one immediatedly
before and one immediately after
commencement of outbreaks

Value of difference
per cow per day**

Producer*

Before (kg)

After (kg)

Difference (kg)

(S)

585

480

-105

-0.95

1395

1605

+ 210

+ 0.71

954

974

+ 20

+ 0.11

*See Fig. 4 for locations of these dairies.

"Average number of lactating cows for producer A = 30, B = 80, C = 50. These producers
received $0.27 gross per kg of milk.

in the spring, but that it declined when black fly outbreaks commenced a few weeks later unless
animals were protected. Producer “A” reported that production did not return to expected
levels during lulls in outbreaks but only after affected animals commenced new lactation cycles,
in many instances long after outbreaks had ceased in the fall. This was particularly noticeable
when stress had been severe and long-lasting, as in 1978. Milk shipment data provided by “A”
indeed showed both immediate and long-term effects (Fig. 9). In 1978 outbreaks commenced
on May 26 and a decline of 3.0 percent in milk production was already evident in the 2-day
accumulation shipped on May 27. By the time outbreaks ended in early September production
had declined by about 40 percent from what it had been during the week before outbreaks
commenced on May 26. This decline did not end until mid-November, about 175 days after
outbreaks had commenced. In contrast, production from herds “B” and “C” increased during
this same 175-day outbreak period (Table 4). Similar long-term effects from outbreaks of S.
luggeri were noted in 1979, but not in 1977, 1980 or 1981 (Fig. 10) when outbreaks of black
flies south of Prince Albert were relatively light.

In 1978 producers received $0.27 per kg of raw milk. Throughout the first 175 days after
outbreaks commenced on May 26, the decline in production in herd “A” resulted in an average
reduction in gross returns of $0.95 per cow per day when compared with average production in
that same herd throughout 175 days immediately preceding May 26 (Table 4). In contrast,
production increased in herds “B” and “C” between these same two 175-day periods, resulting
in average increases in gross returns of $0.71 and $0.1 1 per cow per day. Production trends in
herds “A” and “C” may be compared because both herds were pastured during the summer of

Quaest. Ent., 1985, 21 (2)

204

Fredeen

1978. Management in those two herds differed mainly in that herd “A” grazed out-of-doors all
the time but herd “C” had free access to a barn especially during outbreaks and were fed
indoors as required. The difference in average gross returns between these two herds for the two
consecutive 175-day periods was $1.06 per cow per day for a total of $185.50 per cow. Total
confinement indoors as in herd “B” resulted in an even larger improvement in gross returns.

In those portions of Crop Districts 8 and 9 totalling some 5,700 km2 that were most severely
affected by black flies in 1978, there were estimated to have been about 310 lactating dairy
cows (including herd “A”), which did not have free access to barns during outbreaks. Assuming
that those animals were affected similarly to those in herd “A”, lost production would have
exceeded $57,500 during the first 175 days after outbreaks commenced. Additional to this
would have been costs of supplementary feeding, veterinary services, extra manpower required
to handle hyperactive herds, as well as general milk volume reductions in an additional 32,000
km2 of farm lands that were less severely affected by black flies. Producers also claimed that
there were some delayed conceptions as noted for beef cattle. Thus total losses to the dairy
industry in Saskatchewan due to outbreaks of S. luggeri in 1978 must have greatly exceeded
$57,500.

DISCUSSION AND CONCLUSIONS

The purpose of this paper was to catalogue losses attributed to outbreaks of the black fly S.
luggeri in Saskatchewan, and to attempt to evaluate losses to determine whether demands for
abatement programs by livestock producers and other residents of east-central Saskatchewan
were justified.

Residents in this part of Saskatchewan have had to contend with black fly problems since
the earliest days of settlement because the Saskatchewan River has always provided breeding
sites for large numbers of larvae. Until about 1976, outbreaks of S. arcticum could be expected
every year. Larvae of S. luggeri replaced those of S. arcticum when the river became shallow,
clear and weedy every summer, mainly due to upstream storage of water in new hydropower
reservoirs. Annual outbreaks of S. luggeri commenced on a large scale in 1976 and have
continued to the present time although moderated on many occasions by experimental
injections of methoxychlor larvicide into the Saskatchewan River.

Animals did not die suddenly after attacks by S. luggeri as had happened with S. arcticum.
Nevertheless, outbreaks of S. luggeri inflicted significant losses for several reasons. Losses were
enhanced, in part, by emergence of enormous numbers of adults from larvae breeding in vast
new weed beds in the Saskatchewan River. Also because S. luggeri is multivoltine, in the
absence of larviciding, numbers increase during summer months. In 1978 during the worst
outbreaks on record, black flies emerged from the river throughout late May to late September
and spread into at least 38,000 km2 of surrounding countryside. About one-seventh of this area
was subjected repeatedly to severely damaging outbreaks that year. Searches for larvae in other
nearby breeding sites such as the Carrot and Torch Rivers showed that only the Saskatchewan
River was capable of producing outbreaks of such magnitude. S. luggeri swarmed aggressively
around heads of animals on most warm days that summer forcing animals into almost
continuous hyperactivity. Animals stampeded readily, spent much time on windy hilltops and
greatly reduced breeding activities. People were unable to perform normal outdoor activities
because they too were attacked. Producers, who generally had been satisfied with the
abatement program developed to prevent outbreaks of S. arcticum , demanded similar

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

205

protection from outbreaks of S. luggeri. But whereas single larvicide injections, sometimes
confined to a single branch of the Saskatchewan River, generally were sufficient to prevent
outbreaks of S. arcticum , multiple injections both as regards times and sites often were
required for S. luggeri partly because it was multivoltine and partly because single injections
were no longer effective beyond 20 to 50 km in these weedy rivers. Such an intensive larviciding
program would not be considered acceptable on a continuous basis without adequate economic
justification, not only because of cost of larvicide but also because of concerns for non-target
organisms in the river. Larvicide for a single, annual program as intensive as that required in
1979 (Fredeen, 1983) could cost as much as $25,000 at 1983 prices. Studies of environmental
effects of using methoxychlor as a larvicide, including residue persistance and long-term effects
on non-target invertebrates, indicated that environmental effects were negligable and might be
tolerated in the event of proof Gf need for larviciding (Fredeen et al. 1975; Fredeen, 1983).

Studies of outbreaks of S. luggeri , from 1976 to the present time and especially of severely
disruptive outbreaks in 1978 showed that losses to livestock owners could greatly exceed the
cost of even the most expensive larviciding program envisaged. Losses from a single large
pasture (James Smith Community Pasture) containing about 1,575 beef animals were
estimated to have exceeded $150,000 in 1978. Losses included costs of replacing purebred bulls
which had become impotent due to severe debilitation, fatalities of cows and calves, unrealized
weight gains and losses due to delayed conceptions. About 20 percent of the cows were said to
have missed being bred for at least one oestrus cycle that summer and numbers of cows not
bred at all increased by at least 2.0 percent from the previous year. Not included in the
calculations were costs of supplementary feeding, fence repairs, wages for extra riders,
increased insurance costs, veterinarians’ fees and medications for about 260 cases of pinkeye,
foot rot, scours and pneumonia additional to those seen the previous year.

The James Smith pasture represented less than 0.6 percent of the entire area that was
severely affected by outbreaks in 1978 judging by telephone calls, letters and personal
interviews with producers. In this 5,680 km2 area there were many herds of beef cattle
containing about 650 bulls, 1 1,000 cows, 7,000 weaned cattle and 9,500 suckling calves. Losses
estimated on the same basis as those for the James Smith pasture exceeded $2,900,000. Again,
calculations did not include many miscellaneous costs, nor any losses in the remaining 32,000
km2 of farmlands that were occasionally affected by outbreaks in 1978.

Dairy producers, who were unable to provide housing and indoor feeding for their milking
animals, also suffered considerable losses. Average gross financial returns from two pastured
herds in 1978, one having free access to a barn during outbreaks and one without access to
shelter, differed by $185.50 per animal for the first 175 days after outbreaks commenced on
May 26, when compared with returns during 175 days immediately preceding outbreaks. Total
losses for some 310 unprotected milking cows in 5,680 km2 of severely affected farm lands in
1978 were estimated to have exceeded $57,500. Calculations did not include production losses
from the remaining 32,000 km2 of farm lands affected by lighter outbreaks that year or costs of
supplementary feeding, management or veterinarians’ services.

Fear of chronic outbreaks caused many beef and dairy cattle producers to reduce or even
eliminate herds in recent years. Data from provincial and federal government sources indicated
that trends in animal populations and land-use patterns differed between regions affected by
chronic black fly outbreaks and the rest of the province. For instance, between 1975 and 1981
in Crop District 8 (relatively most affected by outbreaks), numbers of beef cattle declined by
27 percent and dairy cattle by 32 percent. In the remainder of Saskatchewan, numbers declined

Quaest. Ent., 1985, 21 (2)

206

Fredeen

by 21 and 6.5 percent during these same six years. Pasture lands were either converted to less
productive uses or abandoned.

Concerns about black fly outbreaks have not lessened since S. lugger i replaced S. arcticum
in the Saskatchewan River in the early 1970’s. Summer-long outbreaks of S. luggeri have on
occasion caused enormous losses to livestock producers. In addition, people are attacked so
vigorously that sometimes they are not able to work out-of-doors. Control measures against S.
luggeri must be continued and it seems most logical to apply control while black flies are in the
larval stages, confined to relatively limited breeding sites in the Saskatchewan River.
Dispersions of adults are unpredictable and often widespread once they have left breeding sites.
Alternative methods of providing protection for livestock and people are under investigation,
but at present best protection is provided with larvicide used in accordance with permits
renewed annually by federal and provincial authorities.

Experimental manipulation of water flow in the Saskatchewan River seems remote because
approved uses already tax limited water resources. Ongoing tests with insecticides against
adults of S. luggeri indicate that ear tags impregnated with synthetic pyrethroids may provide
relief in large herds. Managers of the 3,230 ha James Smith Community Pasture, already
convinced of their usefulness, have not accepted untagged animals in that pasture since the
spring of 1981. Results from larvicide tests with Bacillus thuringiensis serotype H 14,
conducted by the Canada Biting Fly Centre, Winnipeg, should be known by 1985.

It is hoped this paper will help provide a balanced view for decision makers, when used in
conjunction with an earlier paper on environmental effects of use of methoxychlor larvicide
(Fredeen, 1983). Environmental issues involved must be studied in the broadest sense with
concerns balanced between potential effects of chemical larvicides upon non-target species
inhabiting or otherwise using Saskatchewan River water, and potential effects of black fly
outbreaks upon people in their terrestrial environment, if larvicide is used ineffectively or not at
all.

ACKNOWLEDGEMENTS

I am greatly indebted to livestock producers and employees of Saskatchewan Agriculture
including: Jim Armstrong, Eugene Bendig, Carmen Bibby, Allan Blair, Victor Fremont, Dr. J.
R. Jowsey, Frank Kasko, George O’bertos, Chet Piercy, R. E. Regier, Harold Thompson, the
Board of Directors of the James Smith Community Pasture, and many others who assisted with
this project. I am also indebted to Drs. R. H. Elliott, C. F. Hinks and Ginette Seguin-Swartz,
Agriculture Canada Research Station, Saskatoon, for valuable suggestions during preparation
of this paper.

REFERENCES

Environment Canada. 1980 (a). Historical streamflow summary. Saskatchewan, to 1979.

Minister of Supply and Services, Canada; 339 pages.

Environment Canada. 1980 (b). Historical sediment data summary. Canadian Rivers to 1978.

Minister of Supply and Services, Canada; 121 pages.

Environment Canada. 1981. Surface water data, Saskatchewan, 1980. Minister of Supply and
Services, Canada; 188 pages.

Environment Canada. 1982. Surface water data, Saskatchewan, 1981. Minister of Supply and

Some Economic Effects of Outbreaks of Black Flies in Saskatchewan

207

Services, Canada; 188 pages.

Fredeen, F.J.H. 1958. Black flies (Diptera:Simuliidae) of the agricultural areas of Manitoba,
Saskatchewan and Alberta. Proceedings, Tenth International Congress of Entomology,
Montreal 3: 819-823.

Fredeen, F.J.H. 1974. Tests with single injections of methoxychlor black fly
(Diptera:Simuliidae) larvicides in large rivers. Canadian Entomologist 106: 285-305.

Fredeen, F.J.H. 1975. Effects of a single injection of methoxychlor black fly larvicide on insect
larvae of a 161 km (100 mile) section of the North Saskatchewan River. Canadian
Entomologist 107: 807-817.

Fredeen, F.J.H. 1977 (a). Black fly control and environmental quality with reference to
chemical larviciding in western Canada. Quaestiones Entomologicae 13: 321-325.

Fredeen, F.J.H. 1977 (b). Some recent changes in black fly populations in the Saskatchewan
River system in Western Canada coinciding with the development of reservoirs. Canadian
Water Resources Journal 2: 90-102.

Fredeen, F.J.H. 1981. Keys to the black flies (Simuliidae) of the Saskatchewan River in
Saskatchewan. Quaestiones Entomologicae 17: 189-210.

Fredeen, F.J.H. 1983. Trends in numbers of aquatic invertebrates in a large Canadian river
during four years of black fly larviciding with methoxychlor (Diptera:Simuliidae).
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Fredeen, F.J.H., A.P. Arnason, B. Berck and J.G. Rempel. 1953. Further experiments with
DDT in the control of Simulium arcticum Mall, in the North and South Saskatchewan
Rivers. Canadian Journal of Agricultural Science 33: 379-393.

Fredeen, F.J.H., J.G. Saha and M.H. Baiba. 1975. Residues of methoxychlor and other
chlorinated hydrocarbons in water, sand and selected fauna following injections of
methoxychlor black fly larvicide into the Saskatchewan River, 1972. Pesticides Monitoring
Journal 8: 241-246.

Fredeen, F.J.H. and D.T. Spurr. 1978. Collecting semiquantitative sample of black fly larvae
(Diptera:Simuliidae) and other aquatic insects from large rivers with the aid of artificial
substrates. Quaestiones Entomologicae 14: 41 1-431.

Nicholson, H.P. and C.E. Mickel. 1950. The black flies of Minnesota (Simuliidae). Technical
Bulletin 192, Minnesota Agricultural Experiment Station, 1-64.

Paterson, C.G. and J.R. Nursall. 1975. The effects of domestic and industrial effluents on a
large turbulent river. Water Research 9: 425-435.

Rempel, J.G. and A.P. Arnason. 1947. An account of three successive outbreaks of the black
fly, Simulium arcticum , a serious livestock pest in Saskatchewan. Scientific Agriculture 27:
428-445.

Ryan, J.K. and G.J. Hilchie. 1980. Black fly problem analysis, a survey of biting fly problems
in Athabasca county and Improvement District Number 18. Alberta Department of the
Environment; 80 pages.

Saskatchewan Agriculture. 1970-1982, inclusive. Lands Branch Annual Reports, Regina,
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Saskatchewan Agriculture. 1976 and 1982. Agricultural statistics. Statistics and Public
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Shewed, G.E. 1958. Classification and distribution of arctic and subarctic Simuliidae.
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Statistics Canada. 1973. Census of Canada, 1971: Agriculture, Saskatchewan, Catalogue

Quaest. Ent., 1985, 21 (2)

208

Fredeen

number 96-709, 60 tables of data.

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number 96-909, 35 tables of data.

THE NEARCTIC SPECIES OF THE GENUS PSEUDOMYRMEX (HYMENOPTERA:

FORMICIDAE)

Philips. Ward
Department of Entomology
University of California
Davis, CA 95616
U. S. A.

Quaestiones Entomologicae
21:209-246 1985

ABSTRACT

The Nearctic ants of the genus Pseudomyrmex are revised, with the consequent recognition
of ten species, belonging to three species groups: gracilis group (T. mexicanus Roger),
elongatus group (P. cubaensis Forel, stat. nov./ P. elongatus Mayr), and pallidus group (P.
apache Creighton; P. brunneus F. Smith; P. ejectus F. Smith; P. leptosus Ward, sp.nov. [type
locality: Payne’s Prairie. 8 mi. S. Gainesville, Florida ]; P. pallidus F. Smith; P. seminole
Ward, sp.nov. [type locality: John Pennekamp State Park, Munroe Co., Florida]; P. simplex
F. Smith). P. peruvianus Wheeler, stat. nov., described as a variety of P. ejectus, is considered
a distinct species. The following are proposed as new synonyms of P. simplex: P. delicatulus
Forel = P. capperi Forel = P. panamensis Forel = P. vittatus Forel. The Nearctic
Pseudomyrmex are characterized by marked geographical variation, coupled with the frequent
occurrence of sympatric sibling species.

RESUME

Les fourmis Nearctiques du genre Pseudomyrmex sont revisees, avec la reconnaissance resultante de dix especes,
appartenant de trois groupes d’especes: groupe gracilis (P. mexicanus Roger), groupe elongatus (P. cubaensis Forel, stat.
nov.; P. elongatus Mayr), et groupe pallidus fP. apache Creighton; P. brunneus F. Smith; P. ejectus F. Smith; P. leptosus
Ward, sp. nov. [lieu d’origine du type: Payne’s Prairie, 8 mi. S. Gainesville, Florida]; P. pallidus F. Smith; P. seminole
Ward, sp. nov. [lieu d’origine du type: John Pennekamp State Park, Munroe Co., Florida]; P. simplex F. Smith). P.
peruvianus Wheeler, stat. nov., decrite comme une variete de P. ejectus, est considere une esptce distincte. Les noms
suivant sont proposes comme nouveaux synonymes de P. simplex: P. delicatulus Forel = P. capperi Forel = P.
panamensis Forel = P. vittatus Forel. Les Pseudomyrmex Nearctiques sont caracterises par variation geographique
marquee, accompagnes de I'occurrence frequente des especes sympatriques tres similaires.

INTRODUCTION

The predominantly Neotropical ant genus, Pseudomyrmex , is fraught with species-level
taxonomic problems. Among the few species which occur in the United States, those allied to P.
pallidus have been repeatedly confused ( e.g ., Creighton, 1950). This is partly due to the failure
of earlier taxonomists to examine relevant type material, exacerbated by the existence of sibling
species and extensive geographical variation. Preparatory to a more comprehensive revision of
the pallidus group in the Neotropical region, I here present a review of all Pseudomyrmex
species in the United States, most of which belong to the pallidus group. This allows the
methodological groundwork to be presented and permits clarification of the identity of several
common species.

The North American Pseudomyrmex literature is summarized in D. R. Smith (1979).
Creighton (1950) recognized four species, described one more in 1952, and gave a key to the

210

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United States species in 1955. In the present treatment I recognize nine North American
species, of which two are new. In addition I have included one other species which ranges up to
the margin of the Nearctic region in northern Mexico. Since there has been widespread
misidentification of Nearctic Pseudomyrmex , literature records for most species need to be
reconfirmed. I have attempted to determine the identity of species cited in the North American
literature, in those cases where associated museum material has been available for examination.

A salient feature of the Pseudomyrmex pallidus group (and perhaps a characteristic of the
genus as a whole) is the frequent occurrence of two or more closely related species in a given
locality. There are usually small but reliable morphological differences which serve to
distinguish the members of such sympatric assemblages. However these locally diagnostic
characters show a marked propensity towards geographical variation. Thus, establishing the
diagnostic features of species (as opposed to local populations) requires the examination of
specimens from a broad geographical area. In the present context, this has meant examining
material from both North and Central America since most of the Nearctic species are not
confined to the United States. Moreover, since the characters often involve aspects of size and
shape, it has been necessary to make a rather large number of metric measurements in order to
accurately assess the limits of intra- and interspecific variation. The keys to Nearctic species
depend to some extent on these measurements.

Descriptions of the worker caste of each species have been kept concise, with descriptive
details encapsulated in the ranges of 19 measurements and indices which precede the diagnosis
(expanded to 26 metrics for new species). This helps to standardize and economize the
descriptive process. It also deemphasizes the kind of typological thinking which is likely to
impede taxonomic progress in Pseudomyrmex. In this regard, the illustrations should be used
prudently; they represent “typical” specimens, but reference should also be made to the keys
and species descriptions where the known bounds of variation are indicated.

MATERIALS AND METHODS

Collections are referred to by the following abbreviations:

BMNH British Museum of Natural History, London

GCW G. C. & J. Wheeler collection, San Antonio, TX

LACM Los Angeles County Museum, Los Angeles, CA

MCSN Museo Civico di Storia Naturale, Genoa, Italy

MCZ Museum of Comparative Zoology, Cambridge, MA

MHN Museum d’Histoire Naturelle, Geneva, Switzerland

MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, D. D. R.

NHMB Naturhistorisches Museum, Basel, Switzerland
NHMV Naturhistorisches Museum, Vienna, Austria
PSW P. S. Ward collection, University of California, Davis, CA
UCD Bohart Museum of Entomology, University of California, Davis, CA
USNM National Museum of Natural History, Washington, DC
Scanning electron micrographs were taken with a Philips SEM 501, using gold-palladium
coated specimens. Precautions were taken to avoid distortion of the micrograph by (i) careful
positioning of the specimen on the stub, and (ii) measuring the CRT image with calipers and
making any tilt-correction necessary to restore the true proportions.

Nearctic Species of the genus Pseudomyrmex

211

Terms for integument sculpture are taken from Harris’ (1979) glossary.

Metric measurements were made at 50X power on a Wild microscope, with a Nikon
micrometer wired to an Autometronics digital readout. All measurements were made in
millimeters, to the nearest thousandth of a millimeter. Most have been rounded to two decimal
places for presentation here.

The following measurements are cited (when the head is held in full-face, dorsal view, it is
positioned so that the median ocellus and the frontal carinae lie in the same focal plane):

HW Head width: maximum width of head, including the eyes, measured in
full-face, dorsal view (Figure 1).

VW Vertex width: width of the posterior portion of the head (vertex), measured
along a line drawn through the lateral ocelli, with the head in full-face,
dorsal view (Figure 1).

HL Head length: midline length of head proper, measured in full-face, dorsal
view, from the anterior clypeal margin to the midpoint of a line drawn
across the occipital margin (Figure 1).

EL Eye length: length of compound eye, measured with the head in full-face,
dorsal view (Figure 1).

OD Ocellar distance: distance from the middle of the median ocellus to the
midpoint of a line drawn between the lateral ocelli, measured with the head
in full-face, dorsal view (Figure 1).

OOD Oculo-ocellar distance: distance from the middle of the median ocellus to a
line drawn across the posterior margins of the compound eyes (Figure 1)

(this distance is negative in value if the posterior margin of the compound
eye exceeds the median ocellus).

CD Clypeal distance: distance from the anterior clypeal margin to a line drawn
across the anterior margins of the frontal carinae (Figure 1).

MFC Minimum frontal carinal distance: minimum distance between the frontal
carinae, measured with the head in full-face, dorsal view (Figure 1).

EW Eye width: maximum width of compound eye, measured along its short
axis, in an oblique dorso-lateral view of the head.

SL Scape length: length of the first antennal segment, excluding the radicle.

LF1 Length of first funicular segment: maximum measurable length of the first
funicular segment (pedicel), excluding its basal articulation.

LF2 Length of second funicular segment: maximum measurable length of the
second funicular segment.

FL Forefemur length: maximum measurable length of the forefemur,
measured in posterior view (Figure 3).

FW Forefemur width: maximum measurable width of the forefemur, measured
from the same view as FL, at right angles to the line of measurememt of
FL (Figure 3).

DPL Diagonal length of the propodeum: length of the propodeum, measured in
lateral view along a diagonal line drawn from the metapleural lobe to the

Quaest. Ent., 1985, 21 (2)

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metanotal groove (Figure 2).1

BF Length of the basal (= dorsal) face of the propodeum, measured in lateral
view from the metanotal groove to the point on the surface of the
propodeum which is maximally distant from the diagonal propodeal line
(Figure 2).

DF Length of the declivitous face of the propodeum, measured in lateral view
from the metapleural lobe to the point on the surface of the propodeum
which is maximally distant from the diagonal propodeal line (Figure 2).

MP Depth of metanotal groove (“mesopropodeal impression”), measured in
lateral view from the bottom of the metanotal groove to a line drawn across
the dorsal surface of the mesonotum and propodeum.

PL Petiole length: length of the petiole, measured in lateral view from the
lateral flanges of the anterior peduncle to the posterior margin of the
petiole (Figure 4).

PND Petiolar node distance: distance from the anterior margin of petiole to the
maximum height of the node measured from the same view as PL and
along the same line of measurement (Figure 4).

PH Petiole height: maximum height of the petiole, measured in lateral view at
right angles to PL, but excluding the anteroventral process (Figure 4).

PPL Postpetiole length: length of the postpetiole, measured in lateral view, from
the anterior peduncle (of the postpetiole) to the point of contact with the
fourth abdominal tergite (Figure 4).

DPW Dorsal petiolar width: maximum width of the petiole, measured in dorsal
view.

PPW Dorsal postpetiolar width: maximum width of the postpetiole, measured in
dorsal view.

Indices calculated frrom the preceding measurements include the following ratios:

Cl Cephalic index: HW/HL

01 Ocular index: EW/EL

REL Relative eye length: EL/HL

REL2 Relative eye length, using HW: EL/HW
OOI Oculo-ocellar index: OOD/OD

VI Vertex width index: VW /HW

FCI Frontal carinal index: MFC/HW

CDI Clypeal distance index: CD/HL

SI Scape index: SL/HW

512 Scape index, using EL: SL/EL

513 Scape index, using LF2: SL/LF2

FI Forefemur index: FW/FL

PDI Propodeal index: BF/DF

'In Pseudomyrmex DPL is more appropriate than WL (Weber’s length of
the mesosoma (alitrunk), taken from the anterior pronotal margin to the
metapleural lobe) since the articulation of the pronotum with the
mesothorax renders the measurement of WL imprecise.

Nearctic Species of the genus Pseudomyrme x

213

Figures 1-4. Views of a generalized Pseudomyrmex worker, illustrating some measurements. 1 . Frontal view of head. CD,
clypeal distance; EL, eye length; HL, head length; HW, head width; MFC, minimum distance between frontal carinae;
OD, ocellar distance; OOD, oculo-ocellar distance; VW, vertex width. 2. Lateral view of propodeum. BF, length of basal
(= dorsal) face of propodeum; DF, length of declivitous face of propodeum; DPL, diagonal propodeal length. 3. Posterior
view of forefemur. FL, forefemur length; FW, forefemur width. 4. Lateral view of petiole and postpetiole. PH, petiolar
height; PND, petiolar node distance; PL, petiolar length; PPL, postpetiolar length.

Quaest. Ent., 1985, 21 (2)

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Ward

MPI Metanotal index: MP/HW

NI Petiole node index: PND/PL

PLI Petiole length index: PH/PL

PLI2 Petiole length index, using PPL: PPL/PL

PHI Petiole height index, using PPL: PH/PPL

PWI Petiole width index: DPW/PL

PWI2 Petiole width index, using PPW: DPW /PPW

PPWI Postpetiole width index: PPW /PPL

PPWI2 Postpetiole width index, using HW: PPW/HW

SYNOPSIS

Workers of Pseudomyrmex may be recognized by their large compound eyes (REL
0.39-0.61), closely set frontal carinae and antennal insertions (FCI 0.01-0.07), and short
scapes (SI 0.40-0.51). There is a distinct postpetiole and a well-developed sting.

Among the Nearctic fauna I recognize three species groups, whose workers may be
diagnosed as follows (the gracilis group preceding the other two on the basis of its distinctive
size and habitus):

gracilis group (see also Kempf 1958)

Large black, orange, or bicolored species (HW > 1.20), with broad head, large eyes (REL
> 0.50), and relatively long scapes (SI 0.50); frontal carinae subcontiguous; lateral margins
of pronotum angled; petiole usually long, with a distinct anterior peduncle. Erect pilosity
abundant on body and appendages, including propodeum, scapes, and legs.

elongatus group

Small, brown species (HW 0.56-0.75), with elongate head (Cl < 0.80) and long eyes (REL
0.47-0.58); scapes relatively short (SI 0.45); frontal carinae subcontiguous; lateral margins
of pronotum rounded; petiole very short, without a distinct anterior peduncle (PLI 0.65-0.91;
PWI 0.55-0.74). Erect pilosity present on most parts of body, including mesonotum and
propodeum.

pallidus group

Small yellow, orange or brown species (HW 0.55-1.04), with elongate head (Cl 0.75-0.91);
scapes usually short; frontal carinae contiguous or subcontiguous; lateral margins of pronotum
rounded; petiole usually slender, with an anterior peduncle (PLI 0.43-0.69; PWI 0.38-0.65).
Erect pilosity scarce, lacking on the mesonotum and propodeum.

SYNONYMIC LIST OF NEARCTIC PSEUDOMYRMEX SPECIES

gracilis group

P. mexicanus Roger, 1 863
elongatus group

P. elongatus Mayr, 1870

= P. tandem Forel, 1906
P. cubaensis Forel, 1901, stat. nov.

Nearctic Species of the genus Pseudomyrmex

215

pallidus group

P. apache Creighton, 1952
P. brunneus F. Smith2 1877

= P. nigritus Enzmann, 1945, syn. nov.

P. ejectus F. Smith, 1858
P. leptosus sp. nov.

P. pallidus F. Smith, 1855
P. seminole sp. nov.

P. simplex F. Smith, 1877

= P. delicatulus Forel, 1899, syn. nov.

= P. capperi Forel, 1899, syn. nov.

= P. panamensis Forel, 1899, syn. nov.

= P. vittatus Forel, 1912, syn. nov.

Key to species: workers (excluding P. leptosus new species of which the worker is unknown)

1 (a) Erect hairs conspicuous on most parts of body, including mesonotum and

propodeum 2

(b) Erect pilosity very sparse, lacking on mesonotum and (nearly always)

propodeum 4

2 (a) Large, bicolored orange and black species (HW > 1.40); head

approximately as wide as long (Cl > 0.95); petiole with a long anterior
peduncle (PLI < 0.55) (Figure 12); Florida, Texas south to Panama

P. mexicanus Roger, p. 225

(b) Small, unicolored brown species (HW < 0.80); head notably longer than

wide (Cl < 0.80); petiole short (PLI > 0.60) ( e.g ., Figure 6) 3

3 (a) Larger species (HW > 0.64), with shorter eyes (REL2 0.63-0.73), and a

lower, thinner petiole (PLI 0.65-0.78) (Figures 5,6); Florida, West

Indies P. cubaensis Forel, p. 226

(b) Smaller species (HW < 0.64), with longer eyes (REL2 0.73-0.82), and a
higher, broader petiole (PLI 0.76-0.91) (Figures 7,8); Florida, Texas south
to Colombia P. elongatus Mayr, p. 227

4 (a) Head and gaster usually dark brown; small species (HW < 0.82), with a

deep, wide metanotal groove (e.g., Figure 14) (MPI 0.046-0.097); basal
face of propodeum generally shorter than declivitous face (PDI 0.56-1 .07) 5

(b) Head and gaster golden yellow to orange-brown (fourth abdominal tergite
may have darker fuscous patches); variable in size (HW 0.55-1.04),
metanotal groove usually relatively shallow (e.g., Figure 43) (MPI
0.005-0.054); if metanotal groove very deep (MPI > 0.046), then basal
face of propodeum notably longer than declivitous face (PDI >1.10) 6

5 (a) Basal face of propodeum about one half to three quarters the length of the

declivitous face (PDI 0.56-0.75) (Figure 13); petiole with long anterior
peduncle, the node somewhat displaced posteriorly (NI 0.57-0.65); petiole
and postpetiole very broad (PWI 0.54-0.65; PPWI 1.26-1.54) (Figure 15);

2Occuring no farther north than northern Mexico

Quaest. Ent., 1985, 21 (2)

216 Ward

Mexico P. brunneus F. Smith, p. 231

(b) Basal face of propodeum longer (PDI 0.70-1.07) (Figure 14); summit of
petiolar node in a more anterior position (NI 0.48-0.60); petiole and
postpetiole less broad (PWI 0.40-0.52; PPWI 0.93-1.25) (Figure 16);
southeastern United States, south to Costa Rica

P. ejectus F. Smith, p. 231

6 (a) Fourth abdominal tergite (first “gastric” tergite) smooth and strongly

shining, more or less devoid of appressed pubescence; vertex of head
smooth and shining; broad forefemur (FI 0.45-0.52); small species, with
relatively long eyes (HW 0.55-0.74; OI 0.49-0.55; REL 0.52-0.61);

Florida, West Indies, Mexico south to Brazil

P. simplex F. Smith, p. 238
(b) Fourth abdominal tergite subopaque, covered with a (usually dense) mat of
fine appressed pubescence; vertex of head at least slightly coriarious,
weakly shining to subopaque; longer forefemur (FI < 0.45); generally
larger species, with relatively shorter eyes (HW 0.68-1.04; OI 0.54-0.65;

REL 0.39-0.54) 7

7 (a) Eyes short (REL 0.39-0.44); scapes relatively long, subequal to eye length

(SI2 0.90-1.00); median (protruded) portion of anterior clypeal margin
laterally rounded (Figure 9); frontal carinae relatively well-separated, the
minimum distance between them subequal to the basal width of the scape
(MFC 0.033-0.066, FCI 0.034-0.070); southwestern United States,

northern Mexico

P. apache Creighton, p. 229
(b) Eyes longer (REL 0.43-0.54); scapes notably shorter than eye length (SI2
0.68-0.85); median portion of anterior clypeal margin laterally angulate
(< e.g ., Figure 42); frontal carinae variable, often more closely contiguous so
that the minimum distance between them is notably less than the basal
width of the scape (MFC 0.01 1-0.042, FCI 0.015-0.047) 8

8 (a) Larger species (HW 0.87-0.96); frontal carinae relatively well-separated

(MFC 0.029-0.042, FCI 0.031-0.047); eyes relatively short (REL
0.43-0.48); median portion of anterior clypeal margin weakly angulate,

thus appearing tridentate (Figure 44); Gulf states, Mexico

P. seminole sp. nov., p. 237

(b) Smaller species (HW 0.68-0.89); frontal carinae more closely contiguous
(MFC 0.01 1-0.024, FCI 0.015-0.033); eyes averaging a little longer (REL
0.45-0.54); median portion of anterior clypeal margin usually straight
(Figure 42); southern United States, south to Costa Rica

P. pallidus F. Smith, p. 234

Key to species: queens

1 (a) Large, bicolored orange and black species (HW > 1.45); erect pilosity

abundant on most parts of body, including propodeum; petiole with a long
anterior peduncle (PLI < 0.55); Florida, Texas south to Panama

P. mexicanus Roger, p. 225

Nearctic Species of the genus Pseudomyrmex

217

(b) Smaller species (HW < 1.10); either petiole very short and without a
conspicuous peduncle (PLI > 0.55), or erect pilosity sparse (lacking on

propodeum) 2

2 (a) Head densely punctate, and more than 1.5 times as long as wide (Cl

0.57-0.64); petiole short (PLI 0.58-0.76); usually some erect hairs on the

propodeum 3

(b) Head varying from finely punctate, to coriarious, to smooth and shining,
and no more than 1.5 times as long as wide (Cl 0.66-0.86); petiole
relatively long (PLI 0.43-0.58); propodeum essentially lacking erect setae 4

3 (a) Larger species (HW 0.65-0.72, in a sample of 10 queens); eye length less

than one half head length (REL 0.43-0.47, n = 10); petiole moderately
long (PLI 0.58-0.67, n = 10); Florida, West Indies

P. cubaensis Forel, p. 226

(b) Smaller species (HW 0.56-0.58, n = 5); eye length about one half head
length (REL 0.48-0.53); petiole short (PLI 0.66-0.76, n = 5); Florida,

Texas south to Colombia P. elongatus Mayr, p. 227

4 (a) Head and gaster dark brown; small species (HW 0.62-0.75); frontal

carinae closely contiguous (MFC 0.010-0.018, FCI 0.015-0.026); petiole
long and slender, more than twice as long as high (PLI 0.43-0.48) 5

(b) Head and gaster golden yellow to orange-brown (small fuscous patches
may be present on gaster); mostly larger species (HW 0.57-1.03); if HW
< 0.80, then either the frontal carinae tend to be less closely contiguous
(MFC 0.016-0.035; FCI 0.022-0.046) and/or the petiole is relatively short
(PLI 0.48-0.58) 6

5 (a) Petiole and postpetiole relatively broad, the latter about 1.25 times as wide

as long (PHI 0.72, PWI 0.48, PPWI 1.26, in single specimen examined);
frons opaque, fine punctures more or less obscured by coarse coriarious

sculpture; Mexico P. brunneus F. Smith, p. 231

(b) Petiole and postpetiole less broad, the latter about as wide as long (PHI
0.57-0.67, PWI 0.42-0.48, PPWI 1.00-1.12; n = 13); frons usually
weakly shining, distinctly punctulate on a weaker coriarious background;
southeastern United States south to Costa Rica

P. ejectus F. Smith, p. 231

6 (a) Fourth abdominal tergite (first “gastric” tergite) smooth and shining,

appressed pubescence inconspicuous, hairs (if present) separated by about
their lengths; vertex of head usually smooth and shining, with scattered fine
punctures; small species (HW 0.57-0.75), with contiguous frontal carinae
(MFC 0.008-0.021; FCI 0.014-0.029) and relatively long eyes (REL2
0.65-0.80); SI2 0.52-0.68 (n = 17 for this and preceding measurements);

Florida, West Indies, Mexico south to Brazil

P. simplex F. Smith, p. 238
(b) Fourth abdominal tergite weakly shining to subopaque, with a (usually
dense) mat of appressed pubescence; either vertex of head weakly shining
to subopaque, and coriarious with punctures, or SL about three-quarters of
EL (SI2 0.75-0.77); generally larger species (HW 0.66-1.03); frontal
carinae less closely contiguous (MFC 0.016-0.095; FCI 0.022-0.094); eyes

Quaest. Ent., 1985,21 (2)

218

Ward

usually shorter (REL2 0.52-0.71); SI2 0.64-0.95 (n = 61) 7

7 (a) Larger species (HW 0.85-1.03, HL 1.28-1.46); frontal carinae relatively

well separated (MFC 0.053-0.095; FCI 0.052-0.094); eyes relatively short

(REL2 0.52-0.58) 8

(b) Smaller species (HW 0.66-0.92, HL 0.82-1.15); frontal carinae more
closely contiguous (MFC 0.016-0.035; FCI 0.022-0.046); eyes longer
(REL2 0.59-0.71) 9

8 (a) Eye length (EL) more than 1.25 times scape length (SI2 0.70-0.80; n =

10); petiole longer, with a more slender anterior peduncle (PLI 0.43-0.49,

PWI 0.41-0.51; n = 10); Gulf states, Mexico

P. seminole sp. nov ., p. 237

(b) Eye length less than 1.20 times scape length (SI2 0.85-0.95; n = 10);
petiole shorter and broader (PLI 0.49-0.57, PWI 0.54-0.60; n = 10);

southwestern United States, northern Mexico

P. apache Creighton, p. 229

9 (a) Head, especially upper half, smooth and shining, with scattered fine

punctures (Figure 34); occipital margin broadly rounded, so that VI

0.68-0.75 (n= 11); small species (HW 0.67-0.70; n — 11); Florida

P. leptosus sp. nov., p. 233

(b) Head coriarious and weakly shining, punctures coarser (Figure 36); lateral
margins of occiput more sharply rounded, giving head a more quadrate
shape (VI 0.71-0.88; n = 30); larger species, on average (HW 0.66-0.92;

n = 30); southern United States, south to Costa Rica

P. pal lid us F. Smith, p. 234

Key to species: males (excluding P. brunneus F. Smith, of which the male is unknown).

1 (a) Larger species (HW > 1.30); head wider than long (Cl > 1.05); Florida,

Texas south to Panama P. mexicanus Roger, p. 225

(b) Smaller species (HW < 1.00); head longer than wide (Cl < 0.98) 2

2 (a) Posterior margin of pygidium (eighth abdominal tergite) convex and

pointing posteroventrally (Figure 17); posterior margin of hypopygium

straight or broadly convex 3

(b) Posterior margin of pygidium recurved forward, and forming a pocket
which opens anteroventrally (Figure 18); posterior margin of hypopygium
concave 5

3 (a) Head elongate (Cl < 0.80), upper half densely punctate; eyes relatively

long, EL about three-quarters of head width (REL2 0.71-0.83) 4

(b) Head broader (Cl > 0.80) and not densely punctate; eyes shorter, EL
about one-half head width (REL2 0.50-0.57); southwestern United States,
northern Mexico P. apache Creighton, p. 229

4 (a) Larger species, with very elongate head (HW 0.76-0.82, Cl 0.65-0.71, in a

sample of 6 males); eyes short relative to head length (REL 0.49-0.51; n =

6); SI 0.26-0.28 (n = 6); in dorsal view, outer margin of paramere

indented distally (Figure 25); Florida, West Indies

P. cubaensis Forel, p. 226

Nearctic Species of the genus Pseudomyrmex

219

(b) Smaller species, with less elongate head (HW 0.59-0.62, Cl 0.69-0.76; n =

7); eyes relatively longer (REL 0.55-0.58; n = 7); SI 0.22-0.25 (n = 7); in
dorsal view, outer margin of the paramere not notably indented (Figure

24); Florida, Texas south to Colombia

P. elongatus Mayr, p. 227

5 (a) In lateral view, caudal end of paramere consisting of a large dorsal lobe

preceded by a small, dorsal spine (Figures 29, 30); either forefemur rather
broad (FI > 0.36) or posterior margin of hypopygium with a median,

ventral protuberance 6

(b) In lateral view, caudal end of paramere consisting of a dorsal lobe,
unpreceded by a smaller spine (Figures 26-28); forefemur relatively
elongate (FI < 0.36); posterior margin of hypopygium lacking a distinct
ventral protuberance 7

6 (a) Forefemur relatively elongate (FI 0.30-0.36; n = 7); in dorsal view, inner

caudal margin of paramere strongly concave (Figure 23); posterior margin
of hypopygium with a median, ventral protuberance; southeastern United

States south to Costa Rica P. ejectus F. Smith, p. 23 1

(b) Forefemur broader (FI 0.36-0.51; n = 11); in dorsal view, inner caudal
margin of paramere more or less straight (Figure 22); posterior margin of
hypopygium without a distinct ventral protuberance; Florida, West Indies,

Mexico south to Brazil P. simplex F. Smith, p. 238

7 (a) Larger species (HW 0.81-0.92; n = 6); eyes relatively shorter (REL2

0.56-0.59; n = 6); dorsal lobe of paramere much broadened and bicarinate
along its dorsal margin, and invaginated below the margin so that in lateral
view a distinct lunule is visible (Figure 28); Gulf states, Mexico

P. seminole sp. nov., p. 237

(b) Smaller species (HW 0.61-0.84; n = 17); eyes relatively longer (REL2
0.58-0.71; n = 17); dorsal lobe of paramere consisting of a thin lamella,
without a lunule (Figures 26, 27) 8

8 (a) Larger species (HW 0.67-0.84; n = 12); scape generally subequal in length

to second funicular segment (SI3 0.80-1.13; n = 12); in dorsal view, outer
margin of the paramere indented distally, posterior to a small but distinct
protuberance (Figure 20); southern United States south to Costa Rica

P. pallidus F. Smith, p. 234

(b) Smaller species (HW 0.61-0.68; n = 5); scape length exceeding length of
second funicular segment (SI3 1.11-1.25; n = 5); in dorsal view, outer
margin of the paramere lacking a protuberance and showing no distinct
distal indentation (Figure 19); Florida

P. leptosus sp. nov., p. 233

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Ward

Figures 5-12. Pseudomyrmex workers: frontal views of head, and lateral views of petiole and postpetiole. 5,6. P. cubaensis
(Florida); 7,8. P. elongatus (Florida); 9,10. P. apache (Arizona); 1 1,12. P. mexicanus (Texas).

Figures 13-16. Pseudomyrmex workers. 13, 14, lateral views of mesothorax, propodeum, petiole, and postpetiole. 15, 16,
dorsal views of petiole and postpetiole. 13,15 P. brunneus (Mexico); 14,16. P. ejectus (Mexico).

All drawings to same scale; scale line = 0.5 mm.

Nearctic Species of the genus Pseudomyrmex

221

Figures 17,18. Pseudomyrmex males: lateral views of eighth abdominal tergite (pygidium). 17, P. apache (Texas); 18, P
seminole (Florida). Scale line = 0.5mm.

Figures 19-32. Left parameres of Pseudomyrmex males. 19-25, dorsal views, with caudal end uppermost; 26-32, lateral
views, with caudal end to the right. 19,26, P. leptosus (paratype, Florida); 20,27, P. pallidus (Florida); 21,28, P. seminole
(paratype, Florida); 22,29, P. simplex (Florida); 23,30, P ejectus (Texas); 24,31, P. elongatus (Florida); 25,32, P
cubaensis (Florida). Scale line = 0.5 mm.

Quaest. Ent., 1985, 21 (2)

REL 2 (EL/HW)

222

Ward

0.85 p
0.80 -
0.75 -
0.70 -
0.65 -

elongatus
O Florida

o ° □

□ Texas, Central America

O 08 4 o o „ o

A Jamaica

—

cubaensis

• Florida

♦ Cuba, Haiti,
A Jamaica

, Bahamas

° Yo o 00 °

DD® □ g,D ^

A D ™

•

□

• • _

♦

jt'j

•

♦ ♦

I l l l l l

0.65 0.70 0.75 0.80 0.85 0.90 0.95

PLI (PH/PL)

Figure 33. Plot of relative eye length (REL2) and petiole length index (PLI) in workers of Pseudomyrmex cubaensis and
P. elongatus.

Nearctic Species of the genus Pseudomyrmex

223

Figures 34-39. Pseudomyrmex queens. 34, 36, dorsal views of head; 35, 37, lateral views of mesosoma (part), petiole,
postpetiole, and gaster (part); 38-39, close-up views of head sculpture, from insets. 34,35,38, P. leptosus (holotype,
Florida); 36,37,39, P. pallidus (Florida). Scale lines = 0.5 mm.

224

Ward

Figures 40-45. Pseudomyrmex workers. 40, 42, 44, dorsal views of head; 41, 43, 45, lateral views of mesosoma (part),
petiole, postpetiole, and gaster (part). 40,41, P. simplex (Florida); 42,43, P. pallidus (Florida); 44,45, P. seminole
(holotype, Florida). Scale lines = 0.5 mm.

Nearctic Species of the genus Pseudomyrmex

225

SPECIES ACCOUNTS

gracilis group

Pseudomyrmex mexicanus Roger
(Figs. 11,12)

Pseudomyrma mexicana Roger, 1863, p. 178. Syntype worker(s), Mexico (not in MNHU) [Not examined].

Pseudomyrma gracilis var. mexicana Roger; Wheeler, 1901, p. 204.

Pseudomyrma gracilis var. mexicana Roger; Wheeler, 1908, p. 421.

Pseudomyrma gracilis mexicana Roger; Mitchell & Pierce, 1912, p. 69.

Pseudomyrma gracilis var. mexicana Roger; Wheeler & Bailey, 1920, pp. 259, 262. [Description of larva, and contents of
food pellets] .

Pseudomyrma gracilis subsp. mexicana Roger; Wheeler, 1942, pp. 166,167.

Pseudomyrma gracilis mexicana Roger; Creighton, 1950, p.80.

Pseudomyrmex mexicanus Roger; Whitcomb et al., 1972, pp. 31-33.

Worker Measurements ( n = 13): HL 1.50-1.72, HW 1.53-1.70, MFC 0.036-0.051, Cl
0.98-1.02, OI 0.49-0.54, REL 0.54-0.60, REL2 0.54-0.60, OOI (-0.19)-( + 0.39), VI
0.71-0.81, FCI 0.023-0.033, SI 0.47-0.51, SI2 0.80-0.92, FI 0.37-0.41, PDI 1.03-1.32,
MPI 0.058-0.073, NI 0.62-0.71, PLI 0.46-0.51, PWI 0.41-0.47, PPWI 0.94-1.12.

Worker Diagnosis. — Immediately distinguishable from all other Nearctic Pseudomyrmex
by its large size (worker H W > 1 .40) and bicolored, orange and black markings. Head broad
(Cl 1.00), frontal carinae moderately well separated (MFC 0.04), eyes large (REL
0.58); occipital margin convex to flat, in full-face, dorsal view; pronotum laterally margined;
metanotal groove distinct; basal face of propodeum rounding into declivitous face, and not
distinctly differentiated from it; petiole and postpetiole elongate, the former with a
well-developed anterior peduncle. Integument mostly subopaque, due to fine coriarious or
punctulate sculpture. Appressed and erect hairs common on most parts of the body, including
mesonotum, propodeum, legs, and scapes. Gaster and most of head black; mesosoma ( =
alitrunk), petiole, and postpetiole orange, with varying amounts of black infuscation
(commonly the pronotum is orange, while the mesonotum and propodeum are dark).

Comments. — I have been unable to locate type material of P. mexicanus (not present in
MNHU, according to F. Koch, in litt.). Application of this name to the Nearctic representative
of the gracilis group is based upon the original description and the type locality. P. mexicanus
belongs to a complex of closely related forms of uncertain taxonomic status, distinguished
mainly on the basis of color (the so-called gracilis complex, within the Pseudomyrmex gracilis
group). These taxa are often listed as subspecies of gracilis , despite the fact that some forms
are broadly sympatric. P. mexicanus is similar to the Central American taxon, bicolor Guerin;
the latter is ostensibly darker on average, with a more slender petiole. However, the members of
the gracilis complex exhibit notable variation in color patterns and in the shape of the petiole.
A thorough systematic analysis is needed to disentangle the intra- and inter-specific
components of this variation. The occurrence of modal color patterns and transitional forms
suggests that some of the taxa are incompletely isolated (semispecies).

Biology. — P. mexicanus nests in dead or cavity-ridden branches in a wide variety of trees,
shrubs, and herbs. Museum records include nests from the following plants: Baccharis ,
Cladium , Peperomia , Prosopis, Rhizophora , and Salix. Apparently introduced into Florida in
recent times (first collected in 1960), P. mexicanus is now common in the southern half of the
state, where it occurs in hardwood hammocks, mangrove, and old field second growth habitats.
Whitcomb et al. (1972) provide notes on nesting and feeding behavior in Florida.

Quaest. Ent., 1985, 21 (2)

226

Ward

Material Examined (GCW, LACM, MCZ, PSW, UCD, USNM).—

FLORIDA: Collier Co.: Collier-Seminole St. Pk. (P. S. Ward); Dade Co.: Hialeah (C. Stegmaier); Homestead A.F.B.
(G. C. & J. Wheeler); Long Pine Key (G. C. & J. Wheeler); Mahogany Hammock, Everglades Natl. Pk. (R. Wagner; G.
C. & J. Wheeler); Old Flamingo Rd., Everglades Natl. Pk., 10 m (P. S. Ward); Indian River Co.: Vero Beach (L. & C.
W. O’Brien); Monroe Co.: Bear Lake Trail, near Flamingo, Everglades Natl. Pk. (G. C. & J. Wheeler); John Pennekamp
State Pk. < 5 m (P. S. Ward).

TEXAS: Aransas Co.: Goose I. St. Pk., 5 m (P. S. Ward); Brazoria Co.: 4 mi SW West Columbia (P. S. Ward);
Cameron Co.: 5 mi W Boca Chica (G. C. & J. Wheeler); 10 mi W Boca Chica (W. S. Creighton); Brownsville
(Darlington; Jones & Pratt; D. J. & J. N. Knull; Lattimore & Bottimer; McMillan; C. H. T. Townsend); Harlingen (W.
Buren); Laguna Madre, 25 mi SE Harlingen (D. E. Hardy); no specific locality (Dreyer; D. J. & J. N. Knull); Hidalgo
Co.: Bentsen Rio Grande St. Pk. (E. E. Grissell & A. S. Menke; P. S. Ward); Mission (P. C. Avery); Pharr; Santa Ana
Refuge (P. S. Ward); no specific locality (D. J. & J. N. Knull); Kenedy Co.: 27°10'N, 97°40'W (J. E. Gillaspy); Kleberg
Co.: Kingsville (J. E. Gillaspy); Live Oak Co.: 4 mi S George West (R. Snelling); Nueces Co.: Corpus Christi (R. A.
Cushman; Jones & Pratt); Victoria Co.: Victoria (J. D. Mitchell).

Other material, tentatively identified as P. mexicanus , from Mexico, Guatemala,
Nicaragua, Costa Rica, Panama and Jamaica.

elongatus group

Pseudomyrmex cubaensis Forel stat. nov.

(Figs. 5, 6, 25, 32)

Pseudomyrma elongata var. cubaensis Forel, 1901, p. 342. Holotype (unique syntype) worker, Bahia Honda, Cuba
(MHN) [Examined].

Pseudomyrma elongata ; Wheeler (nec Mayr), 1905, pp. 85-87 (partim).

Pseudomyrma elongata var. cubaensis Forel; Forel, 1913, p. 215 [Description of queen].

Pseudomyrma elongata var. cubaensis Forel; Wheeler, 1913a, pp. 484-485.

Pseudomyrma elongata var. cubaensis Forel; Wheeler & Mann, 1914, p. 18.

Pseudomyrma elongata var. cubaensis Forel; Mann, 1920, p. 405

Pseudomyrma elongata; Wheeler & Bailey (nec Mayr), 1920, pp. 260, 265 [Description of larva, and contents of food
pellets].

Pseudomyrma elongata ; Wheeler (nec Mayr), 1932, p. 4 (partim).

Pseudomyrma elongata ; Wheeler (nec Mayr), 1942, p. 165.

Pseudomyrma elongata ; Creighton (nec Mayr), 1950, pp. 79-80 (partim).

Pseudomyrmex elongata ; Creighton (nec Mayr), 1955, pp. 17-20 (partim).

Pseudomyrmex elongatus ; Wheeler & Wheeler (nec Mayr), 1956, p. 384 [Description of larva].

Worker Measurement s{n = 21, except for HL, HW, Cl, REL, REL2, and PLI, where n =
52): HL 0.84-1.05, HW 0.64-0.75, MFC 0.017-0.029, Cl 0.69-0.77, OI 0.52-0.58, REL
0.47-0.51, REL2 0.63-0.73, OOI 0.41-0.95, VI 0.74-0.83, FCI 0.024-0.044, SI 0.44-0.48,
SI2 0.64-0.74, FI 0.42-0.48, PDI 1.09-1.44, MPI 0.038-0.084, NI 0.53-0.63, PLI
0.65-0.78, PWI 0.55-0.69, PPWI 1.01-1.25.

Worker Diagnosis. — A small, brown species with elongate head (HW 0.64-0.75, Cl

O. 69-0.77) and with erect pilosity on the mesonotum and propodeum. Very similar to P.
elongatus Mayr (q.v.), except averaging larger, with relatively short eyes (REL2 0.63-0.73)
and a longer petiole and postpetiole (PLI 0.65-0.78). Head punctate, the punctures maximally
separated by about their diameters.

Comments. — Originally described as a variety of P. elongatus, P. cubaensis was
synonymized with the former by Creighton (1955, p. 18). However it appears to be consistently
distinct from the smaller elongatus- like form with which occurs sympatrically in south Florida.
The most important differences are in the relative length of the eye and the shape of the petiole.
A two-dimensional plot of REL2 and PLI cleanly separates all Floridian and most other
material into two taxa (Figure 33). In Jamaica the two forms are less distinct. It is possible that

P. elongatus and P. cubaensis represent a remnant circular Rassenkreis stretching around the

Nearctic Species of the genus Pseudomyrmex

227

Gulf of Mexico, with intermediate populations in Jamaica.

Apart from the differences in eye length and petiole shape, P. cubaensis also tends to have a
broader head, narrower forefemur (FI 0.42-0.48), longer postpetiole (PPWI 1.01-1.25), and
fewer (but longer) erect setae on the petiole, postpetiole, and fourth abdominal tergite. The
body sculpture and appressed pubescence is lighter than in Florida P. elongatus, producing a
shinier appearance, particularly on the occiput, propleuron, petiole, and postpetiole. (Elsewhere
P. elongatus may have an equally shiny integument, e.g. in Texas.)

Differences between queens and males of the two species are given in the respective keys.
Biology. — In Florida, I have collected P. cubaensis in dead twigs of Rhizophora mangle
and Conocarpus erectus. There are museum records of nests in Tillandsia (Florida) and
Cladium (Bahamas), and of workers foraging on Ficus aurea , mangrove, sea grape, and acacia.
Wheeler’s (1905) records of Bahamaian “ elongatus ” in culms of Uniola and Cladium and in
hollow twigs of gum mastic, sea grape, and buttonwood, refer in part to P. cubaensis (see also
discussion of Pseudomyrmex subater Wheeler & Mann under P. elongatus ).

Material Examined (LACM, MCZ, PSW, UCD, USNM).—

FLORIDA: Collier Co.: Collier-Seminole St. Pk. (P. S. Ward); Marco (W. T. Davis); Dade Co.: Biscayne Bay
(Slosson); Cards Point (W. M. Wheeler); Long Pine Key (W. M. Wheeler); Miami Beach (W. E. Brown; A. C. Cole);
Paradise Key (D. Fairchild; W. M. Wheeler); no specific locality (J. N. Knull); Highlands Co.: Archbold Biol. Stn. (R.
Silberglied); Highlands Hammock State Park (L. & C. W. Obrien); near Sebring (R. W. Klein); no specific locality (F. J.
Moore); Hillsborough Co.: no specific locality (J. C. Bowyer); Lake Co.: no specific locality (W. A. Hiers); Lee Co.: Ft.
Meyers [ = Ft. Myers]; Monroe Co.: Lower Matecumbe Key (W. M. Wheeler); N. Key Largo (R. W. Klein); Key West;
No Name Key (P. S. Ward); Osceola Co.: Lake Alfred (M. H. Muma); Palm Beach Co.: Boynton Beach (Wood &
Davidson); Sarasota Co.: Long Branch Key (A. C. Cole); Sarasota (A. C. Cole); 30 mi SE Sarasota (J. Longino).

BAHAMAS: Andros Island (W. M. Wheeler); Mangrove Cay, Andros Island (B. Cole); Conception Island (G.
Greenway); Gun Point, Crooked Island (B. Valentine & R. Hamilton); New Providence (B. Cole).

CUBA: Aguada de Pasajeros (W. M. Wheeler); Anafe, Havana (G. Aguayo); Carnoa, Havana (G. Aguayo);
Cayamas (Baker; E. A. Schwartz); Cienaga de Japata (W. M. Wheeler); Guanajay, Pinar del Rio (E. O. Wilson);
Guavivo Cave, Soledad (F. Smith); Jiquari (Barbour & Shaw); La Milpa, near Cienfuegos (G. Salt); Pinares Oriente (W.
M. Mann); Santa Clara, Las Villas Prov. (E. O. Wilson); Soledad, Cienfuegos (C. T. & B. B. Brues; W. S. Creighton; J.
G. Myers; F. Smith; N. A. Weber).

HAITI: Grande Riviere (W. M. Mann); Mtns. N. of Jacmel (W. M. Mann).

JAMAICA: Troy (Wight); Balaclava (Wight).

Pseudomyrmex elongatus Mayr
(Figs. 7,8, 24,31)

Pseudomyrma elongata Mayr, 1870, p. 413. Syntype worker(s), Colombia, (Lindig) (not in NHMV) [not examined],
Pseudomyrma elongata var. tandem Forel, 1906, p. 228. Syntype workers, El Hiquito, near San Mateo, Costa Rica (P.

Biolley) (MNHN) [Examined] [Synonymy by Creighton, 1955, p. 18].

Pseudomyrma elongata Mayr; Wheeler, 1932, p. 4 (partim).

Pseudomyrma elongata Mayr; Creighton, 1950, pp. 79-80 (partim).

Pseudomyrmex elongata Mayr; Creighton, 1955, pp. 17-20 (partim).

Pseudomyrmex elongatus Mayr; Wilson, 1964, p. 4.

Worker Measurements (n = 24, except for HL, HW, Cl, REL, REL2, and PLI, where n =
50): HL 0.78-0.91, HW 0.56-0.64, MFC 0.013-0.024, Cl 0.68-0.74, OI 0.52-0.58, REL
0.50-0.58, REL2 0.73-0.82, OOI 0.13-0.48, VI 0.82-0.90, FCI 0.021-0.041, SI 0.42-0.48,
SI2 0.55-0.64, FI 0.45-0.56, PDI 0.95-1.45, MPI 0.036-0.072, NI 0.55-0.63, PLI
0.76-0.91, PWI 0.62-0.74, PPWI 1.09-1.40.

Worker Diagnosis. — A small, brown species with elongate head and eyes (HW 0.56-0.64,
Cl 0.68-0.74); frontal carinae subcontiguous; occipital margin flat or slightly concave, in
full-face dorsal view; basal and declivitous faces of propodeum well differentiated; petiole
short, broad, and rounded (PLI 0.76-0.91); postpetiole wider than long. Head densely punctate

Quaest. Ent., 1985, 21 (2)

228

Ward

and usually more or less opaque; remainder of body finely punctate or coriarious-imbricate,
varying from opaque to sublucid. Erect pilosity and fine appressed pubescence present on most
parts of body, including mesonotum and propodeum; fourth abdominal tergite with a rather
dense mat of appressed pubescence.

Comments. — This is the smaller of two elongatus- like species in North America. I am
considering it conspecific with P. elongatus Mayr on the basis of (i) the original description of
P. elongatus, particularly the indication that HL is 1.5 times HW, and (ii) worker material
from Costa Rica (leg. Biolley) (MNHN, NHMV) determined as P. elongatus by Mayr and
Forel. There is a confusing variety of elongatus-like forms in Central and South America,
which require detailed taxonomic study. Until such a study is carried out, it seems expedient to
refer to the North American species as P. elongatus and to leave P. tandem Forel as a
provisional synonym.

P. subater Wheeler & Mann (1914), originally described as a subspecies of P. elongatus,
was recognized as a distinct species by Creighton (1955). It may be distinguished from P.
elongatus and P. cubaensis by the shinier integument, conspicuous pilosity (grading insensibly
from appressed pubescence to fine suberect and erect setae), broad head (Cl 0.83-0.88), short
eyes (REL2 0.54-0.58 in P. subater, > 0.62 in P. elongatus and P. cubaensis ), distinct petiolar
shape (gradually inclined anterior face rounded into a sharply declining posterior face so that
NI 0.61-0.72), and conspicuous anteroventral tooth on the postpetiole. Apart from two “cotype”
workers in the MCZ from Haiti, I have seen material of P. subater (misidentified as P.
elongatus ) from the Bahamas (Andros Island, Nassau) and the same, or a closely related
species, from Jamaica (Kingston)). Wheeler’s (1905) record of “ elongatus ” from the Bahamas
appears to be based on a combination of P. subater and P. cubaensis, judging from material in
the MCZ.

Recent collections of P. subater from the Bahamas by Blaine Cole show that this species has
striking bright orange queens, which look superficially like those of P. pallidus. Cole also made
a collection from a single Cladium culm which contained both P. subater and P. cubaensis
workers. These findings suggest that Wheeler’s (1905) and Mann’s (1920) records of dulotic
associations between “ flavidula ” and “ elongata ” may have been based in part on pure colonies
of P. subater, or mixed colonies of P. subater and P. cubaensis.

Biology. — P. elongatus nests in dead twigs in a variety of woody shrubs and trees. I have
collected P. elongatus colonies in twigs of Avicennia germinans, Baccharis halimifolia,
Laguncularia racemosa and Rhizophora mangle in Florida, and in Gliricidia sepium,
Helicteres, and Inga in Costa Rica and Panama. Among museum material there are records of
P. elongatus nesting in a “climbing vine” and “mangrove stems” in Florida, in Quercus
virginiana and Prosopis (Texas), and in Quercus fusiformis (Nuevo Leon, Mexico).

Material Examined (GCW, LACM, MCZ, PSW, UCD, USNM).—

FLORIDA: Collier Co:. Everglade[s] (W. T. Davis); Dade Co:. Coconut Grove; Miami; Rattlesnake Hammock,
Homestead (R. Gregg); Shark Valley, Everglades Natl. Pk. (P. S. Ward); no specific locality (J. N. Knull); Highlands
Co: Archbold Biol. Stn., Lake Placid (T. C. Schneirla); Lee Co: Ft. Myers (W. M. Barrows); Monroe Co: Big Pine Key
(E. O. Wilson); John Pennekamp St. Pk., < 5 m (P. S. Ward); Key Largo; Key West (E. O. Wilson); N. Key Largo (R.
W. Klein); Plantation Key (E. O. Wilson).

TEXAS: Cameron Co: 5 mi W Boca Chica (G. C. & J. Wheeler); 10 mi W Boca Chica (R. R. Snelling); Harlingen
(W. Buren); Hidalgo Co: Mission (W. Buren); Monte Alto (W. S. Creighton); Kenedy Co: 26 mi N Raymondsville (W.
S. Creighton).

MEXICO: Nayarir. Maria Magdalena, Is. Tres Marias (R. R. Snelling); Nuevo Leon : El Pastor, Montemorelos, 2000
ft (W. S. Creighton); San Luis Potosi: Rio Amahac, Tamazunchale, 300 ft (W. S. Creighton); 3 mi N. Valles (W. S.
Creighton); Sinaloa: Mazatlan (P. J. Spangler); Tamaulipas: Canon de el Abra, 1000 ft (W. S. Creighton).

COSTA RICA: Guanacaste Prov: 1 km SW Pto. Coyote, < 5 m (P. S. Ward); Puntarenas Prov: Llorona, Corcovado
Natl. Park, 10 m (P. S. Ward); Manuel Antonio Natl. Pk., 5 m (P. S. Ward); Monteverde, 1200 m (P. S. Ward); Sirena,

Nearctic Species of the genus Pseudomyrmex

229

Penin. Osa, 50 m (J. Longino).

JAMAICA: Ford 1 mi SE Stony Hill (E. A. Chapin).

PANAMA: 2 km W Gamboa, Canal Zone, 30 m (P. S. Ward); 6 km NW Gamboa, Canal Zone, 50 m (P. S. Ward); 6
km NW Gatun Dam, Canal Zone, 75 m (P. S. Ward).

pallidus group

Pseudomyrmex apache Creighton
(Figs. 9, 10, 17)

Pseudomyrmex apache Creighton, 1952, p. 134. Nidoparatype workers, females, males. Brown Canyon, Baboquivari
Mtns., Arizona, 4400 ft, 2.ix. 1 95 1 , in Quercus oblongifolia 502 (W. S. Creighton) (LACM, MCZ) [Examined],
Pseudomyrma pallida', Wheeler (nec F. Smith), 1908, p. 420 (partim).

Pseudomyrmex apache Creighton; Creighton, 1954, pp. 9-15 [Distribution],

Pseudomyrmex apache Creighton; Wheeler & Wheeler, 1956, p. 380 [Description of larva].

Pseudomyrmex apache Creighton; Creighton, 1963, pp. 1-4 [Biology].

Pseudomyrmex apache Creighton; Wheeler & Wheeler, 1973, pp. 41-42.

Worker Measurements (n = 35): HL 1.02-1.30, HW 0.83-1.04, MFC 0.033-0.066, Cl
0.75-0.84, OI 0.58-0.65, REL 0.39-0.44, REL2 0.48-0.54, PPI 1.10-2.00, VI 0.74-0.84,
FCI 0.034-0.070, SI 0.46-0.51, SI2 0.90-1.00 FI 0.38-0.44, PDI 0.87-1.19, MPI
0.007-0.044, NI 0.51-0.64, PLI 0.54-0.61, PWI 0.48-0.56, PPWI 1.02-1.19.

Worker Diagnosis. — Head broad; eyes short (EL subequal to SL); anterior clypeal margin
laterally rounded; frontal carinae subcontiguous, MFC subequal to the basal width of scape;
occipital margin flat to broadly convex, in full-face, dorsal view; pronotum with weak lateral
margination; metanotal groove usually weak; petiole relatively short, broad, and high with a
rather sharply inclined anterior face. Head opaque to sublucid, densely punctulate on a
coriarious background; mesosoma and petiole subopaque, coriarious-imbricate; postpetiole and
gaster opaque to sublucid, covered with numerous, fine piligerous punctures. Erect setae
sparsely present on scape, head, pronotum, petiole, postpetiole, gaster, and legs (generally
absent on mesonotum and propodeum); typically four pairs of erect setae on dorsum of head,
and two or three pairs each on pronotum, petiole, and postpetiole. Fine, appressed pubescence
scattered over body, including fourth abdominal tergite. Rich orange-brown, the head (and
sometimes legs and gaster) a little darker.

Comments. — Although I have placed P. apache in the pallidus group as a matter of
convenience, it is a rather distinct species showing only superficial resemblance to other
members of the group. It is the only species to possess such well separated frontal carinae,
laterally rounded anterior clypeal margin, short eyes relative to scape length, and (in the male)
ventrally pointed pygidium. P. apache workers also tend to be larger, more densely sculptured
(hence less shiny), and more setose than those of other pallidus group species. Size alone
(worker HW > 0.83) will separate P. apache from all species except P. pallidus and P.
seminole. Apart from character differences outlined in the keys (of which eye size relative to
scape length and shape of male terminalia are most distinctive), P. apache can usually be
distinguished from P. pallidus and P. seminole by the presence of a pair of erect setae, one on
either side of the median ocellus, in the worker. In P. apache workers these two setae are
always present and usually as long as the ocellar distance (OD). In P. seminole and P. pallidus
workers these setae are either absent or shorter than OD.

Biology. — A denizen of xeric habitats, P. apache nests in sizable dead branches (1-12 cm
diameter) of various trees (especially live oaks) and large woody shrubs, usually taking
advantage of beetle-bored cavities. By state and country, nest-site records are as follows:

Quaest. Ent., 1985, 21 (2)

230

Ward

Texas: Prosopis glandulosa , Quercus grisea.

Arizona: Populus sp., Prosopis sp., Quercus arizonica , Q. emoryi, Q. grisea, Q.
oblongifolia, Q. turbinella.

California: Arctostaphylos manzanita , Fraxinus gall, Pinus attenuata cone, Quercus
chrysolepis, Q. mslizenii, Umbellularia californica.

Mexico: Prosopis sp., Quercus emoryi , Q. fusiformis, Q. oblongifolia, Q. santaclarensis.

Of 13 nests which I have dissected (from Texas, Arizona and California), five contained no
dealate females, six contained a single queen, one contained two functional (i.e. inseminated)
queens, and one contained 6 dealate queens. Thus this species is at least occasionally
polygynous and (judging from the queenless nests) polydomous. For two of the five queenless
nests, queenright nests were located on the same tree or shrub.

I have seen two instances of lone foraging (presumably colony founding) dealate queens: one
on the trunk of a Quercus arizonica tree in September (Arizona) and the other on an
Arctostaphylos bush in February (northern California). The latter queen was dissected and
found to be inseminated but possessing preoviposition ovaries (ovarioles short; corpora lutea
absent). Alates of P. apache have been collected in March, April, and July to November,
suggesting that mating may occur in more than one season.

Material Examined (BMNH, GCW, LACM, MCZ, PSW, UCD, USNM).—

ARIZONA: Cochise Co.: Carr Canyon, Huachuca Mtns., 5400 ft (W. S. Creighton), 6200 ft (C. W. O’Brien); Cave
Crk. Ranch, Chiricahua Mtns., 5000 ft (G. E. Wallace); Chiricahua Mt. (D. J. and J. N. Knull); Chiricahua Mtns. (J. N.
Knull); Chiricahua Natl. Monum. Cpgrd., 5400 ft (W. S. Creighton); Cochise Stronghold, Dragoon Mtns., 5200 ft (W. S.
Creighton); Coronado Peak, 2020 m (P. S. Ward); Garden Canyon, Huachuca Mtns., 5800 ft (W. S. Creighton);
Huachuca Mt. (J. N. Knull); Miller Canyon, Huachuca Mtns. (W. S. Creighton); Portal (G. Alpert); 3 km SW Portal,
1510 m (P. S. Ward); 7 km SE Sunnyside, 1670 m (P. S. Ward); Gila Co.: Globe (Nuttig); Graham Co.: Cottonwood
Canyon, Peloncillo Mtns., 4800 ft (W. S. Creighton); Graham Mtns., 3500-4500 ft (R. M. Bohart); Post Canyon, Pinaleno
Mtns., 5000-6000 ft (W. M. Wheeler); Mohave Co.: Hualapai Mtns., S. of Kingman, 1450 m (E. Schlinger); Pima Co.:
Abra Wash, Growler Mtns., Organpipe Cactus Natl. Monum., 1300 ft (W. S. Creighton); Alamo Canyon, Ajo Mtns.,
Organpipe Cactus Natl. Monum., 2200 ft (W. S. Creighton); Brown Canyon, Baboquivari Mtns., 4400 ft (W. S.
Creighton); Forestry Cabin, Baboquivari Mtns., 3500 ft (W. S. Creighton); Organpipe Cactus Natl. Monum. (E. R.
Tinkham); Sabino Canyon (V. L. Vesterby); San Miguel (E. D. Algert); Tucson (J. Knull); Santa Cruz Co.: Canelo Pass,
5300 ft (W. S. Creighton); Madera Canyon, Santa Rita Mtns. (W. S. Creighton), 4880 ft (C. R. Kovacic; V. L. Vesterby);
Nogales (Burdine; D. J. & J. N. Knull); Pena Blanca Springs, 3700 ft (W. S. Creighton); Sweetwater, Santa Rita Mtns.,
4000 ft and 6000 ft (W. S. Creighton); Tumacacori Mt. (D. J. & J. N. Knull); county unknown: Catal Springs (Hubbard
& Schwartz); Santa Catalina Mtns. (M. Chrisman); Santa Rita Mtns. (R. M. Bohart; J. Knull).

CALIFORNIA: Butte Co.: 6 km N Feather Falls, 600 m (P. S. Ward); Colusa Co.: 1 km W Fout Springs, 600 m (P.
S. Ward); Imperial Co.: Winterhaven (R. L. Westcott); Lake Co.: Borax Lake (E. L. Westcott); Los Angeles Co.:
Altadena; Eaton Canyon Pk. (M. E. Thompson); Foothill, Pasadena (A. H. Sturtevant); 3 mi N. Mt. Baldy (E. Weidert);
Tanbark Flat (R. C. Bechtell; R. M. Bohart); Napa Co.: Mt. St. Helena (J. S. Buckett); 4 km E summit Mt. St. Helena,
450 m (P. S. Ward); Orange Co.: Irvine Pk. (K. Brown); Trabuco Canyon (M. E. Irwin); Trabuco east (E. Eidert);
Riverside Co.: Blythe (R. M. Hardman); Deep Canyon (W. P. MacKay); Dripping Springs, Agua Tibia Mtns., 1500 ft
(W. S. Creighton); Pinon Flat, San Jacinto Mtns. (R. L. Macdonald); Poppet Flats (G. Clark); Riverside (E. I. Schlinger);
Whitewater (A. L. Melander); Winchester (W. Icenogle); San Bernardino Co.: nr. Cajon pass; San Diego Co.: Jacumba
(D. J. & J. N. Knull); 5.9 mi NE Ramona, Hwy. 78 (S. & S. ;Fromer, S. Larisch); 5.2 mi NW Ramona, Hwy. 78 (S. & S.
Frommer, S. Larisch); no specific locality; Santa Barbara Co.: Canyon del Medio, Santa Cruz I. (R. O. Schuster & E. C.
Toftner); Santa Clara Co.: no specific locality; Solano Co.: Cold Canyon, 420 m (P. S. Ward); Tehama Co.: 26 km WSW
Red Bluff, 240 m (P. S. Ward); Tulare Co.: Ash Mtn. Powerhouse #3 (D. J. Burdick; J. A. Halstead); Ash Mtn., Sequoia
Natl. Pk. (M. G. Fitton); Horse Creek Rd. (O. L. Brawner); Ventura Co.: Saticoy (R. E. Barrett); Yolo Co.: 3 km SW
Guinda, 150 m (P. S. Ward); 4 km NW Rumsey, 150 m (P. S. Ward); county unknown: mtns. near Claremont [Los
Angeles or San Bernardino Co.] (Baker).

DISTRICT OF COLUMBIA: Washington (B. P. Currie) [Dubious locality record].

TEXAS: Bexar Co.: San Antonio (P. S. Ward); Duval Co.: Freer (R. R. Rodgers); San Diego; Edwards Co.: Camp
Wood (C. R. Ward); Goliad Co.: no specific localilty (J. D. Mitchell); Hidalgo Co.: Monte Alto, 60 ft (W. S. Creighton);
La Salle Co.: Fowlerton, 300 ft (W. S. Creighton); Maverick Co.(?): El Indigo [= El Indio?] (D. H. Bixby); Presidio Co.:
Arsaca Canyon, Chinati Mtns., 4800 ft (W. S. Creighton); Starr Co.: no specific locality (D. J. & J. N. Knull); Travis
Co.: Austin (P. S. Ward); Uvalde Co.: no specific locality (D. J. & J. N. Knull).

Nearctic Species of the genus Pseudomyrmex

231

MEXICO: Baja California Norte : 25 mi N El Arco (W. H. Ewart); Baja California Sur. 72 mi NW La Paz, 100 ft
(R. R. Snelling); San Jose del Cabo; 7 mi NW Santa Rosalia, 850 ft (R. R. Snelling); 2.7 mi SE Valle Peridido;
Chihuahua: 3 mi S Encinillas, 4900 ft (W. S. Creighton); 16 mi W Gral. Trias, 5800 ft (W. S. Creighton); Nogales
Ranch, Sierra de en Medio, 5200 ft (W. S. Creighton); Ojo del Cerro Chilicote (C. H. T. Townsend); 23 mi S Parral, 5500
ft (W. S. Creighton); 34 mi S Parral, 5800 ft (W. S. Creighton); Durango: Villa Ocambo [ = Ocampo], 5700 ft (W. S.
Creighton); Nuevo Leon: China, 600 ft (W. S. Creighton); Sonora: 30 mi SE Agua Prieta (V. Roth); 4.8 mi S Cananea
(V. Roth).

Pseudomyrmex brunneus F. Smith
(Figs. 13, 15)

Pseudomyrma brunnea F. Smith (1877), p. 63. Holotype (unique syntype) worker, Mexico (BMNH) [Examined],
Pseudomyrma brunnea var. nigrita Enzmann, 1945, p. 82. Syntype workers, Mirador, Mexico (E. Skwarra) (MCZ)
[Examined]. Syn. nov.

Worker Measurements ( n = 10): HL 0.77-0.86, HW 0.67-0.72, MFC 0.012-0.024, Cl
0.83-0.90, OI 0.57-0.63, REL 0.51-0.56, REL2 0.58-0.64, OOI 0.67-1.20, VI 0.70-0.78,
FCI 0.017-0.035, SI 0.43-0.50, SI2 0.69-0.80, FI 0.41-0.45, PDI 0.56-0.75, MPI
0.052-0.075, NI 0.57-0.65, PLI 0.47-0.58, PWI 0.54-0.65, PPWI 1.26-1.54.

Worker Diagnosis. — Small, brown species, with wide head and broadly rounded occiptal
margin (HW 0.67-0.72, Cl 0.83-0.90). Similar to P. ejectus F. Smith {q.v.) except as follows:
basal face of propodeum between one half and three quarters the length of declivitous face
(PDI 0.56-0.75); petiole shorter and broader than that of P. ejectus (PWI 0.54-0.65); petiolar
node somewhat displaced posteriorly; postpetiole short and wide (PPWI 1.26-1.54). Front of
head opaque, densely coriarious-imbricate; sculpture becoming weaker towards the vertex, with
scattered, fine punctures on a (usually) sublucid, coriarious background; mesosoma subopaque,
coriarious to coriarious-imbricate; petiole, postpetiole, and gaster increasingly (in that order)
less coriarious and more smooth and shining. Erect pilosity very sparse; appressed pubescence
scattered over body; abdominal tergite IV with appressed hairs separated by about their
lengths, and not obscuring the shiny integument. Body dark brown, little or no contrast in color
between the head, mesosoma, petiole, postpetiole, or gaster; mandibles and apical ends of tarsi
luteous.

Comments. — This appears to be an uncommon Mexican species, which occurs
sympatrically with P. ejectus. The major differences between the two species are in the shapes
of the propodeum, petiole, and postpetiole (note especially the non-overlapping values of PWI
and PPWI). In addition, P. brunneus tends to have a more densely sculptured (and opaque)
head than P. ejectus and to exhibit less contrasting light and dark brown coloration on the
mesosoma, petiole and postpetiole.

Biology. — At Cola de Caballo, near Monterrey, I collected workers and larvae of P.
brunneus in dead twigs of a small tree, probably Melia azedarach, and in the dead stalk of an
unidentified mint. The latter nest contained a single dealate queen. P. ejectus was also found
nesting in a dead mint stalk at this locality.

Material Examined (BMNH, MCZ, PSW, UCD).—

MEXICO: Nuevo Leon: Cola de Caballo, 38 km SSE Monterrey, 600 m (P. S. Ward); Veracruz: Mirador (E.
Skwarra); state unknown: “Mexico.”.

Pseudomyrmex ejectus F. Smith
(Figs. 14, 16, 23,30)

Pseudomyrma ejecta F. Smith (1858), p. 157. Two syntype workers, “ Brazil?” (BMNH) [Examined]. One syntype here
designated as LECTOTYPE.

Quaest. Ent., 1985, 21 (2)

232

Ward

Pseudomyrma brunnea ; Wheeler (nec F. Smith), 1908, pp. 420-421.

Pseudomyrma brunnea ; Mitchell & Pierce (nec F. Smith), 1912, p. 69.

Pseudomyrma brunnea ; Wheeler (nec F. Smith), 1913b, p. 240.

Pseudomyrma brunnea ; Wheeler (nec F. Smith), 1932, p. 3.

Pseudomyrma brunnea-, Creighton (nec F. Smith), 1950, p. 79.

Pseudomyrmex brunneus ; Wheeler & Wheeler (nec F. Smith), 1956, p. 382 [Description of larva].

Worker Measurements (n = 50): HL 0.65-0.97, HW 0.56-0.81, MFC 0.007-0.021, Cl
0.78-0.89, 01 0.52-0.62, REL 0.51-0.59, REL2 0.61-0.70, OOI 0.46-1.52, VI 0.64-0.78,
FCI 0.010-0.030, SI 0.43-0.49, SI2 0.64-0.76, FI 0.36-0.50, PDI 0.70-1.07, MPI
0.046-0.097, NI 0.48-0.60, PLI 0.43-0.57, PWI 0.40-0.52, PPWI 0.93-1.25.

Worker Diagnosis. — Head wide, broadly rounded (HW 0.56-0.81, Cl 0.78-0.89); anterior
clypeal margin medially straight, laterally angulate; frontal carinae closely contiguous;
occipital margin convex, flat, or slightly concave, in full face, dorsal view; pronotum broadly
rounded; metanotal groove wide and rather deep (MPI 0.046-0.097); basal face of propodeum
subequal in length to declivitous face, and poorly differentiated from it (PDI 0.70-1.07);
petiole and postpetiole relatively long and slender (PWI 0.40-0.52); anteroventral tooth present
on petiole. Head subopaque to weakly shining, the frons punctulate on a coriarious imbricate
background; sculpture weakening towards the vertex, which is correspondingly more shiny;
mesosoma and petiole sublucid, coriarious-imbricate; postpetiole and gaster more or less
smooth and shining. Erect pilosity very sparse; appressed pubescence inconspicuous; appressed
hairs on abdominal tergite IV not forming a dense mat nor obscuring the shiny integument.
Head (except clypeus and mandibles), mesonotum, propodeum, and gaster dark brown;
pronotum, petiole, and postpetiole generally a lighter brown, of varying contrast; clypeus,
mandibles, and apices of legs light brown to pale luteous.

Comments. — The lectotype and paralectotype workers of P. ejectus in the BMNH agree
well with the common species in southeastern United States which has been masquerading
under the name “brunneus” . I have also seen material which I would consider conspecific with
P. ejectus from Mexico, Jamaica, Belize, and Costa Rica, but not from South America. (There
are other ejectus- like taxa in Central and South America, some undescribed.) It seems likely
that the types of ejectus came from the United States or Central America rather than Brazil.
Differences between P. ejectus and P. brunneus are discussed under the latter species.

Described as a variety P. ejectus , P. peruvianus Wheeler (1925, p. 1 1) is here considered to
be a distinct species (stat. nov.). Three syntype workers in the MCZ from Chaquimayo, Peru
(leg. Holmgren) have a smooth, shining, puncticulate head whose light brown color contrasts
with the dark brown gaster; more clearly differentiated basal and declivitous faces of the
propodeum than P. ejectus (PDI 1.06-1.09); and a short, high, and (in dorsal view) thin petiole
such that PLI 0.60-0.65 and PHI 0.77-0.83 (PLI 0.43-0.57 and PHI 0.54-0.78 in P. ejectus ).

Biology. — P. ejectus nests in dead twigs or stalks of woody and herbaceous plants. Among
the P. ejectus nest series which I have examined there are records from the following plant
genera: Carya, Cladium, Conostegia, Prosopis, Quercus, Rhus , Spilanthes , Vernonia, and
Vitis. In southeastern United States alates have been collected in the months of March, June,
July, and September. Three out of five nests which I dissected (from Florida, Texas, and
Mexico) contained a single dealate female; the other two nests were queenless. In Florida some
nests of this species are polygynous (R. W. Klein, pers. comm.).

Material Examined (BMNH, GCW, LACM, MCSN, MCZ, NHMB, PSW, UCD,
USNM). —

ALABAMA: Baldwin Co.: Jackson’s Oak (W. S. Creighton); Marlow’s Ferry, Fish R. (W. S. Creighton); Mobile Co:.
Dog R., Mobile (W. S. Creighton); Irvington (Van Aller); Mobile (W. D. Pierce); Theodore (A. H. Sturtevant); Whistler
(A. H. Sturtevant); county unknown: Kushla (A. H. Sturtevant).

Nearctic Species of the genus Pseudomyrmex

233

FLORIDA: Alachua Co:. Gainesville (R. W. Klein; N. L. H. Krauss); Collier Co: Everglades (W. M. Barrows),
Immokalee (M. Deyrup); Royal Palm Pk. (Melander). Dade Co: Coconut Grove; Homestead (W. F. Buren; G. B.
Merrill); Homestead Air Force Base (G. C. & J. Wheeler); Long Pine Key (W. M. Wheeler); Miami (W. T. Davis); Old
Flamingo Rd., Everglades Natl. Pk. 10 m (P. S. Ward); Paradise Key (W. M. Wheeler); Pinelands Trail, Everglades Natl.
Pk. (G. C. and J. Wheeler); Shark Valley, Everglades Natl. Pk. (P. S. Ward); Visitor Centre, Everglades Natl. Pk. (G. C.
& J. Wheeler); no specific locality (J. N. Knull); Duval Co.:. Fort George; Highlands Co: Archbold Biol. Stn., Lake
Placid (T. C. Schneirla, J. Walker); Highlands Hammock St. Pk. (P. S. Ward); Sebring (I. E. Harper); Hillsborough Co:
Pine Crest [ = Pinecrest] (W. M. Wheeler); no specific locality (J. C. Bowyer); Indian River Co: Vero Beach (L. & C. W.
O’Brien); Leon Co: Anders Branch, Tall Timbers Res. Stn. (J. F. Lynch); Tallahassee (G. C. & J. Wheeler); Monroe Co:
Key Large (H. V. Weems); 12 mi N Key Largo (P. S. Ward); N. Key Largo (R. W. Klein); Orange Co.: no specific
locality (C. Nelson; B. L. Smith; E. Storrs); Palm Beach Co: Boynton Beach (Wood & Davidson); Palm Beach; Pinellas
Co: Belle Air [= Belleair] ; Dunedin (Blatchley); Largo (Bradley & Knorr); Taylor Co: Williams Landing (R. Smith);
Volusia Co: Haw Creek (T. Pergande?), county unknown: Everglades Natl. Pk. (G. C. & J. Wheeler).

GEORGIA: Chatham Co: Savannah (H. T. Vanderford); Decatur Co: no specific locality (Kannowski); Glynn Co.:
Brunswick (N. L. H. Krauss); Seminole Co: no specific locality (Kannowski).

LOUISIANA: Beauregard Co: DeRidder (W. Buren).

MARYLAND: St. Marys Co: Leonardtown (O. L. Cartwright).

SOUTH CAROLINA: Chesterfield Co: Cheraw St. Pk. (G. C. & J. Wheeler).

TEXAS: Bexar Co: Ft. S. Houston (R. B. Kimsey); San Antonio (P. S. Ward; R. Williams); San Antonio NE
Preserve (R. B. Kimsey); Brazoria Co: 4 mi SW West Columbia (P. S. Ward); Cameron Co: 10 mi W Boca Chica (W. S.
Creighton); Brownsville (P. J. Darlington; J. Knull); Comal Co: New Braunfels (W. M. Wheeler); Hidalgo Co: Bentsen
R. Grande St. Pk., Mission (W. S. Creighton; P. S. Ward); Santa Ana Refuge (P. S. Ward); no specific locality (J. Knull);
Kenedy Co.: 26 mi N Raymondsville (W. S. Creighton); Live Oak Co: Three Rivers (W. S. Creighton); McLennan Co:
Waco; Travis Co: Austin (G. Bush & W. L. Brown; P. S. Ward); Victoria Co: Victoria (W. E. Hinds; J. D. Mitchell).

MEXICO: Guerrero: 18 mi S Chilpancingo (F. D. Parker & L. A. Stange); Nuevo Leon: Cola de Caballo, 38 km SSE
Monterrey, 600 m (P. S. Ward); Quintana Roo: San Miguel, Cozumel I. (N. L. H. Krauss); Tabasco: Frontera (R.
Andrews); Tamaulipas: Matamoros (F. F. Bibby); Veracruz: Los Tuxtlas (R. L. Jeanne); Mirador (E. Skwarra); Palma
Sola (R. Andrews); Remutadero (E. Skwarra); Tinajas (F. D. Parker & L. A. Stange).

BELIZE: Rideau Camp (P. Broomfield).

COSTA RICA: Guanacaste Prov: Agua Caliente (W. M. Wheeler); Puntarenas Prov: Monteverde, 1200 m (P. S.
Ward); San Jose Prov: Alfombra, 850 m (P. S. Ward); San Jose (W. M. Wheeler).

JAMAICA: Lapland, Catadupa; Ford 1 mi SE Stony Hill (E. A. Chapin).

Pseudomyrmex leptosus Ward sp. nov.

(Figs. 19, 26, 34,35, 38)

Holotype queen. — Payne’s Prairie, 8 mi S Gainesville, Alachua Co., Florida, 5.viii. 1 982, R. W. Klein (MCZ)
(ex lab colony established from a mixed nest of P. leptosus and P. ejectus originally collected 16.V.1982). HW 0.70, HL
0.86, EL 0.41, PL 0.53, PH 0.27.

Paratype queens, males. — Payne’s Prairie, 8 mi S Gainesville, R.W. Klein (ex lab colony established from
mixed nest of P. leptosus and P. ejectus originally collected 16.V.1982); Gainesville, Alachua Co., Florida, R. W. Klein (ex
lab colony established from mixed nest of P. leptosus and P. ejectus originally collected 1 5.v. 1 982) (BMNH, LACM,
MCZ, PSW, UCD, USNM).

Worker. — Unknown.

Queen Measurements ( n = 11): HL 0.83-0.91, HW 0.67-0.70, MFC 0.019-0.027, Cl
0.77-0.82, 01 0.53-0.59, REL 0.47-0.51, REL2 0.59-0.63, 001 0.92-1.58, VI 0.68-0.75,
FCI 0.028-0.039, CDI 0.047-0.057, SI 0.45-0.48, SI2 0.75-0.77, FI 0.42-0.47, NI
0.60-0.68, PLI 0.47-0.51, PLI2 0.72-0.84, PHI 0.59-0.68, PWI 0.43-0.51, PWI2
0.50-0.57, PPWI 1.02-1.18, PPWI2 0.61-0.67.

Queen Diagnosis. — A small orange species with broadly rounded, shiny head (HW
0.67-0.70, VI 0.68-0.75); anterior clypeal margin medially straight or slightly convex, laterally
angulate; distance between frontal carinae less than basal width of scape; occipital margin
convex, flat, or weakly concave, in full-face, dorsal view; lateral margins of pronotum rounded;
basal face of propodeum rounding into declivitous face; petiole relatively slender, twice as long
as high; anterior face of petiole convex, in lateral view; postpetiole as wide or wider than long.
Front of head finely but densely punctulate, on a more or less smooth, shining background; fine

Quaest. Ent., 1985,21 (2)

234

Ward

punctures becoming less dense towards the occiput, which is also smooth and shining; petiole
and most of mesosoma sublucid, finely or obscurely punctulate on a weak coriarious
background; propleuron subopaque, coriarious-imbricate; postpetiole and gaster weakly
shining, the sheen partially obscured by numerous, fine piligerous punctures. Erect pilosity very
sparse; several erect setae on dorsum of head, pronotum, mesonotum, metanotum, petiole,
postpetiole and gaster; erect pilosity essentially absent from propodeum, scapes, and mid and
hind femora. Fine appressed pubescence present, but not obscuring sculpture on most parts of
body; appressed pubescence moderately dense on abdominal tergite IV, only partially obscuring
the shiny integument. Body light orange-brown; a conspicuous pair of anterolateral, dark
fuscous patches on abdominal tergite IV.

Comments. — This species is known only from queens and males collected in, or reared
from, two mixed nests of P. leptosus and P. ejectus from the vicinity of Gainesville, Florida (R.
W. Klein, leg.).3 The original colonies each contained two dealate queens of P. leptosus
(together with P. ejectus workers, and brood of both species), and Klein subsequently reared P.
leptosus alates in the laboratory.

P. leptosus queens are readily distinguishable from those of P. apache and P. seminole on
the basis of size alone (HL > 1.25 in P. apache and P. seminole , HL < 0.95 in P. leptosus). P.
leptosus queens differ from those of P. simplex by the possession of a wider, more broadly
rounded head (Cl > 0.77 VI < 0.75, in P. leptosus ; Cl < 0.77, VI >0.80, in P. simplex ),
shorter eyes (REL2 < 0.63 in P. leptosus ; REL2 > 0.65 in P. simplex ), more divergent
frontal carinae, and narrower forefemur (FI 0.42-0.47 in P. leptosus , FI 0.49-0.55 in P.
simplex). The differences between P. leptosus and P. pallidus queens are more subtle. All of
their measurements and indices overlap, although P. leptosus tends to be smaller and (more
importantly) to possess a more broadly rounded head, so that VI 0.68-0.75 (VI 0.71-0.88 in P.
pallidus). The most important distinction between the two species lies in the sculpture of the
frons and vertex: finely punctate on a predominantly smooth, shiny background in P. leptosus ;
more coarsely punctate on a sublucid, coriarious background in P. pallidus (Figures 38, 39).
Despite some size-related sculptural variation in P. pallidus even the smallest P. pallidus
queens possess more strongly developed coriarious sculpture on the head than P. leptosus
queens. The postpetiole and gaster of P. leptosus also tend to be shinier than those of P.
pallidus. There are slight differences in the male genitalia of the two species, as outlined in the
key to males.

Biology. — This species is apparently a workerless, social parasite of P. ejectus. Details on
the life history and behavior of P. leptosus will appear elsewhere (R. W. Klein, in prep.).

Material Examined (BMNH, LACM, MCZ, PSW, UCD, USNM).—

FLORIDA: Alachua Co:. Gainesville (R. W. Klein); Payne’s Prairie, 8 mi S Gainesville (R. W. Klein).

Pseudomyrmex pallidus F. Smith
(Figs. 20, 27, 36, 37, 39, 42, 43)

Pseudomyrma pallida F. Smith (1855), p. 160. One syntype queen (dealate), one syntype worker, “U.S.” (BMNF1)
[Examined], Syntype worker here designated as LECTOTYPE.

Pseudomyrma jlavidula\ Wheeler (nec F. Smith), 1905, pp. 83-85, 87 (partim)

Pseudomyrma flavidula\ Wheeler (nec F. Smith), 1908, p. 419.

3I recently received an additional collection of P. leptosus , from Munroe
Co., Florida (Hwy. 94, 10 mi. W Tamiami Ranger Station, 26.xi.1984, P.
leptosus queens in nest with P. ejectus workers, Blaine Cole leg.).

Nearctic Species of the genus Pseudomyrmex

235

Pseudomyrma pallida F. Smith; Wheeler, 1908, pp. 419-420 (partim).

Pseudomyrma Jlavidula, Mitchell & Pierce (nec F. Smith), 1912, p. 69.

Pseudomyrma pallida F. Smith; Mitchell & Pierce, 1912, p. 69.

Pseudomyrma Jlavidula, Mann (nec F. Smith) 1920, p. 405 (partim).

Pseudomyrma Jlavidula-, Wheeler & Bailey (nec F. Smith), 1920, pp. 260, 265 [ Description of larva, and contents of food
pellets ] .

Pseudomyrma flavidula var. delicatula\ Wheeler & Bailey (nec Forel), 1920, p. 265 (partim) [ Description of food pellet
contents].

Pseudomyrma Jlavidula ; Wheeler (nec F. Smith), 1932, p. 4 (partim).

Pseudomyrma pallida F. Smith; Wheeler, 1932, p. 4 (partim).

Pseudomyrma pallida F. Smith; Creighton, 1950, pp. 80-82 (partim).

Pseudomyrmex pallidus F. Smith; Wilson, 1964, pp. 4-5 (partim).

Pseudomyrmex pallidus F. Smith; Wheeler & Wheeler, 1973, pp. 41-44.

Worker Measurements ( n = 70): HL 0.78-1.06, HW 0.68-0.89, MFC 0.011-0.024, Cl
0.77-0.91, 01 0.54-0.62, REL 0.45-0.54, REL2 0.53-0.65, 001 0.78-2.08, VI 0.67-0.84,
FCI 0.015-0.033, SI 0.41-0.49, SI2 0.68-0.85, FI 0.37-0.45, PDI 1.10-1.52, MPI
0.022-0.054, NI 0.54-0.67, PLI 0.47-0.62, PWI 0.38-0.52, PPWI 0.85-1.18.

Worker Diagnosis. — Medium-sized species (for the pallidus group), with moderately broad
head (HW 0.68-0.89, Cl 0.77-0.91); anterior clypeal margin medially flat, laterally angulate:
distance between frontal carinae less than basal width of scape; eyes moderately long, EL
greater than scape length; occipital margin convex, flat or weakly concave, in full-face, dorsal
view; lateral margins of pronotum rounded; metanotal groove present but shallow; basal face of
propodeum longer than declivitous face, and more or less differentiated from it; petiole slender,
with a distinct anterior peduncle and anteroventral tooth. Head subopaque to weakly shining;
frons densely punctulate on a coriarious background; punctures less dense on the vertex which
remains (at least weakly) coriarious; dorsum of mesosoma and petiole sublucid,
coriarious-punctulate, becoming coriarious-imbricate laterally; postpetiole and gaster weakly
shining, covered with numerous, very fine piligerous punctures. Erect pilosity sparse, lacking on
mesonotum, propodeum, and mid and hind femora; one to several erect setae on dorsum of
head, pronotum, petiole, postpetiole, and abdominal tergite IV. Fine, appressed pubescence
present on most parts of body, forming a moderately dense mat on abdominal tergite IV, which
partially obscures the sheen of the integument. Body orange-brown, with paler mandibles and
appendages; a pair of anterolateral fuscous patches sometimes present on abdominal tergite IV.

Comments. — This is the most common and widespread member of the pallidus group. P.
pallidus shows considerable geographical variation in size, sculpture, and body proportions
(note wide ranges of some metrics). However the workers are consistently orange-brown in
color, with contiguous frontal carinae (MFC < 0.025), moderately long eyes (REL2 > 0.52),
and (at least weakly) coriarious-punctulate sculpture on the vertex. No other Nearctic species
possesses this combination of characters. Specific differences between P. pallidus and other
orange Pseudomyrmex ( P . apache , P. leptosus , P. seminole, and P. simplex ) are discussed
under those species.

Biology. — P. pallidus exhibits diversity in its choice of nesting sites. While it shows a
preference for dead stalks or culms of herbaceous plants, it will also nest in dead twigs or
branches of shrubs and trees in some localities.

By state, the Nearctic nest-site records are from the following plants (based on personal

observations or on museum material which I have examined):

Florida: Ambrosia artemisiifolia , Andropogon, Bidens, Cladium jamaicense , Uniola

paniculata.

Georgia: Callicarba.

Texas: Baccharis , Heterotheca subaxillaris , Iva ciliata , Melia azedarach , Prunus , Ptelea
trifoliata , Uniola paniculata.

Quaest. Ent., 1985, 21 (2)

236

Ward

Arizona: Gossypium thurberi, Quercus emoryi, Q. oblongifolia.

California: Acacia greggii, Hyptis emoryi.

The number of functional queens in a colony varies widely. The majority of P. pallidus nests
which I dissected from Texas and Florida were queenless or monogynous, but sometimes larger
numbers of mated, dealate queens cohabited (up to a maximum of 22). Since P. pallidus
colonies are often polydomous, the number of queens per colony may be higher.

P. pallidus alates have been collected in every month of the year, indicating that mating
occurs in more than one season.

Material Examined (BMNH, GCW, LACM, MCSN, MCZ, MHN, NHMB, PSW, UCD,
USNM). —

ALABAMA: Mobile Co.: Mobile (Van Aller); Spring Hill, Mobile (W. S. Creighton); Theodore (A. H. Sturtevant);
Whistler (A. H. Sturtevant); county unknown : Kushla (A. H. Sturtevant)

ARIZONA: Cochise Co.: Carr Canyon, Huachuca Mtns. 5400 ft (W. S. Creighton); Huachuca Mtns., T.24S, R.20E,
sec. 4, SW quadr., 5850-5900 ft (R. R. Snelling); Portal, Chiricahua Mtns. (R. M. Bohart); Ramsey Canyon, Huachuca
Mtns., (W. S. Creighton); Graham Co.: Cottonwood Pass, Peloncillo Mtns., 4800 ft (W. S. Creighton) Maricopa Co.:
Tempe (W. M. Wheeler); Pima Co.: Brown Canyon, Baboquivari Mtns, 4400 ft (W. S. Creighton); Forestry Cabin,
Baboquivari Mtns., 3500 ft (W. S. Creighton); Molino Basin, Santa Catalina Mtns., 4200 ft (W. S. Creighton); Santa
Cruz Co.: Bathtub Canyon, Santa Rita Mtns. (L. F. Byars); Nogales (C. A. Geesey; C. H. Spitzer); county unkown: Santa
Rita Mtns., (J. Knull).

CALIFORNIA: Riverside Co.: Deep Canyon (G. C. & J. Wheeler); San Bernardino Co.: 49 Palms, Joshua Tree
Natl. Monum., 900 m (P. S. Ward); Yucca Valley (Melander).

FLORIDA: Alachua Co.: Gainesville (C. J. Drake; R. W. Klein); Payne’s Prairie, 8 mi S Gainesville (R. W. Klein);
Collier Co.: Marco (W. T. Davis); Dade Co.: Agri. Res. Educ. Centre, Homestead (R. W. Klein); Dodge I., Miami (G.
Stegmaier); Homestead (C. W. O’Brien); Homestead Air Force Base (G. C. & J. Wheeler); Old Flamingo Rd.,
Everglades Natl. Pk., 10 m (P. S. Ward); Miami Beach (W. Wirth); no specific locality (J. Knull): Duval Co.: Fort
George; Glades Co.: Fisheating Creek, Palmdale (M. Deyrup); Highlands Co.: Archbold Biol. Stn., Lake Placid (T. C.
Schneirla); Highlands Hammock St. Pk. (C. W. O’Brien); Leon Co.: Sheep I., Tall Timbers Res. Stn. (J. F. Lynch);
Monroe Co.: Bahia Honda Recr. Area, 5m (P. S. Ward); Big Pine Key (P. S. Ward; E. O. Wilson); John Pennekamp St.
Pk., < 5m (P. S. Ward): 12 mi N Key Largo (P. S. Ward); 16 mi N Key Largo (P. S. Ward); Key West (T. Pergande);
Loggerhead Key, 1.9 mi S Cudjoe Key (R. Thorington, J. Layne & P. Cone); Lower Matecumbe Key (W. M. Wheeler);
Mrazek Pond, Everglades Natl. Pk. (L. &C. W. O’Brien): No" Name Key (P. S. Ward); Refuge Nature Trail, Big Pine
Key, 10 m (P. S. Ward); Pinellas Co.: Dunedin (Blatchley); Sarasota Co.: Long Branch Key, Sarasota (A. C. Cole);
county unknown: “Florida” (T. Pergande; S. Henshaw).

GEORGIA: Chatham Co.: nr. Savannah (R. A. Cushman); Richmond Co.: Augusta (R. R. Snelling).

LOUISIANA: Beauregard Co.: DeRidder (W. Buren); Caddo Co.: Shreveport (W. Buren; R. A. Cushman); Madison
Co.: Tallulah (E. R. Kalmbach).

MISSISSIPPI: Adams Co.: Sibley (A. Fleming); Jackson Co.: Pascagoula; Lauderdale Co.: Meridian (H. T.
Vanderford); Smith Co.: Taylorsville (W. S. Creighton)

NEW JERSEY: Cape May Co.: Dias Creek.

NORTH CAROLINA: New Hanover Co.: Wrightsville [ = Wrightsville Beach?] (W. T. Davis).

TEXAS: Bexar Co.: San Antonio (G. A. Prucia; P. S. Ward); 10 mi NW San Antonio (W. S. Ross); Brazos Co.:
College Station (R. S. Peigler); Cameron Co.: Brownsville (J. Knull); Comal Co.: New Braunfels (Darlington); Fort Bend
Co.: Richmond (Cushman & Pierce); Goliad Co.: no specific locality (J. D. Mitchell); Gonzales Co.: Palmetto St. Pk. (P.
S. Ward); Hidalgo Co.: Bentsen Rio Grande St. Pk. (P. S. Ward); Santa Ana Refuge (P. S. Ward); no specific locality (D.
J. & J. N. Knull); Kleberg Co.: Padre I. Natl. Seashore (P. S. Ward); Matagorda Co.: Wadsworth (P. S. Ward); Nueces
Co.: Mustang I. St. Pk. (P. S. Ward); Port Aransas, 5 m (P. S. Ward); 3.4 km SW Port Aransas, 5 m (P. S.Ward); 1 1 km
SW Port Aransas, 5 m (P. S. Ward); 18 km SW Port Aransas, 5 m (P. S. Ward); Travis Co.: Austin (D. Tupa; W. M.
Wheeler); Barton Creek, Austin (P. S. Ward); Brackenridge Field Stn., Austin (P. S. Ward); Victoria Co.: Victoria (J. D.
Mitchell); Willacy Co.: 7 mi N. Rio Hondo (W. S. Creighton); county unknown: Devils River (E. A. Schwartz).

MEXICO: Baja California Sur: Las Barrancas (W. M. Mann); 7 mi N. Santiago (W. H. Ewart); 2.7 mi SE Valle
Perdido (R. R. Snelling); Chiapas: Tonola (A. Petrunkewitch); Tuxtla Gutierrez (N. L. H. Krauss); Chihuahua: El Paso,
Texas, POE (V. J. Shiner); Durango: 6 mi E San Lucas, 6200 ft (W. S. Creighton); Guerrero: Acapulco (Baker); 18 mi S.
Chilpancingo (F. D. Parker & L. A. Stange); Revolcadero, nr. Acapulco (N. L. H. Krauss); Hidalgo: San Miguel (W. M.
Mann); Morelos: Cuernavaca (N. L. H. Krauss); Nayarit: Tepic; Nuevo Leon: Iturbide, 1800 m (P. S. Ward); Quintana
Roo: San Miguel, Cozumel I. (N. L. H. Krauss); Sinaloa: 1.1 mi W El Quelite (M. L. Siri); Isabel I. (H. H. Keifer);
Mazatlan (R. M. Bohart; P. J. Spangler); 20 mi S Villa Union (E. I. Schlinger); Sonora: Alamos (A. Mintzer); 4.8 mi S
Cananea (V. Roth); Cocorit (F. D. Parker & L. A. Stange); 5 mi N Santa Cruz, 4700 ft (W. S. Creighton); Tamaulipas:
Brownsville [= Matamoros?]; Veracruz: Cordoba; Jalapa (N. L. H. Krauss); La Buena Ventura (A. Petrunkewitch); Los
Tuxtlas (R. L. Jeanne); Mirador (E. Skwarra); Veracruz; state unknown: Tetela [Oaxaca or Puebla].

Nearctic Species of the genus Pseudomyrmex

237

BAHAMAS: Mangrove Cay, Andros I. (W. M. Mann); Nassau (W. M. Wheeler); San Salvador I. (J. F. Lynch);
South Bimini I. (C & P. Vaurie): Watlings I (J. Greenway).

BELIZE: Augustine (J. Reiskind); Rideau Camp (P. Broomfield).

COSTA RICA: Cartago Prov .: Cartago (N. L. H. Krauss); Paraiso (N. L. H. Krauss); Guanacaste Prov .: 15 km SW
Bagaces, Comelco (H. V. Daly); Finca la Pacifica, 7 km SW Canas (H. V. Daly); Hacienda la Pacifica, nr. Canas, 50 m
(P. S. Ward); 1 km SW Pto. Coyote, < 5 m (P. S. Ward); Santa Rosa Natl. Pk., < 5 m, 270 m (P. S. Ward); Limon
Prov.: Linda Vista, 540 m (P. S. Ward); Puerto Viejo, < 5 m (P. S. Ward); Puntarenas Prov.: Monteverde (H. V. Daly);
Monteverde, 1220 m, 1 350 m, 1400 m (P. S. Ward); San Jose Prov.: 1 km N La Ese, 1400 m (P. S. Ward); Pavas, 1000 m
(P. S. Ward); San Jose (W. M. Wheeler); Hamburg Farm (F. Nevermann).

CUBA: Santa Barbara, Isla de Pinos (S. C. Bruner).

EL SALVADOR: Cerro Verde (L. J. Bottimer); La Libertad (N.L.H. Krauss); San Salvador (L. J. Bottimer).
GUATEMALA: Antigua (W. M. Wheeler); San Lucas, Toliman (W. M. Wheeler).

HONDURAS: La Ceiba (F. J. Dyer).

Pseudomyrmex seminole Ward, sp. nov.

(Figs. 18,21,28,44, 45)

Pseudomyrma flavidula ; Wheeler (nec F. Smith), 1905, pp. 83-85 (partim).

Pseudomyrma pallida ; Wheeler (nec F. Smith), 1932, p. 4 (partim).

Holotype worker: John Pennekamp State Pk., Munroe Co., Florida, < 5 m, 14.ix.1982, ex colony in dead
Andropogon culm, roadside near mangrove, P. S. Ward acc. no. 5723 (MCZ). HW 0.90, HL 1.05, EL 0.50, PL 0.59,
PH 0.29.

Paratype workers, queens, males: Two nest series from John Pennekamp St. Pk., Munroe Co., Florida,
1 4.ix. 1 982, ex dead Andropogon culms, P. S. Ward acc. nos. 5722, 5723; two nest series and ground foragers from 12
mi N. Key Largo, Munroe Co., Florida, 1 0.i. 1 979, ex dead Andropogon culms, and foraging on ground, P. S. Ward
acc. nos. 3199, 3202, 3203 (BMNH, LACM, MCZ, PSW, UCD, USNM).

Type series is restricted to material from these two adjacent localities. Other specimens
believed to be conspecific are listed below under “Material Examined”.

Worker Measurements (n = 26): HL 0.98-1.16, HW 0.87-0.96, MFC 0.029-0.042, Cl
0.81-0.89, 01 0.57-0.63, REL 0.43-0.48, REL2 0.53-0.56, OOI 1.04-1.94, VI 0.75-0.85,
FCI 0.031-0.47, SI 0.42-0.47, SI2 0.75-0.85, FI 0.38-0.44, PDI 1.05-1.40, MPI
0.017-0.045, NI 0.53-0.62, PLI 0.46-0.53, PWI 0.39-0.47, PPWI 0.93-1.09.

DPL 0.75-0.87, MP 0.015-0.042, CDI 0.040-0.064, PLI2 1.11-1.34, PHI 0.56-0.67,
PWI2 0.50-0.59, PPWI2 0.48-0.57.

Worker Diagnosis:. — Relatively large species, with broad head (HW 0.87-0.96, Cl
0.81-0.89); median portion of anterior clypeal margin obtusely (and weakly) angulate, sharply
angulate laterally (Figure 44); distance between frontal carinae subequal to, or slightly less
than, basal width of scape; eyes relatively short (REL 0.43-0.48); occipital margin convex or
flat, in full face, dorsal view; lateral margins of pronotum rounded; metanotal groove wide but
shallow; basal face of propodeum rounding into declivitous face, the former equal to, or longer
than, the latter; petiole and postpetiole long and slender, the former with a distinct anterior
peduncle and (usually) prominent, rounded anteroventral tooth. Mandibles very weakly striate,
with scattered punctures; head subopaque to sublucid, densely punctulate on a coriarious
background; mesosoma and petiole subopaque, coriarious-punctulate, becoming
coriarious-imbricate laterally; postpetiole and gaster subopaque, with numerous fine piligerous
punctures. Erect pilosity sparse, lacking on mesonotum and propodeum; one to several pairs of
erect setae on dorsum of head, pronotum, petiole, postpetiole and abdominal tergite IV. Fine
appressed hairs present on most parts of body, and forming a rather dense mat on abdominal
tergite IV. Body orange brown; mandibles and apices of appendages variably paler.

Comments. — Essentially a Gulf Coast species, P. seminole occurs sympatrically with the
closely related P. pallidus. Workers of P. seminole may be recognized by the less convergent

Quaest. Ent., 1985,21 (2)

238

Ward

frontal carinae (MFC > 0.029 in P. seminole, < 0.024 in P. pallidus), shorter eyes, and
weakly angulate median portion of the anterior clypeal margin (compare Figures 42 and 44).
The angulate clypeus of P. seminole tends to be a little more produced than that of P. pallidus
workers (CDI 0.040-0.064 in P. seminole , 0.030-0.054 in P. pallidus ). On average, the petiole
and postpetiole of P. seminole are longer and more slender than those of P. pallidus, but there
is sufficient variation in both species that the relevant metrics overlap broadly. Differences
between the queens of the two species are more pronounced and the male genitalia of P.
seminole are quite distinct (see keys to queens and males).

Biology. — I have collected nests of P. seminole in dead stalks or culms of Andropogon,
Heterotheca subaxillaris, Uniola paniculata, and an unidentified woody legume. None of these
nests was polygynous; some were queenless, indicating that this species is polydomous.

On Padre Island, east Texas P. seminole is patchily distributed in a continuous population of
P. pallidus. Both species use the same nest sites ( Heterotheca stalks and Uniola culms). I have
observed incipient P. seminole colonies consisting of (i) a single, dealate queen, (ii) a single,
dealate queen with brood, and (twice) (iii) a single dealate queen in association with P.
pallidus workers. In one of the latter instances a dealate P. pallidus queen and five workers
occupied one Uniola internode, while the P. seminole queen occupied an adjacent cavity; in the
second instance, the P. seminole queen coexisted with seven P. pallidus workers (but no queen)
plus brood of unknown identity, in a single dead Heterotheca stalk. These observations suggest
that P. seminole may be a facultative, temporary social parasite of P. pallidus.

Alates or alate pupae have been collected in most months of the year, indicating a rather
continual production of sexuals.

Material Examined (BMNH, GCW, LACM, MCZ, PSW, UCD, USNM).—

FLORIDA: Alachua Co.: Gainesville (R. W. Klein); no specific locality (T. H. Hubbell); Dade Co.: Agric. Res. Educ.
Centre, Homestead (R. W. Klein); Homestead Air Force Base (G. C. & J. Wheeler); Paradise Key (H. & A. Howden);
De Soto Co.: Prairie Creek, 11 mi S. Arcadia (M. Deyrup); Highlands Co.: Highlands Hammock St. Pk. (C. W. O’Brien);
Hillsborough Co.: no specific locality (B. P. Moore); Indian River Co.: Vero Beach (L. & C. W. O’Brien); Monroe Co.:
John Pennekamp St. Pk. < 5 (P. S. Ward); Key Largo (A. C. Cole); 12 mi N Key Largo (P. S. Ward); Osceola Co.:
Kissimmee; Pinellas Co.: Dunedin (Blatchley); Polk Co.: Lakeland (W. T. Davis).

LOUISIANA: East Baton Rouge Co.: Baton Rouge (M. R. Smith); Iberia Co.: New Iberia (A. H. Sturtevant).

MISSISSIPPI: Harrison Co.: Gulfport.

TEXAS: Cameron Co.: Brownsville (J. Knull); Nueces Co.: Mustang I. St. Pk. (P. S. Ward); Port Aransas, 5 m (P. S.
Ward); 3.4 km SW Port Aransas, 5 m (P. S. Ward); 1 1 km SW Port Aransas, 5 m (P. S. Ward).

MEXICO: Tamaulipas: 7 km WSW El Encino, 140 m (P. S. Ward)

BAHAMAS: Nassau (W. M. Wheeler).

Pseudomyrmex simplex F. Smith
(Figs. 22, 29,40,41)

Pseudomyrma simplex F. Smith 1877, p. 64. Holotype (unique syntype) worker, Sao Paulo [“St. Paul”], Brazil (BMNH)
[Examined].

Pseudomyrma delicatula Forel, 1899, p. 93. Syntype workers, one dealate queen, Kingston, Jamaica (Forel) (MHN)
[Examined.] Syn. nov.

Pseudomyrma delicatula var. panamensis Forel, 1899, p. 93. Holotype (unique syntype) worker, Pantaleon, Guatemala,
1700 ft (Champion) (MHN) [Examined]. Syn. nov.

Pseudomyrma delicatula var. capperi Forel, 1899, p. 93. Syntype workers, Jamaica (Capper) (MHN) [Examined]. Syn.
nov.

Pseudomyrma acanthobia race delicatula, var. vittata Forel, 1912, p. 26. Syntype workers, Ceara, Brazil (Rocha)
(MHN) [Examined]. Syn. nov.

Pseudomyrma flavidula var. delicatula Forel; Wheeler, 1913, p. 484.

Pseudomyrma flavidula var. delicatula Forel; Wheeler & Mann, 1914, p. 17.

Pseudomyrma flavidula; Wheeler (nec F. Smith), 1932, p. 4 (partim).

Pseudomyrma pallida-, Creighton (nec F. Smith), 1950, pp. 80-82 (partim).

Nearctic Species of the genus Pseudomyrmex

239

Pseudomyrmex pallidus Wilson (nec F. Smith), 1964, pp. 4-5 (partim).

Worker Measurements ( n = 46): HL 0.70-0.92, HW 0.55-0.74, MFC 0.009-0.017, Cl

0.75-0.85, OI 0.49-0.55, REL 0.52-0.61, REL2 0.62-0.77, 001 0.55-1.92, VI 0.68-0.83,

FCI 0.013-0.025, SI 0.40-0.48, SI2 0.56-0.72, FI 0.45-0.52, PDI 1.27-1.94, MPI
0.005-0.043, NI 0.53-0.65, PLI 0.50-0.69, PWI 0.39-0.60, PPWI 0.97-1.31.

Worker Diagnosis. — Relatively small species, with moderately elongate head (HW

O. 55-0.74, Cl 0.75-0.85); median portion of anterior clypeal margin straight, laterally
angulate; frontal carinae very closely contiguous (minimum distance between them much less
than the basal width of scape); eyes relatively long (REL 0.52-0.61); occipital margin concave,
flat, or slightly convex, in full face, dorsal view; lateral margins of pronotum rounded;
metanotal groove very weak and shallow; basal face of propodeum longer than declivitous face,
and more or less clearly differentiated from it; petiole relatively short, with an anterior
peduncle and anteroventral tooth; postpetiole often wider than long, somewhat globose in
lateral view. Head predominately smooth and shining; frons with scattered fine punctures of
variable size, on a smooth or obsoletely coriarious background; punctures less dense on the
vertex, which is smooth and shining; mesosoma and petiole sublucid, dorsally weakly
coriarious-punctulate, laterally coriarious-imbricate; postpetiole and gaster more or less smooth
and shining. Erect pilosity sparse, lacking on mesonotum, propodeum, mid and hind femora,
and (often) petiole; typically a pair of erect setae on pronotum, postpetiole, and adjacent to the
eyes. Fine, appressed pubescence very sparse, notably so on postpetiole and abdominal tergite
IV. Light orange brown, mandibles and clypeus a paler luteous; a pair of anterolateral fuscous
patches usually present (sometimes weak) on abdominal tergite IV.

Comments. — This species is the smallest one of a trio of orange Pseudomyrmex ( P .
pallidus , P. seminole , P. simplex ) which have been confused repeatedly in the United States.

P. simplex workers are recognizable by their smooth, shiny, puncticulate head; broad
forefemur (FI > 0.45); and shining fourth abdominal tergite which is devoid of a dense mat of
appressed pubescence. In addition the workers have long eyes, closely contiguous frontal
carinae, a very shallow metanotal groove, and a short petiole and postpetiole. The simplex and
delicatulus types share these essential features, along with the other material which I have
examined. There is a fair range of variation in size and body proportions (see metrics). P.
simplex tends to be lighter in color than related species, and the fuscous patches on abdominal
tergite IV are usually conspicuous, at least in Florida populations (less so in Central and South
America).

Biology. — P. simplex shows a preference for nesting in dead twigs of woody shrubs or trees,
rather than in dead stalks of herbaceous plants. In Florida, I have collected nests in dead twigs
or stalks of Baccharis halimifolia, Cladium jamaicense, Laguncularia racemosa Metopium
toxiferum, and Nectandra coriacea\ there are also museum records from Carya floridana and
Swietenia mahagoni.

In Costa Rica I recorded nests of P. simplex in dead twigs of Anacardium, Ardisia revoluta,
Avicennia germinans, Conocarpus erectus, Gliricidia sepium. Hibiscus tiliaceus , and
Terminalia catappa.

Most P. simplex nests I dissected were queenless (indicating a high level of polydomy),
some were monogynous, and one contained two functional (i.e. inseminated, with
v ell-developed ovaries) lealate queens.

Alates have been collected in May, June, and September in Florida.

Material Examined fBMNM, GCW LACM, MCSN, MCZ. MHN, NHMB PSW, UCD,
USNM). —

Quaest. Ent., 1985, 21 (2)

240

Ward

FLORIDA: Charlotte Co.: Punta Gorda (W. T. Davis); Collier Co.: Marco (W. T. Davis); Dade Co.: Biscayne Bay
(A. Slosson); Long Pine Key, Everglades Natl. Pk., 10 m (P. S. Ward); Mahogany Hammock, Everglades Natl. Pk. (G. C.
& J. Wheeler); Miami (G. B. Merill; C. F. W. Muesebeck; C. Stegmaier); Paradise Key (H. S. Barber); Pinelands Trail,
Everglades Natl. Pk. (G. C. & J. Wheeler); no specific locality (J. N. Knull); Highlands Co.: Archbold Biol. Stn., Lake
Placid (J. Walker); Monroe Co.: Big Pine Key (P. S. Ward; E. O. Wilson) John Pennekamp St. Pk.,< 5 m (P. S. Ward);
N Key Largo (R. W. Klein); 16 mi N Key Largo (P. S. Ward); Refuge Nature Trail, Big Pine Key, 10 m (P. S. Ward);
Key West; Pinellas Co.: Dunedin (Blatchley); Sarasota Co.: Long Branch Key, Sarasota (Cole).

MEXICO: Quintana Roo: San Miguel, Cozumel I. (N. L. H. Krauss); Tamaulipas: Tampico (F. C. Bishop).
BAHAMAS: Gun Point, Crooked I. (B. Valentine & A. Hamilton); Mangrove Cay, Andros I. (B. Cole); New
Providence (B. Cole).

BELIZE: Belize (N. L. H. Krauss); El Cayo (N.L.H. Krauss); Punta Gorda (P. Broomfield).

CAYMAN IS.: Grand Cayman (M. E. C. Giglioli)

COSTA RICA: Guanacaste Prov.: Hacienda la Pacifica, nr. Caiias, 50 m (P. S. Ward); 1 km SW Pto. Coyote, < 5 m
(P. S. Ward); Santa Rosa Natl. Pk., < 5 m, 5 m, (P. S. Ward); Taboga Hill (C. R. Carroll); Limon Prov.: Cahuita Natl.
Pk. < 5 m (P. S. Ward); Puntarenas Prov.: Lagarto, 120 m (P. S. Ward); Llorona, Corcovado Natl. Pk., 10 m (P. S.
Ward); Manuel Antonio Natl. Pk. 5 m, 20 m (P. S. Ward); Osa Peninsula, Corcovado (J. Longino); San Jose Prov.: San
Jose (W. M. Wheeler); province unknown: “Costa Rica” (Tonduz).

CUBA: Cogimar (W. M. Wheeler); Yunquede, Baracoa, Ote (P. J. Darlington).

EL SALVADOR: Quezaltepeque (M. Irwin & D. Cavagnaro).

GUATEMALA: Escuintla (P. J. Spangler); Livingston (Barber & Schwartz); Pantaleon (Champion); “Guatemala”
(Stoll).

HAITI: Cape Haitien (W. M. Mann).

HONDURAS: La Ceiba (F. J. Dyer); Tegucigalpa (F. J. Dyer).

JAMAICA: Balaclava (Wight); Kingston (A. Forel; P. Vogel); Lapland, Catadupa; Mandeville (Wight); Montego
Bay; “Jamaique” (Capper).

PANAMA: Ancon, Canal Zone (W. M. Wheeler); Barro Colorado I., Canal Zone (W. L. Brown and E. S.
McCluskey; Zetek); Cristobal, Canal Zone (H. F. Dietz); 2 km SE Fort Kobbe, Canal Zone, 10 m (P. S. Ward); 5 km
WNW Gatun Dam, Canal Zone, 160 m (P. S. Ward).

PUERTO RICO: Mayagiiez (M. R. Smith).

TRINIDAD: Port of Spain (R. Thaxter); St. George (J. Noyes).

WEST INDIES: St. Lucia (N. A. Weber).

BRAZIL: Amazonas: Rio Taruma Mirim-Igapo (J. Adis); Bahia: Bondaz; Ceara: no specific locality (Rocha); Para:
Ourem; Santarem, Taperinha (R. L. Jeanne); Tacura; Tucurul (W. L. Overal); Paraiba: Independencia (Mann & Heath);
Rio de Janeiro: Mendes (Eidmann); Sao Paulo: Sao Paulo.

COLOMBIA: Huila (B. & E. MacKay); Serrania de Macuira, 6-8 km S. Nazareth, 70-200 m (W.L. Brown & R. C.
Kugler).

ECUADOR: Rio Palenque (L. Gillespie).

PERU: Piura (Townsend).

DISCUSSION

Coexistence of congeners

Every Nearctic species of Pseudomyrmex occurs sympatrically with two or more congeners
in at least some portion of its range. Where two or more species co-occur, they often use a
broadly overlapping array of nest-sites. For example, in the Florida Keys, Cladium culms are
occupied by both P. pallidus and P. simplex , although P. simplex also nests in woody twigs,
and P. pallidus will nest in Andropogon culms (a nest-site shared with P. seminole ); on Padre
Island, east Texas P. pallidus and P. seminole occupy the same nest-sites ( Uniola culms and
Heterotheca stalks); in northern Mexico, P. ejectus and P. brunneus were both recorded
nesting in dead mint stalks at the same location. In none of the above instances were workers of
the coexisting species found together in the same individual nest-site, but they could be found
in adjacent stalks separated by only a few meters. The impression to be gained from these field
observations is that there is a rather high degree of overlap among related species using the
dead stalks or culms of herbaceous plants. These nest-sites can be expected to have a short
half-life, relative to dead woody twigs or branches. The ephemeral nature and continual
production of such sites may allow the coexistence of nest-site competitors, in a manner
analagous to competing fish on coral reef patches (Sale, 1977).

Nearctic Species of the genus Pseudomyrme x

241

Figure 46. Northern limits of some Nearctic Pseudomyrmex. All of these species, except P. seminole, range south through
Central America.

Geographical distribution and speciation

Although most Nearctic Pseudomyrmex species show extensive overlap of their
geographical ranges, each species has a rather distinctive northern limit (Figure 46). This
variable penetration into North America of essentially Neotropical taxa results in a gradient of
species diversity which is maximal in southern Texas and southern Florida. The disjunct
distributions across the Gulf of Mexico suggest a possible basis for previous differentiation and
speciation. Thus P. cubaensis may represent an earlier Florida-Antillean isolate cut off from
Central American populations of P. elongatus by a cooling trend. By this interpretation,
contemporary populations of P. elongatus in Florida and Texas (which show some
morphological differentiation) represent the severance of a more recent Gulf Coast connection.

Other closely related species of Nearctic Pseudomyrmex have rather different distribution
patterns. In three cases, the range of one member of a sibling species pair is rather limited in

Quaest. Ent., 1985,21 (2)

242

Ward

extent and is completely enclosed within the range of the other member ( P . brunneus by P.
ejectus, P. leptosus by P. pallidus, P. seminole by P. pallidus ). This suggests that the more
localized species was derived from a divergent, daughter population of the widespread species
(Type lb allopatric speciation in the parlance of Bush (1975)). One might even question
whether the differentiation always proceeded allopatrically, since P. seminole shows evidence of
being a temporary social parasite of P. pallidus , its presumptive ancestor. Moreover the social
parasitic species P. leptosus is very localized and is surrounded by, sympatric with, and
morphologically similar to P. pallidus (although its only known host is P. ejectus , a less closely
related species). In any event, differentiation to the point of attaining reproductive isolation
appears to be a plausible event on both a local and a broad geographical scale.

ACKNOWLEDGEMENTS

I am grateful to the following persons for loans of material or access to collections: Cesare
Baroni-Urbani (NHMB), Claude Besuchet (MHN), Barry Bolton (BMNH), Max Fischer
(NHMV), A1 Newton (MCZ), Roberto Poggi (MCSN), David R. Smith (USNM), Roy
Snelling (LACM), and G. C. & J. Wheeler (GCW). Additional valuable material was received
from Blaine Cole and Rudi Klein. I thank Bill Brown for comments on the manuscript. This
work was supported by NSF DEB-8204230.

LITERATURE CITED

Bush, G. 1975. Modes of animal speciation. Annual Review of Ecology and Systematics
6:339-364.

Creighton, W. S. 1950. The ants of North America. Bulletin of the Museum of Comparative
Zoology 104:1-185.

Creighton, W. S. 1952. Pseudomyrmex apache , a new species from southwestern United
States. Psyche 59:131-142.

Creighton, W. S. 1954. Additional studies on Pseudomyrmex apache. Ibid., 61:9-15.

Creighton, W. S. 1955. Observations on Pseudomyrmex elongata. Journal of the New York
Entomological Society 63:17-20.

Creighton, W. S. 1963. Further observations on Pseudomyrmex apache. American Museum
Novitates 2156:1-4.

Enzmann, E. V. 1945. Systematic notes on the genus Pseudomyrma. Psyche 51:59-103.

Forel, A. 1899. Formicidae. Biologia Centrali-Americana. Hymenoptera 3:1-160.

Forel, A. 1901. Varietes myrmecologiques. Annales de la Societe Entomologique de Belgique
45:334-382.

Forel, A. 1906. Fourmis neotropiques nouvelles ou peu connues. Ibid., 50:225-249.

Forel, A. 1912. Formicides neotropiques. IV. Sous-famille Myrmicinae (suite). V. Sous-famille
Dolichoderinae. VI. Sous-famille Camponotinae. Memoires de la Societe Entomologique de
Belgique 20:1-92.

Forel, A. 1913. Fourmis d’Argentine, du Bresil, du Guatemala et de Cuba. Bulletin de la
Societe Vaudoise des Sciences Naturelles 49:203-250.

Harris, R. A. 1979. A glossary of surface sculpturing. California Department of Food and
Agriculture Occasional Papers in Entomology No. 28.

Kempf, W. W. 1958. Estudos sobre Pseudomyrmex. II. Studia Entomologica 1:433-462.

Nearctic Species of the genus Pseudomyrmex

243

Mann, W. M. 1920. Additions to the ant fauna of the West Indies and Central America.

Bulletin of the American Museum of Natural History 62:403-439.

Mayr, G. 1870. Formicidae novogranadenses. Sitzunberichte der Akademie der
Wissenschaften Wien 61:370-417.

Mitchell, J. D. and W. D. Pierce. 1912. The ants of Victoria County, Texas. Proceedings of the
Entomological Society of Washington 14:67-76.

Roger, J. 1863. Die neu aufgefuhrten Gattungen und Arten meines Formiciden-Verzeichnisses.
Berliner Entomologische Zeitschrift 7:131-214.

Sale, P. F. 1977. Maintenance of high diversity in coral reef fish communities. American
Naturalist 111:337-359.

Smith, D. R. 1979. Formicoidea. In Krombein, K. V. et tf/.(Eds.). Catalogue of Hymenoptera
in America north of Mexico. Vol. 2. Washington, D.C.: Smithsonian Institution Press, pp.
1323-1467.

Smith, F. 1855. Descriptions of some species of Brazilian ants belonging to the genera
Pseudomyrma, Eciton and Myrmica. Transactions of the Entomological Society of London
3:156-169.

Smith F. 1858. Catalogue of hymenopterous insects in the collections of the British Museum.
Part VI. Formicidae.

Smith, F. 1877. Descriptions of new species of the genera Pseudomyrma and Tetraponera,
belonging to the family Myrmicidae. Transactions of the Entomological Society of London,
1877, pp. 57-72.

Wheeler, G. C. and J. Wheeler. 1956. The ant larvae of the subfamily Pseudomyrmecinae.

Annals of the Entomological Society of America, 49:374-398.

Wheeler, G. C. & J. Wheeler. 1973. Ants of Deep Canyon. Philip L. Boyd Deep Canyon Desert
Research Center, University of California, Riverside.

Wheeler, W. M. 1901. Notices biologiques sur les fourmis mexicaines. Annales de la Societe
Entomologique de Belgique 45:199-205.

Wheeler, W. M. 1905. The ants of the Bahamas, with a list of the known West Indian species.

Bulletin of the American Museum of Natural History 21:79-135.

Wheeler, W. M. 1908. The ants of Texas, New Mexico, and Arizona. I. Ibid., 24:399-485.
Wheeler, W. M. 1913a. The ants of Cuba. Bulletin of the Museum of Comparative Zoology
54:479-505.

Wheeler, W. M. 1913b. Ants collected in the West Indies. Bulletin of the American Museum
of Natural History 32:239-244.

Wheeler, W. M. 1925. Neotropical ants in the collections of the Royal Museum of Stockholm.
Arkiv for Zoologi 1 7 A(8): 1 —55.

Wheeler, W. M. 1932. A list of the ants of Florida with descriptions of new forms. Journal of
the New York Entomological Society 40:1-17.

Wheeler, W. M. 1942. Studies of neotropical ant-plants and their ants. Bulletin of the Museum
of Comparative Zoology 90:1-262.

Wheeler, W. M. and 1. W. Bailey. 1920. The feeding habits of pseudomyrmine and other ants.

Transactions of the American Philosophical Society 22:235-279.

Wheeler, W. M. and W. M. Mann. 1914. The ants of Haiti. Bulletin of the American Museum
of Natural History 33:1-61.

Whitcomb, W. H., H. A. Denmark, W. F. Buren and J. F. Carroll. 1972. Habits and present
distribution in Florida of the exotic ant, Pseudomyrmex mexicanus. Florida Entomologist

Quaest. Ent., 1985,21 (2)

244

Ward

55:31-33.

Wilson, E. O. 1964. The ants of the Florida keys. Breviora 210:1-14.

Appendix I. Pseudomyrmex queens and males. Ranges of metric measurements and indices.

Nearctic Species of the genus Pseudomyrmex

245

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QUAESTIONES ENTOMOLOG1CAE ISSN 0033-5037

A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.

Volume 21 Number 3 1985

CONTENTS

Peck and Anderson-Taxonomy, phylogeny and biogeography of the Carrion Beetles

of Latin America (Coleoptera: Silphidae) 247

Hilchie-The tiger beetles of Alberta (Coleoptera: Carabidae, Cicindelini) 319

Ball-Characteristics and evolution of elytral sculpture in the tribe Galeritini

(Coleoptera: Carabidae) 349

Book Review-Manson, D.C.M. 1984. Fauna of New Zealand; Number 4 369

TAXONOMY, PHYLOGENY AND BIOGEOGRAPHY OF THE CARRION BEETLES OF
LATIN AMERICA (COLEOPTERA: SILPHIDAE)

Stewart B. Peck
Department of Biology
Carleton University
Ottawa, K1S 5B6, CANADA

Robert S. Anderson
Department of Entomology
University of Alberta

Edmonton, T6G 2E3, CANADA Quaes tiones Entomologicae

21:247-317 1985

ABSTRACT

The species of Silphidae, excluding Agyrtidae, are reviewed for Mexico, Central America,
and South America. Keys are provided for the identification of adults of six genera (Necrodes,
Heterosilpha, Oiceoptoma, Thanatophilus, Oxelytrum, Nicrophorus) and 24 species known or
suspected to occur in Mexico or southward. No new species are proposed. The following new
synonymies are presented: Silpha microps Sharp is a junior synonym of Oxelytrum anticola
(Guerin-Meneville) and Hyponecrodes opacus Portevin is a junior synonym of Oxelytrum
erythrurum ( Blanchard ).

Given for each species, as appropriate, are: synonymy, diagnosis, comments on variation,
geographic distribution, seasonality, ecological data, and illustrations of important structural
characteristics. Geographic distributions are mapped for all species.

Oxelytrum is regarded as the sister genus to Ptomaphila of the Australian region.
Oxelytrum, represented by eight species, probably originated and diversified during the
Tertiary in South America when this continent was isolated. Two lineages of Oxelytrum are
recognized based on adult characters. The emarginatum group has four species found in
northern and eastern lowland and mid-elevation montane habitats, with only one species, O.
discicolle, ranging into Central America and north to extreme southern Texas. The lineatocolle
group has four species found in south-western coastal lowlands and low to high elevation
montane habitats.

Nearctic species of Necrodes, Oiceoptoma, Heterosilpha and Thanatophilus which also
occur in Latin America range no further south than the Isthmus of Tehuantepec. Only T.
graniger is endemic, found in high elevation habitats of central and northern Mexico.

Nicrophorus is represented by nine species in three species groups in Latin America. All
groups are of northern origin. The five species of Nicrophorus endemic to Latin America are
all members of the orbicollis group. Ancestors of the three South American endemic species
probably moved south along the mountain axis of Central and South America in late
Cretaceous or early Tertiary time and likely diversified in the Tertiary following
fragmentation of forest habitats. The two Middle American endemics are probably the result
of a second, but mid-Tertiary, inter-island dispersal of a northern ancestor. The four
remaining species of Nicrophorus represent two species groups. All of these species occur in
the United States with three ranging into arid areas of central and northern Mexico: the

248

Peck and Anderson

fourth ranges south to El Salvador.

A classification of New World Nicrophorus is also presented. Thirteen of the fifteen New
World species are placed in four species groups based on larval and adult characters. Two
species are Incertae sedis. A reconstructed phytogeny is presented for the New World members
of each species group.

RESUMEN

Se revisan las especies de Silphidae, excluyendo Agyrtidae, de Mexico, Centroamerica y Suramerica. Se proveen
claves para la identificacion de los adultos de los 6 generos (Necrodes, Heterosilpha, Oiceoptoma, Thanatophilus,
Oxelytrum, Nicrophorusj y 24 especies que se conoce o se sospecha que existen en Mexico o hacia el sur. No se proponen
nuevas especies. Se presentan, a continuacion, los nuevos sinonimos: Silpha microps Sharp es un sindnimo reciente de
Oxelytrum anticola ( Guerin-Meneville ) e Hyponecrodes opacus Portevin es un sindnimo reciente de Oxelytrum
erythrurum ( Blanchard ).

Se dan para cada especie, segUn sea apropiado: la sinonimia, el diagnbstico, comentarios sobre la variacion, la
frecuencia por estacion, datos ecologicos e ilustraciones de caracteristicas estructurales importantes. Se presentan las
distribuciones geograficas de todas las especies.

Oxelytrum se considera el genero hermano de Ptomaphila de la region australiana. Oxelytrum, representado por ocho
especies, probablemente se origino y diversified en Suramerica durante el Terciario cuando este continente se encontraba
aislado. Basados en las caracteristicas de los adultos, se reconocen dos linajes en Oxelytrum. El grupo emarginatum
tiene 4 especies que se encuentran en habitats montanos de baja y media altitud en el este y norte de norteamerica, con
una sola especie, O. discicolle, que se extiende a Centroamerica y de alii hacia el norte hasta el extremo sur de Texas. El
grupo lineatocolle tiene 4 especies que se encuentran en areas de baja elevacion en la costa suroeste y en habitats
montanos de alta y baja altitud.

Las especies neoarticas Necrodes, Oiceoptoma, Heterosilpha y Thanatophilus que tambien se presentan en
Latinoamerica se extienden, hacia el sur, no mas alia del Istmo de Tehuantepec. T. graniger es la t mica endemica,
encontrandose en habitats de alta elevacion en el centro y norte de Mexico.

Nicrophorus esta representado por 9 especies en 3 grupos de especies. Todos los grupos tienen su or i gen en el norte.
Las 5 especies de Nicrophorus, endemicas para Latinoamerica, son todas miembros del grupo orbicallis. Los ancestros de
las 3 especies endemicas de suramerica probablemente se dispersaron al sur a traves del eje montahoso de Centro y Sur
America en el Cretaceo tardio o a comienzos del Terciario y, es muy posible, que se diversificaran en el Terciario,
siguiendo la fragmentacion de los habitats forestales. Las dos endemicas de Mesoamerica son, probablemente, el
resultado de una dispersion secundaria a traves de islas, a partir de un ancestro norteho durante el Terciario medio. Las
cuatro especies restantes de Nicrophorus representan dos grupos de especies. Todas estas especies se encuentran en los
Estados Unidos con tres que se extienden hasta zonas aridas del centro y norte de Mexico; la cuarta, se extiende, hacia el
sur, hasta El Salvador.

Se presenta tambien una clasificacion de Nicrophorus para el Nuevo Mundo. Trece de las quince especies del Nuevo
Mundo se agrupan en 4 grupos de especies basados en caracteres de las larvas y los adultos. Dos especies son Incertae
sedis. Se presenta una reconst ruccibn filogenetica para los miembros de cada grupo de especies del Nuevo Mundo.

INTRODUCTION

The Silphidae, or carrion beetles, are the predominant beetles scavenging on dead terrestrial
vertebrate remains in temperate and sub-arctic regions in the Northern Hemisphere. Silphids
also occur in tropical lowlands, as well as in tropical montane and south temperate regions.
However, their role in the carrion-feeding insect guild is noticeably less in tropical than in
temperate regions (Cornaby, 1974; Jiron and Cartin, 1981). They are probably less abundant
in lowland tropical regions because they are less able to compete with increased rates of
bacterial decomposition and feeding of ants and fly larvae, and with the greater abundance of
carrion scavenging vertebrates (Arnett, 1946; Janzen, 1976). Recent reviews of the silphid
fauna of North America north of Mexico recognized 29 species in 8 genera (Anderson and
Peck, 1985), most of which have their relationships with species in Europe and Asia. Some of
these also have distributions extending south into Mexico. However, most of the Latin
American (used herein to indicate all of Mexico, Central America, and South America) fauna

Carrion Beetles of Latin America

249

consists of species in the primarily South American genus Oxelytrum and of endemic species in
the genus Nicrophorus. No silphids are known to occur on the islands of the Carribean.

The Latin American silphids were last revised by Portevin (1926). His liberal use of
infraspecific categories, inadequately illustrated and complex keys, which also served as
descriptions, and vague distributional data have led to problems in interpreting the species in
Latin America. This present work attempts to alleviate these problems by reviewing and
revising available knowledge about classification, distribution, and relationships of the Latin
American silphid fauna.

NATURAL HISTORY

Silphid beetles are commonly called carrion beetles because of their association with dead
vertebrate carcasses. Both adults and larvae of most species are scavengers and eat carrion.
Based on studies of Nearctic species, silphids feed in two different ways. In the first, both
adults and larvae of the sub-family Silphinae feed on comparatively large carcasses which
remain exposed on the soil surface. No parental care of larvae is known. In the second, adults of
the genus Nicrophorus feed at large and exposed carcasses, but they must also secure a
comparatively small carcass and bury it for reproduction and subsequent larval maturation.
Adults remain with the developing larvae and care for them until they pupate. The Oriental
genus Ptomascopus exhibits behavior combining aspects of the life histories of both Silphinae
and Nicrophorus , but does not exhibit the parental care of larvae typical of Nicrophorus (Peck,
1982). These differing methods of carrion use are also found in Palearctic Silphidae.

Detailed studies have not been made on the Neotropical species, but there is no reason to
suspect that their feeding and reproductive behaviors differ. Our field observations indicate
that all species of Oxelytrum behave as typical Silphinae in feeding and breeding primarily on
large carcasses. Species of Oxelytrum differ from most Nearctic and Palaearctic silphines in
that most are nocturnal instead of being diurnal.

Unfortunately, natural history data are few for Latin American species of Nicrophorus but
they indicate that at least some of the species are nocturnal. There is no reason to suspect that
Latin American Nicrophorus differ in other aspects of their natural history from Holarctic
Nicrophorus. Rearings have not been attempted, and larvae are unknown for most species.

Anderson (1982a, 1982b) and Anderson and Peck (1985) review more detailed accounts of
the natural history of silphids.

METHODS

We do not include the Agyrtidae, previously considered as part of the Silphidae, but now
separated as a distinct family (Lawrence, 1982; Lawrence and Newton, 1982). The only
agyrtids known from Latin America are three Mexican species of Apteroloma (reviewed in
Bolivar and Hendrichs, 1972, who list them as Pterolomo ).

Full synonymies for North American genera and species are given in Peck and Miller (in
press). Type-species of genus-group names are in Madge (1980). Full synonymies are not given
here for species which also occur in the U.S. and Canada; they are given in Peck and Miller (in
press), and Anderson and Peck (1985). Full synonymies are listed only for those species limited
in their distribution to Latin America. All original literature has been checked unless otherwise
noted. We give the first use of a name or combination, and only references that contribute new

Quaest. Ent., 1985,21 (3)

250

Peck and Anderson

data. It is not our intention to give references to every use of a name in the older literature. We
do not cite “aberrations” but only usage of a name as a “variety”, because it may be interpreted
as having subspecific rank (International Code of Zoological Nomenclature, 1974, art. 45 (e)
(i)). For species that occur in the U.S. and Canada as well as Latin America, only the
synonyms pertaining to material from Latin America are given. Nomina nuda are not cited.
Such are listed by Portevin (1926), Hatch (1928), and Blackwelder (1944). Depositories are
indicated for type material we examined. In those instances where type material was not
examined we have indicated the probable depository according to Horn and Kahle (1937), and
noted this with a question mark.

Keys for identification of species north of Mexico were published by Miller and Peck(1979),
Peck (in press), and Anderson and Peck (1985) and should be consulted to confirm species
identifications of specimens from northern Mexico. Keys presented here include only species
known or likely to occur in Latin America. Keys for larvae are not presented because of lack of
species descriptions. Keys to larvae of some genera and species which occur in Latin America
are in Anderson and Peck (1985).

Species that may in the future be found to naturally occur in Latin America, but are not yet
recorded, are mentioned below. These are excluded from the detailed species discussions and
keys.

Nicrophorus americanus, N. carolinus, N. investigator , and TV. tomentosus and some
silphines of the southeastern United States may yet be found in Mexico because they occur in
bordering states to the north. A significant component of the biota of this region does occur in
temperate forests in northeastern Mexico (Martin and Harrell, 1957; Rosen, 1978). We have
seen one specimen of TV. sayi labelled “Mexico, N.L., Sierra de Gaucamayas, Zarogoza,
2-3. VII. 69, J.M. Matthieu, M.W. Sanderson, trampa de luz negra” (SBPC) but cannot accept
this single record for this far northern species as evidence of its occurrence in Mexico. Another
doubtful record is one of Necrophila americana labeled “Cuepayaca” (Cuernavaca) IX-46, H.
Field (FMNH).

Old records that we believe are erroneous, doubtful, or un-substantiated by recent specimens
are cited in Portevin (1926), Hatch (1928), and Blackwelder (1944). We do not discuss these
any further.

We have been unable to establish the identity of the names Nicrophorus quadricollis Gistel
from Mexico and Necrodes pronotus Gistel (1857:94) from Brazil. The types were supposedly
in the collections of the Zoological Museum in Munich (Horn and Kahle, 1937), but are now
considered lost (G. Scherer, in litteris 1984).

Distributions of all species are mapped based on personal examination of specimens.
Because of space limitations, full label data are not given but are available from the first
author. We cite only condensed locality (under State or Department or Province names in large
countries) and ecological data in alphabetical order, month (if on label) and number of
specimens if more than one. Specimen repository information or literature references follow the
records for each country. Obscure localities from Matthews (1888) in the Biologia Centrali
Americana were verified or located in Selander and Vaurie (1962).

All drawings were prepared with a camera lucida or an ocular grid and squared paper.
Measurements of length are from the anterior margin of pronotum to the elytral apex.

Phylogenies are reconstructed following Hennig (1966) and Wiley (1981). As do most
systematists, we adopt the biological species concepts as outlined by Mayr (1963). Since there
is no direct information available about reproductive isolation in Latin American Silphidae,

Carrion Beetles of Latin America

251

such isolation is inferred from differences in structural features, distribution, and available
information about natural history. We do not attempt to recognize subspecies. Adequate
population samples are not available to investigate the significance of variation in coloration
such as occur in some species of Oxelytrum and Nicrophorus.

MATERIALS

We have borrowed and examined material, totaling more than 4580 specimens, from the
following individuals and collections through the kindness of their owners or curators as follows:

AFNC Alfred F. Newton, Jr. Collection, Cambridge, Mass., U.S.A.

BMNH British Museum (Natural History), London; England; R.B. Madge.

CASC California Academy of Sciences, San Francisco, California, U.S.A.; D.H.
Kavanaugh.

CBMV Carlos Bordon Collection, Maracay, Venezuela; C. Bordon.

CMNH Carnegie Museum of Natural History, Pittsburg, Penn., U.S.A. ; G. Ekis
CNCI Canadian National Collection of Insects, Ottawa, Ont., Canada; A.

Smetana.

FMLC Fundacion M. Lillo, Tucuman, Argentina; R. Golbach.

FMNH Field Museum of Natural History, Chicago, 111., U.S.A.; H.S. Dybas.

FSCA Florida State Collection of Arthropods, Gainesville, Fla., U.S.A.; R.E.
Woodruff.

GMNH Geneva Museum of Natural History, Geneva, Switzerland; I. Lobl.

INPA Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil; N.D.

Penny.

ITMM Instituto Technologico de Monterrey, Mexico; Juan Contreras.

IZAV Instituto de Zoologia Agricola, Maracay, Venezuela; F. Fernandez- Yepez.

LACM Natural History Musuem of Los Angeles County, Los Angeles, California,

U.S.A.; C.L. Hogue.

LPMCN La Plata Museo de Ciencias Naturales, La Plata, Argentina; L. De Santis.

MCZC Musuem of Comparative Zoology, Harvard University, Cambridge, Mass.,

U.S.A.; A.F. Newton, Jr.

MHNM Museo de Historia Natural de la Ciudad de Mexico, Mexico City, Mexico;

P. Reyes-Castillo.

MNHN Museum National d’Histoire Naturelle, Paris, France; N. Berti.

MNSC Museo Nacional de Historia Natural, Santiago, Chile; G.A. Santic.

MZUSP Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, Brasil; C.

Costa.

OSCU Ohio State University, Department of Entomology, Columbus, Ohio,

U.S.A.; C.A. Triplehorn.

PURC Purdue University Entomology Collection, Lafayette, Ind., U.S.A.; R.D.

Waltz.

RDCC R.D. Cave Collection, Auburn, Ala., U.S.A.

RSAC Robert S. Anderson Collection, Edmonton, Alta., Canada.

SDMC San Diego Natural History Musuem, San Diego, Calif., U.S.A.; S.E.

Miller.

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252

Peck and Anderson

SBPC Stewart B. Peck Collection, Ottawa, Ont., Canada.

TMMC Texas Memorial Museum Collection, University of Texas, Austin, Tx.,

U.S.A.; J. Reddell.

UAIC University of Arizona Insect Collection, Tuscon, Ariz., U.S.A.; F.G.

Werner.

UFPB Universidade Federal do Parana, Curitiba, Parana, Brasil; D. Urban.

UICM University of Idaho, Department of Entomology, Moscow, Idaho, U.S.A.;

W.F. Barr.

USNM United States National Museum of Natural History, Smithsonian
Institution, Washington, D.C., U.S.A.; T.J. Spilman.

UTDZ University of Texas, Department of Zoology, Austin, Tx., U.S.A.; J.

Rawlings.

No fossil silphids are known from Latin America. Churcher (1966) tentatively reported
silphids among the insects found in the Talara late Pleistocene tar seeps of Peru. We examined
these fossils, deposited in the Royal Ontario Museum, Toronto, Ontario, Canada, and found
that they belong to other beetle families.

SYSTEMATICS

Key to Adults of Latin American Genera

1 Antenna clavate, the antennomeres gradually widening into an apical club

(fig. 1); fronto-clypeal suture absent (fig. 3); abdominal tergum V lacking
mid-dorsal stridulatory files: subfamily Silphinae

1' Antenna with the apical four antennomeres forming an abrupt club (fig.

2); fronto-clypeal suture present (fig. 4); abdominal tergum V bearing a
pair of mid-dorsal stridulatory files (hidden in many specimens by apices of

truncate elytra): subfamily Nicrophorinae

Nicrophorus Fabricius, p. 265

2 (1) Elytra with at least some reddish markings (fig. 5); pronotal postcoxal lobe

short and broadly rounded (fig. 6) Necrodes Leach, p. 253

2' Elytra wholly black; pronotal postcoxal lobe large, prominent (fig. 7)

3 (2') Elytra with ramose or branching sculpturing (fig. 9)

Heterosilpha Portevin, p. 253
3' Elytra lacking ramose or branching sculpturing

4 (3') Pronotum with two or four pairs of low, broad longitudinal costae on disc

(fig. 1); widest at or before middle, lacking hairs on dorsal surface

Oxelytrum Gistel, p. 257

4' Pronotum lacking costae; widest behind middle, bearing at least some hairs

on dorsal surface

5 (4') Head with a short row of long, erect hairs behind the eyes (fig. 11); elytra

tricostate Oiceoptoma Leach, p. 254

5' Head lacking a short row of erect hairs behind the eyes; elytra either

lacking costae entirely (fig. 12) or with numerous tubercles on dorsal
surface (Figs. 13, 14) Thanatophilus Leach, p. 255

Carrion Beetles of Latin America

253

SUBFAMILY SILPHINAE
NECRODES LEACH 1815

Four species are known in this genus, one of which occurs in North America where it is
widespread (Anderson and Peck, 1985; Ratcliffe, 1972). Adults are easily recognized by the
key characters, large size and form of elytra (Fig. 5).

Necrodes surinamensis (Fabricius)

Figures 5, 6, 17

Silpha surinamensis Fabricius, 1775: 72. Type locality: “ in America meridianali." . Syntypes: in Hunterian collection.

University of Glasgow (Ratcliffe, 1972). Ratcliffe, 1972; Anderson and Peck, 1985.

Diagnosis. — Length 15-25 mm. Eyes large, separated by distance about twice width of an
eye in dorsal view. Apical three antennomeres black. Pronotum black, sparsely punctate;
orbicular, widest near middle (Fig. 17). Pronotal postcoxal lobe short, broadly rounded (fig. 6).
Elytra tricostate, black with red markings present in apical quarter (fig. 5), some specimens
also with red markings along lateral margin near midlength. Some males with hind femora
greatly expanded.

Distribution. — Because TV. surinamensis occurs in counties bordering the Rio Grande in
Texas, it most likely occurs in bordering Mexico (Ratcliffe, 1972). The species name suggests
that it occurs in South America. We have seen a single specimen (MCZC) labeled “Ucayale
P(eru?). Maranon R., C. Sarkady” (which seemingly means where the Ucayale and Maranon
Rivers meet). In the absence of any other verifiable records, and our inability to find the species
in extensive collecting in Latin America, we cannot now accept the presence of the species any
farther south than possibly northern Mexico.

Ratcliffe (1972) has reviewed the natural history and distribution of the species in the
United States.

HETEROSILPHA PORTEVIN 1926

Two species of this endemic North American genus are known. Among Latin American
silphids, they are easily recognized by the ramose or branching sculpturing on the elytra.

Key to species

1 Males with pro- and mesotarsomeres 1-4 broadly expanded and with

elytral apex not prolonged (fig. 9); females with elytral apex somewhat
prolonged (fig. 8); male genitalia thicker and broader, parameres with
apices thicker and down-curved (figs. 18, 19); elytra without metallic lustre

H. ramosa (Say), p. 254

V Males with pro- and mesotarsomeres 1-4 not expanded; male and female

elytral apices similar, not prolonged (fig. 10); male genitalia more thin and
slender, parameres more narrow and straight (figs. 20, 21); elytra of some
specimens with metallic lustre H. aenescens (Casey), p. 254

Quaest. Em.. 1985, 21 (3)

254

Peck and Anderson

Heterosilpha ramosa (Say)

Figures 8, 9, 18, 19; Map 1

Silpha ramosa Say, 1823: 193. Type locality: “the upper Missouri (River)”. Neotype: in MCZC (designated by Miller and
Peck, 1979: 93). Miller and Peck, 1979; Anderson and Peck, 1985.

Diagnosis. — Length 14-18 mm. Eyes small, separated by distance about four times width
of an eye in dorsal view. Apical three antennomeres black. Pronotum black, uniformly and
densely punctate; transverse, widest near base. Pronotal postcoxal lobe large, rounded at apex.
Elytron tricostate, wholly black, with well developed branching sculpturing (fig. 9). Males with
pro- and mesotarsomeres 1-4 laterally expanded, densely pubescent beneath; females with
elytral apices slightly prolonged (fig. 8). Male genital characters as in key (figs. 18, 19).

Distribution. — The species is widespread throughout much of western North America
(Anderson and Peck, 1985). Its natural history in Colorado is described by Brewer and Bacon
(1975). Matthews (1888: 95) reports one specimen of the species from an unspecified site in
northern Sonora. Horn (1894) lists it from San Pedro Martir, Baja California. We have seen
the following record: Mexico. Baja California. Tijuana, III, 3. FMNH.

Heterosilpha aenescens (Casey)

Figures 10, 20, 21

Silpha aenescens Casey, 1886: 171. Type locality: San Francisco, California. Lectotype: in USNM (designated by Miller
and Peck, 1979: 93). Miller and Peck, 1979.

Heterosilpha aenescens (Casey), Portevin, 1926: 85.

Diagnosis.-— Length 14-18 mm. Eyes small, separated by distance of about four times
width of an eye in dorsal view. Apical three antennomeres black. Pronotum black, uniformly
and densely punctate; transverse, widest near base. Pronotal postcoxal lobe large, rounded at
apex. Elytron tricostate, wholly black but many specimens with a metallic lustre, with well
developed branching sculpturing. Males with pro- and mesotarsomeres 1-4 not expanded, not
densely pubescent beneath; male and female with elytral apices similar, not prolonged in
female. Male genital characters as in key (figs. 20, 21).

Distribution. — The species is known from southern to northern coastal California and
southern Oregon (Miller and Peck, 1979), and may occur in northwestern Mexico. We have
seen records from as far south as San Diego, California and suspect it occurs in northern Baja
California, Mexico.

OICEOPTOMA LEACH 1815

Three species of the Holarctic genus Oiceoptoma are known from North America, one of
which may enter extreme northeastern Mexico. Six species in this genus are known from the
Palearctic region.

Oiceoptoma rugulosum (Portevin)

Figures 11, 16, 23

Silpha inaequalis rugulosa Portevin, 1903: 333. Type locality: Savannah, Georgia. Type: in MNHN?, not seen.

Diagnosis Length 13-15 mm. Head with short row of long, erect hairs behind eyes (fig.
11). Eyes small, separated by distance about four to five times width of eye in dorsal view.
Apical three antennomeres black. Pronotum black, uniformly moderately densely punctate;

Carrion Beetles of Latin America

255

with few blackish hairs; transverse, widest near base. Pronotal postcoxal lobe broad, with
right-angled apical point. Elytron black, tricostate; some specimens with elytral intervals with
transverse rugose sculpturing. Elytral humeri each with a single tooth (fig. 16). Females with
elytral apices prolonged to sharp points; males with apices broadly rounded.

Taxonomic status and distribution. — The species has usually been considered conspecific
with 0. inaequale, which is widespread in both eastern Canada and the United States
(Anderson and Peck, 1985). Adults of the two taxa are separated most readily by those of 0.
rugulosum having a narrow elytral epipleuron (on the posterior half the upper oblique part is
subequal to the lower vertical part, fig. 23) and by those of 0. inaequale having a wide elytral
epipleuron (on the posterior half the upper oblique part is at least twice the width of the lower
vertical part, fig. 22). We know 0. rugulosum to occur from Florida to Indiana to Texas. It is
probably active in winter or spring. Portevin (1903: 333; 1926) cites the species from Mexico,
which is possible, but records are not known to us, and from Guiana, which is an obvious error.

THAN A TOPHILUS LEACH 1815

Six species of this widespread genus are known to occur in North America. Three of these
occur in Latin America, from central to Northern Mexico. Although primarily a northern
cold-adapted taxon, some species of Thanatophilus occur in southern desert grasslands and
shrublands. Other than two species which occur in the grasslands of southern Africa, members
of the genus are Holarctic in distribution. Where they occur at more southerly latitudes they
usually do so at higher elevations.

Key to species

1 Elytra abruptly truncate, lacking costae (fig. 12)

T. truncatus (Say), p. 255

1' Elytra not abruptly truncate; tricostate and with tubercles interspersed

between the costae (figs. 13, 14) 2

2 (T) Head and pronotum with abundant, long yellow-grey hairs; metasternal

hairs yellow-grey; posterior margin of abdominal sternum VII of female

unornamented, with marginal hairs only (fig. 24)

T. lapponicus (Herbst), p. 256

2' Head and pronotum with shorter and darker hairs which do not entirely

obscure the basal sculpture; metasternal hairs brown; posterior margin of
abdominal sternum VII of female with numerous coarse crenulations or
tooth-like projections (figs. 25a, 25b) T. graniger (Chevrolat), p. 256

Thanatophilus truncatus (Say)

Figure 12; Map 2

Silpha truncata Say, 1823: 193. Type locality: eastern Colorado. Neotype: in MCZC (designated by Peck and Miller.

1982: 154). Matthews, 1888:95.

Philas truncata (Say), Portevin, 1903: 331.

Diagnosis. — Length 11-14 mm. Eyes small, separated by distance four to five times width
of eye in dorsal view. Apical three antennomeres black. Pronotum black, uniformly and densly
punctate; with short appressed blackish hairs over entire surface; transverse, widest near base.

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256

Peck and Anderson

Pronotal postcoxal lobe large, rounded at apex. Mesosternal hairs black. Elytra black, lacking
costae, apices in both sexes abruptly truncate (fig. 12).

Natural history and distribution. — The species occurs in arid and open habitats in the
southwestern United States and extends in such habitats onto the Mexican Plateau. Adults are
known to be active from June to September. We have seen 76 specimens representing the
following records:

MEXICO. Chihuahua. Pinos Altos. Santa Clara. Coahuila. Canon del Fuenteno, Sierra de la Madera, VII. Districto
Federal. Mexico City. Durango. Durango. El Salto, 2450-2750m, VI, 27. 32 km E El Salto, 2450 m, IX, 2. Pedicena (not
located), VI, 8. Guerrero. Chilpancingo. Michoacan. Rio Balsas (Rio Mescales). Tancitaro. Nayarit. Jesus Maria, VII, 7.
La Mesa, VII, 3. Puebla. Puebla. Veracruz. Jalapa. Zacatecas. 40 km W Fresnillo, 2400 m, VI, 9. BMNH, CNCI,
FMNH, SBPC, USNM.

Horn (1895: 227) mentions a record from “Sierra San Lazaro”, Baja California. We do not
know this locality. The specimen was probably destroyed in the 1905 San Francisco earthquake
and fire.

Thanatophilus lapponicus (Herbst)
Figures 13, 24; Map 2

Silpha lapponica Herbst, 1793: 209, plate 52; Fig. 4. Type locality: Lappland. Type: in Berlin?, not seen. Anderson and

Peck, 1985. Not the species listed by Matthews, 1888: 96.

Thanatophilus lapponicus (Herbst), Portevin, 1926: 33.

Diagnosis .— Length 10-14 mm. Eyes small, separated by distance of four to five times
width of eye in dorsal view. Apical three antennomeres black. Pronotum black, uniformly and
densly punctate; with long yellow-grey hairs variably distributed over surface; transverse,
widest near base. Pronotal postcoxal lobe large, rounded at apex. Mesosternal hairs
yellow-grey. Elytra shorter, black, tricostate, with numerous tubercules interspersed between
costae (fig. 13). Females with elytral apices prolonged and rounded (fig. 13); males with apices
rounded, but not prolonged. Female with posterior margin of abdominal sternum VII
unornamented, with marginal hairs only (fig. 24).

Natural history and distribution. — The species is widespread in North America, especially
at higher altitudes or latitudes (Anderson and Peck, 1985). Portevin (1926: 136) states that T.
californicus Mannerheim, a synonym of T. lapponicus , is distributed from California, through
Central America, and along the Andes to Bolivia. This is an error. Records of this species in
Matthews (1888) refer to T. graniger.

A single Mexican record is known to us: MEXICO; Baja California. Tijuana, III, 5. FMNH.

Thanatophilus graniger (Chevrolat)

Figures 7, 14, 25a, 25b; Map 1

Oiceoptoma granigera Chevrolat, 1 833: 1 . Type locality: Mexico. Type: in MNHN?, not seen.

Silpha lapponica Herbst, misidentification of Matthews, 1888: 96.

Diagnosis.— Length 10-14 mm. Eyes small, separated by distance of four to five times
width of eye in dorsal view. Apical three antennomeres black. Pronotum black, uniformly and
densely punctate; with short yellow-grey hairs variably distributed over surface; transverse,
widest near base. Pronotal postcoxal lobe large, rounded at apex (fig. 7). Mesosternal hairs
dark brown. Elytra longer, black, tricostate, with numerous tubercules interspersed between
costae (fig. 14). Females with elytral apices prolonged and rounded (Fig. 14); males with apices
rounded but not prolonged. Females with posterior margin of abdominal sternum VII with
numerous coarse crenulations or tooth-like projections, nearly as long as marginal hairs (figs.

Carrion Beetles of Latin America

257

25a, 25b).

Taxonomic notes, natural history and distribution. — This species has usually been
considered conspecific with T. lapponicus, however, the female abdominal character clearly
separates the two as distinct taxa.

The species is known only from the higher elevations of the Transverse Volcanic Sierra of
Mexico, and Cerro Potosi in northeastern Mexico. Adults were collected during February,
May, June, and November.

We have seen 38 specimens representing the following records:

MEXICO. Districto Federal. Mexico City. Hidalgo. Guerrero Mills, 5. Apan, V, 1. Jalisco. Sayula. Mexico. Salazar,
3000 m, IX, 6. Morelos. Km 50 Mexico to Cuernavaca Road, VIII, 2. Nuevo Leon. Galeana, Cerro Potosi, 3750 m, VI, 16.
Puebla. Ciudad Serdan (San Andres Cholchicomula). Vera Cruz. Jalapa. Las Vigas. No locality, II. No data, 2. BMNH,
ITMM, MCZC, MHNM, SBPC, USNM.

OXELYTRUM GISTEL 1848

Several generic and subgeneric names have been used for the species of Oxelytrum. We are
unable to find enough characters which combine to form a consistent suite justifying
recognition of more than a single genus.

Adults of this genus are easily recognized by tricostate elytra, lacking ramose sculpturing, a
pronotal disc lacking hairs and bearing two or four low longitudinal costae (fig. 1), and a large
pronotal postcoxal lobe. We recognize eight species in the genus, most of which are confined to
South America. Only the very widespread O. discicolle enters extreme southern Texas.

Key to species

1 Eyes small, not prominently protruding, separated by distance distinctly
greater than three times width of an eye in dorsal view (figs. 26a, 26b)

Y Eyes large, prominently protruding, separated by distance about three

times or less than width of an eye in dorsal view (figs. 1,3)

2 (1) Pronotum with quadrangular reddish spot in posterolateral corner;

pronotum and elytra with vague blue-green iridescence

O. biguttatum (Philippi), p. 258

2' Pronotum wholly black; pronotum and elytra lacking trace of iridescence

3 (2') Apical three antennomeres orange-yellow O. apicale (Brulle), p. 258

3' Antenna black O. anticola (Guerin-Meneville), p. 259

4 (1') Pronotum and elytra concolorous chestnut brown to black; apical

antennomere orange

O. lineatocolle (Laporte), p. 260
4' Pronotum with margins orange-yellow, disc mostly or partly blackish (fig.

1 ); antennae various in color

5 (4') Elytral humeri rounded (fig. 27); pronotum with costae distinctly elevated

5' Elytral humeri toothed (fig. 28); pronotum with costae present but

indistinct

6 (5) Apical antennomere orange-yellow; elytra with apices emarginate, sutural

angles sharp (Fig. 32) O. emarginatum (Portevin), p. 260

6' Apical antennomere black; elytra with apices not emarginate, the sutural

angles evenly rounded or only slightly prolonged

O. erythrurum (Blanchard), p. 261

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258

Peck and Anderson

7 (5') Pronotum with postcoxal lobe wholly orange-yellow; pronotum with dark

coloration confined to very middle of disc; elytral apices produced and

blunt (Figs. 35, 36) O. cayennese (Sturm), p. 262

7 7 Pronotum with postcoxal lobe wholly black; pronotum with dark coloration

more extensive (fig. 1); elytral apices sharply pointed (figs. 37, 38)

O. discicolle (Brulle), p. 263

Oxelytrum biguttatum (Philippi), NEW COMBINATION
Figures 26b, 29; Map 3

Necrodes biguttatus Philippi, 1859: 664. Type locality: Chile. Type in: MNSC; syntypes no. 2219-2220, not
seen.

Silpha biguttula Fairmaire and Germain, 1859: 350. Type locality: Straits of Magellan. Type in: MNHN?,
not seen.

Necrodes biguttulus (Fairmaire and Germain), Fairmaire, 1888: 27.

Hyponecrodes biguttatus (Philippi), Berg, 1901: 327. Schouteden, 1905: 199.

Paranecrodes biguttatus (Philippi), Portevin, 1921: 81. 1926: 131.

Silpha ( Paranecrodes ) biguttata (Philippi), Hatch, 1928: 116.

Diagnosis. — Length 12-19 mm. Head with eyes not prominent, separated by distance five
times width of an eye in dorsal view (fig. 26b); frontal depressions absent; occipital-frontal crest
acute. Antennae black. Pronotum black with vague blue-green iridescence, with quadrangular
reddish spot in each posterolateral corner; transverse, about 0.6 times as long as wide; posterior
angles obtusely angulate; margins not reflexed upwards; pronotal costae present but effaced.
Pronotal postcoxal lobe black. Elytra black with blue-green iridescence, elytral humeri not
toothed. Abdomen of both sexes with segment VII and apical portion of segment VIII
orange-red, otherwise black. Males with elytral apices abruptly rounded (fig. 29); in females,
slightly more prolonged and evenly rounded.

Natural history and distribution. — The species occurs in forests and open habitats in the
southern half of Chile and adjacent Argentina. Adults were collected from October to April.

We have seen 59 specimens representing the following records:

ARGENTINA. Chubut. No data, 1. No locality, XI. Neuqen. Nahuel Huapi, 2. Neuquen, III, IV, 4. Pucara, Parque
Nacional Lanin, XII, 2. Rio Negro. Bariloche. Gutierrez, XI. Santa Cruz. Lago Argentino. Lago Blanco. Valle Tunel (not
located), 2. Ventisquero Moreno, Los Glacieres, I. No data, 1. Tierra del Fuego. Bahia San Sebastian, Cerrillos, IV. Rio
Grande. Rio McClelland (not located). San Sebastian. Ushuaia, II. No Locality, I. No data, 2. BMNH, FMLC, FMNH,
LPMCN, USNM.

CHILE. Aisen. Golfo de Penas, I, 5. Laguna San Rafael, Taitao, X, XII. Puerto Cisnes, II. Chiloe. Palena.
Llanquihue. Frutillar, I. Malleco. 20 km E Manzanar, 1100m, XII, 12. Termas de Tolguaca, II. Magallanes. Dawson
Island. Esperanza (not located), I. Isla Navarina, XI. Isla Riesco, Mina Elena, II. Puerto Eden, XII. Punta Arenas, II.
Useless Bay. No data, 1. Osorno. Parque Nac. Puyehue, Antillanca Rd., 965 m, Nothofagus forest, XII, 3. Valdivia.
Corral. Enco, III. AFNC, BMNH, CNCI, MCZC, MNSC, SBPC, USNM.

Oxelytrum apicale (Brulle), NEW COMBINATION
Figures 26a, 30a, 30b; Map 4

Silpha apicalis Brulle, 1840: 74, in Brulle and Blanchard, 1840. Type locality: Potosi, Bolivia. Type in: MNHN?, not
seen.

Hyponecrodes apicalis (Brulle), Kraatz, 1876: 375.

Hyponecrodes (Katanecrodes) apicalis (Brulle), Portevin, 1921: 82.

Silpha ( Katanecrodes ) apicalis (Brulle), Hatch, 1928: 113.

Diagnosis Length 9-11 mm. Head with eyes not prominent, separated by distance
about five times width of an eye in dorsal view (fig. 26a); frontal depressions moderately deep;
occipital-frontal crest obtuse. Apical three antennomeres orange red. Pronotum black;

Carrion Beetles of Latin America

259

transverse, about 0.75 times as long as wide; posterior angles broadly rounded; margins not
reflexed upwards; pronotal costae distinctly elevated. Pronotal postcoxal lobe black. Elytra
black; middle costa of some specimens effaced at basal one third; elytral humeri not toothed.
Abdomen of both sexes black except for yellow-orange segment VIII in females. Males with
elytral apices abruptly rounded (fig. 30a); in females very slightly prolonged immediately
lateral to sutural apex and more evenly rounded (fig. 30b).

Natural history and distribution. — The species is known to us only from high and low
elevation open habitats in northwestern Argentina and adjacent Bolivia. Adults were collected

from November to April. We have seen 107 specimens representing the following records:

ARGENTINA. Catamarca. Catamarca, II. El Manchado, 3000 m, I; 4000 m, I. El Suncho Experimental Station, II,
6. Famabulasto (not located). La Cienaga, 3700 m, III, 2. Las Mansas (not located). III. Los Angeles, II. Nevados del
Aconquija, Quebrada de los Cazedores, 4500 m, XI; between Ingehio and Puesto de los Ojos (not located), II. Santa Maria
Puesto de los Ojos (not located), II, 5. Cordoba. Cordoba, 4. Pampa de Achala, II. Jujuy. Abra Pampa, III. La Rioja.
Velasco, II. Mina da Esperanza (not located), II, 5. Tucuman. Amaicha, 2000 m, II; Quebrada Amaicha, IX. Between El
Nagalar and Santa Maria, III. Infiernillo, XI, 12; XII, 33; Quebrada Honda, 3400 m. San Jose, 2500 m, IV, 2. Siambon,
III. Tafi del Valle, II, 3; III, 2; XI, 8; no date, 7. Tafi Viejo, II. Trancas, San Pedro Colalao, II. No locality, 3. FMLC,
GMNH, LPMCN, MZUSP.

BOLIVIA. Pongo de Quime, VI. USNM.

Oxelytrum anticola Guerin-Meneville), NEW COMBINATION
Figure 31; Map 5

Silpha anticola Guerin-Meneville, 1855: 592. Type locality: Ecuador. Type in: Brussels Museum?, not seen.

Hyponecrodes anticola (Guerin-Meneville), Kraatz, 1876: 375.

Silpha microps Sharp, 1891: 40. NEW SYNONYMY. Type locality: Quito, Ecuador, 2895 m (9500 feet). Type in
BMNH, seen.

Hyponecrodes (Katanecrodes) andicola (Guerin-Meneville), Portevin, 1921: 82.

Silpha (Katanecrodes) anticola (Guerin-Meneville), Hatch, 1928: 1 14.

Silpha ( Katanecrodes ) microps (Sharp), Hatch, 1928: 114.

Diagnosis. — Length 9-11 mm. Head with eyes not prominent, separated by distance
about five times width of an eye in dorsal view; frontal depressions moderately deep;
occipital-frontal crest obtuse. Antennae black. Pronotum black; transverse, about 0.75 times as
long as wide; posterior angles broadly rounded; margins not reflexed upwards; pronotal costae
distinctly elevated. Pronotal postcoxal lobe black. Elytra black; middle costa of some specimens
effaced at basal one-third; elytral humeri not toothed. Abdomen of both sexes black except for
yellow-orange segment VIII in females. Males with elytral apices abruptly rounded (fig. 31);
female apices very slightly prolonged immediately lateral to sutural apex and more evenly
rounded.

Notes about synonymy. — Silpha microps Sharp is placed in synonymy with O. anticola
because we find no features on the type which separate it from the description or specimens of
Oxelytrum anticola from the same general locality.

Natural history and distribution. — The species is known mostly from high elevation open
habitats in the Andean countries of Ecuador, Peru, and Bolivia. Portevin (1926) cites the
species from Colombia, which is possible, but we know of no records. Adults were collected
during the months of December through April and in July. We have seen 17 specimens
representing the following records:

BOLIVIA. La Paz, II; El Alto, 4100 m, XII; no date, 2. Oruro, 3700 m. BMNH, USNM

ECUADOR. Latacunga, I. Machachi, VII. Quito (at Miami in aircraft, quarantine intercept), VII. 16 km N. Quito.
Mitad del Mundo, III. PURC, USNM.

PERU. Cajacey, 2650 m, IV. Carumas, 2200 m, IV. Chiquata, near Arequipa, 3100 m, II. Hlancayo (not located),
III. Otoyo (not located), 4000 m. Tacana Libra (Totora)(not located), 2. BMNH, FMLC. FMNH, LPMCN, MCZC,
USNM.

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Oxelytrum lineatocolle (Laporte)
Figures 33, 34; Map 6

Silpha lineatocollis Laporte, (Comte de Castelnau) 1840: 5. Type locality: Chile. Type in: MNHN?, not seen. Fairmaire
and Germain, 1859: 350.

Necrodes gayi Solier, 1849: 359: Type locality: Chile. Type in: MNHN?, not seen.

Hyponecrodes lineatocollis (Laporte), Kraatz, 1876: 375.

Hyponecrodes lineaticollis (Laporte), Berg, 1901: 329.

Hyponecrodes (Hyponecrodes) lineatocollis (Laporte), Portevin, 1921: 84.

Silpha ( Oxelytrum ) lineaticollis (Laporte), Hatch, 1928: 115.

Diagnosis. — Length 15-22 mm. Head with eyes prominent, separated by distance about
twice width of an eye in dorsal view; frontal depressions present but shallow; occipital-frontal
crest acute. Antennae black. Pronotum wholly black to dark brown; transverse, about 0.65
times as long as wide; posterior angles obtusely angulate; margins not or else very slightly
reflexed upwards; pronotal costae elevated. Pronotal postcoxal lobe black. Elytra black to dark
brown; costae continuous throughout; elytral humeri not toothed. Abdomen of both sexes black
except for yellow-orange anterior portion of segment VIII. Males with elytral apices abruptly
rounded (fig. 33); apices in female slightly more prolonged and more evenly rounded (fig. 34).

Natural history and distribution. — The species is known from central Chile and adjacent
Argentina. Adults have been collected from October to July, and are associated with both open

and forested habitats. We have seen 358 specimens representing the following records:

ARGENTINA. Neuquen. Pampa Central, III. Patagonia. No other data. “NW Patagonia,” 300-900 m. Rio Negro.
El Bolson, III. BMNH, LPMCN, LACM.

CHILE. Arauco. Caramavida, X. Cordillera Nahuelbuta, XII, 4. Cautin. Bellavista, Lago Villarrica, 310 m,
Valdivian rainforest, XII, 13. 3 km NE Token, 3 m, II, 4. Volcan Villarrica, Nothofagus forest, 1 120 m, XII, 23; 1250 m,
XII, 4. Chiloe. Chiloe Island. Concepcion. Ejido Pinares, XII. Concepcion, III, 3; IV; V, 4. 6 km S San Pedro, 360 m,
Pinus forest, XII, 101. Curico. El Coigo, III. Llanquihue. Chamiza, III. Lago Chapo, 11.7 km E Correntoso, 320 m,
Valdivian rainforest, XII, 58; 13.5 km e Correntoso, 310 m, Valdivian rainforest, XII, 24. Maullin, I. Petrohue, 600 m, I,
2. Malleco. Malalcahuello, 6.5 km E, 1080 m, Nothofagus forest, XII, 8; 14 km E, 1570 m, forest, XII. Manzanar, 1 100
m, XII. 15 km W Victoria, 200 m, XII, 2. Maule. No locality, 1. El Pantanillo, 17 km SE Constitucion, 250 m, XI. Nuble.
Alto Trequalemu, 500 m, 20 km SE Chovellen, I; XII, 3. Las Trancas, 1400 m, 70 km E Chilian, XII. Termas de Chilian,
I. 3 km NE Tolten, 3 m, II, 4. 17.5 km S Curanipe, 50 m, I, 2. Osorno. Parque Nacional Puyehue, 600 m, Aguas
Calientes, II, 9; Antillanca Rd., 720 m, Nothofagus forest, XII, 3; 4.1 km NE Anticura, 430 m, Valdivian rainforest, XII,
16. 15 km W Termas de Puyehue, Rancho Mirado, 205 m, II. 7.7 km NE Termas de Puyehue, 200 m, Valdivian
rainforest, XII, 206. No locality, 6. Santiago. Leyda, VI. Santiago. Tarapaca. Mocha (doubtful record). Valdivia. 4.1 km
W Anticura, 270 m, Valdivian rainforest, XII, 44. Enco, 120 m. III. Valdivia, X, 5; XII; no date, 5. Valparaiso. Quillota,
VII, 2. Valparaiso, 6. AFNC, BMNH, CMNH, CNCI, FMNH, GMNH, LPMCN, MNSL, MZUSP, SBPC, USNM.

Oxelytrum emarginatum (Portevin)

Figure 3; Map 7

Hyponecrodes emarginatus Portevin, 1920b: 506. Type locality: Brazil. Type in: MNHN, seen.

Hyponecrodes ( Hyponecrodes ) emarginatus Portevin, Portevin, 1921: 83.

Silpha (Oxelytrum) emarginata (Portevin), Hatch, 1928: 1 14.

Diagnosis. — Length 14-18 mm. Head with eyes prominent, separated by a distance of
about three times the width of an eye in dorsal view; frontal depressions present, shallow;
occipital-frontal crest acute. Apical antennomere orange. Pronotum with margins orange-red,
disc black; transverse, from 0.60-0.65 times as long as wide; posterior angles obtusely angulate;
margins reflexed upwards in most specimens; pronotal costae elevated. Pronotal postcoxal lobe
black. Elytra black; inner two costae effaced near base, middle costa effaced at basal one-third
in some specimens; elytral humeri not toothed. Abdomen of males and females with segment IX
and apical portion of segment VIII orange-red. Males and females with elytral hind angles
abruptly rounded, apex emarginate, the sutural apices prolonged to needle-like points (fig. 32).

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261

Variation. — One female from the Boraceia Biology Station, Salesopolis, Brazil, (MZUSP),
has distinctly emarginate elytral apices but only a vaguely orange-red apical antennal segment.

Natural history and distribution. — The species is known to us only from the coastal ranges
and highlands of southeastern Brazil. We have seen 1 1 specimens representing the following
records:

BRAZIL. Minas Gerais. Sapucai-Mirim, Cidade Azul, 1400 m. Virginia, Faz Campos, 1500 m. Rio de Janeiro.
Itatiaia. Rio de Janeiro. Sfto Paulo. Boraceia, 850 m, X, 3; XI. Pindamonhangaba. Salesopolis, VI. Viradouro. CNCI,
MNHN, MZUSP, USNM.

Oxelytrum erythrurum (Blanchard)
Map 7

Silpha erythrura Blanchard, 1840: 75, in Brulle and Blanchard, 1840. Type Locality: Montevideo, Uruguay. Type in:
MNHN, seen.

Hyponecrodes erythrurus (Blachard), Kraatz, 1876: 376.

Hyponecrodes erythrura (Blanchard), Berg, 1901: 328.

Hyponecrodes ( Hyponecrodes ) erythrurus (Blanchard), Portevin, 1921: 85.

Hyponecrodes erythrurus var. melanurus Portevin, 1926: 129. Type locality: not given. Type in: MNHN?, not seen.
Hyponecrodes (Hyponecrodes) erythrurus var. pygialis Portevin, 1921: 83. Type locality: not given. Type in: MNHN?,
not seen.

Hyponecrodes (Hyponecrodes) erythrurus var. melancholicus Portevin, 1921: 83. Type locality: not given. Type in:
MNHN?, not seen.

Silpha ( Oxelytrum ) erythrura (Blanchard), Hatch, 1928: 114.

Hyponecrodes (Hyponecrodes) opacus Portevin, 1921: 83. NEW SYNONYMY. Type locality: Bolivia. Type in: MNHN,
seen.

Hyponecrodes (Hyponecrodes) opacus var. tristis Portevin, 1921: 83. Type locality: not given. Type in: MNHN?, not seen.
Silpha ( Oxelytrum ) opaca (Portevin), Hatch, 1928: 1 14.

Diagnosis .— Length 10-19 mm. Head with e} s prominent, separated by distance about
three times width of an eye in dorsal view; frontal depressions present, shallow; occipital-frontal
crest acute. Antennae black. Pronotum with margins orange-red, disc black; transverse, about
0.65 times as long as wide; posterior angles obtusely angulate; margins slightly to strongly
relexed upwards in some specimens; pronotal costae elevated. Pronotal postcoxal lobe black.
Elytra black; inner two costae effaced at very bases, otherwose strongly raised and glabrous;
elytral humeri not toothed. Males with abdominal tergum VIII and IX with basal portion
orange-red to varying degree; abdominal sternum VIII entirely black. Females with abdominal
segment VIII entirely yellow-orange. Elytral apices broadly and evenly rounded in male;
slightly prolonged immediately laterally of sutural apices in female. Some males with
posteriorly directed lateral expansions of sternite of abdominal segments V and VI, but not
appearing as prominent spines as in O. cayennensis.

Variation and Synonymy. — The amount of orange coloration on abdominal tergum IX (in
males) varies. Portevin used this in establishing varieties. We also find coloration to be related
to sex of the specimens, and cannot find any coherent pattern of geographic variation. There is
also individual variation in the extent to which elytral costae are effaced. This was the main
character used to define Hyponecrodes opacus Portevin. Portevin (1921) described the species
from only Five specimens, supposedly from Brazil and Bolivia. Of these, we have seen only the
type, a female. The only distinctive feature we see in it is that it has the second elytral costa
completely effaced in the anterior half. It bears the following labels: a square hand written label
“Bolivie / Standiger”, a square machine printed label “Museum Paris / Coll. A. Grouvelle
1915”, a square machine printed label in red type “TYPE”, and an apparently recent square
label “H. opacus Port”. One specimen, labelled O. opacus , in the BMNH has narrower first
elytral costae in the basal one third than does O. erythrurum. Specimens of O. erythrurum

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from NW Argentina tend to have the second elytral costa more effaced. We conclude that the
type of O. opacum is within the range of variation we have seen in O. erythrurum, and since we
know of no other characters to separate it, consider the two names synonyms.

Natural history and distribution. — This species is widespread and apparently common in
northern Argentina, Bolivia, southern Brazil, Paraguay, southeastern Peru, and Uruguay. It
seemingly occurs in open and forested habitats. Adults have been collected in all months of the

southern summer. We have seen 230 specimens representing the following records:

ARGENTINA. Buenos Aires. Ajo. Bahia Blanca, XI. Buenos Aires, I, 2; VI, 3; XI; no date, 3. Lago de Gomez, Junin,
XII. Lago Monte, San Miguel del Monte, XII. Las Flores, Ciudad, X. La Plata, 23. Palermo, II; III, 4. Punta Lara, X.
Rincon de Ajo, 5. San Fernando, XI, 8. Tiore (not located), II. Vitel, N of Chascomus, XI. Catamarca. Andalgata. El
Manchado, 3000 m, I. El Rodeo, I. Los Hoyos Mesaga (not located), 1700 m, IV, 3. Cordoba. Alta Gracia, XI. Cordoba.
Corrientes. Corrientes, VII. Entre Rios. Gualequay. Primero de Mayo, X. Jujuy. Digue la Cienaga, III (not located).
Estero Uyto, II, 2. Sunchal, IX. Misiones. Iquazu, X, 3. No data, I. Salta. Capital, II. Coronel Moldes, II, 20. El
Corralito, 15 km S Campo Quijano, II, 9. Rio Blanco, Campo Quijano, I, 11. Rosario de Lerma, II, 7. San Antonio, XI.
San Lorenzo, I; XI. Santa Fe. Rosario, 4. Santa Fe, 3. Tucuman. Aguadita (not located), I, 5. Guabatal (not located), I, 2.
Infiernillo, III, 6. La Higuera, IV. Parque Aconguija, XII, 2. Rio Los Sosa, 900 m. III, 2. San Pedro de Colalao, I, 3.
Siambon, II, 3; III, 6; VII, 2; XII. Tacanas, I, 2; II; XII, 2. Tafi del Valle, II; XI, 16. Tafi, Taficillo, 1500 m, XI. Tucuman,
Ciudad Universitaria, II; 11. Tucuman, II, 2. BMNH, CASC, FMLC, FMNH, FSCA, GMNH, LPMCN, MZUSP,
SBPC, USNM.

BOLIVIA. La Paz, II. USNM.

BRAZIL. Mato Grosso. Maracaju. Parana. Curitiba, 900 m, XI. Serrinha Parana, XII, 4. Rio Grande do Sul.
Pelotas, X; XII, 2. Rio Grande. Santa Catarina. No data. Sho Paulo. Sao Paulo, 2. BMNH, CMNH, MCZC, MZUSP,
USNM.

PARAGUAY. Caaguazu. Paso Yobai (not located), X. Cordillera. Caacupe, X, Blacklight. FMLC, RDCC.

PERU. Junin. Valle Chanchamayo, 1400 m, IV. FMLC.

URUGUAY. Las Piedras, Canelones, II. Maldonado. Montevideo, II; XI; XII, 2; no date, 2. BMNH, FSCA,
GMNH, USNM.

Oxelytrum cayennense (Sturm)

Figures 28, 35, 36; Map 8

Silpha cayennensis Sturm, 1826: 61. Type locality: Cayenne, French Guiana. Type in: Munich?, not seen.

Hyponecrodes cayennensis (Sturm), Kraatz, 1876: 375.

Hyponecrodes ( Hyponecrodes ) cayennensis (Sturm), Portevin, 1921: 85.

Oxelytrum occidentale Gistel, 1848: 190. Type locality: Brazil. Type in: probably lost.

Oxelytrum aequinoctiale G istel, 1848: 190. Type locality: Brazil. Type in: probably lost. Madge, 1980: 357.

Silpha ( Oxelytrum ) cayennensis (Sturm), Hatch, 1928: 115.

Diagnosis. — Length 13-19 mm. Head with eyes prominent, separated by distance about
two times width of an eye in dorsal view; frontal depressions present, shallow; occipital-frontal
crest acute. Antennae black. Pronotum with margins and most of disc orange-red or yellow,
only very middle of disc black; transverse, about 0.6 - 0.65 times as long as wide; posterior
angles obtusely angulate; margins not reflexed upwards; pronotal costae present but effaced;
pronotal postcoxal lobe entirely orange yellow. Elytra black; inner two elytral costae partially to
almost completely effaced from midlength to basal one-third in most specimens, otherwise
elevated; elytral humeri each with a single tooth (fig. 28). Males and females with abdominal
segment VIII entirely orange-red; abdominal segment VII either black or with orange-red area
at middle of apical margin. Males with elytral apices obliquely truncate, sutural angle evenly
rounded (Fig. 35); apices in female prolonged and sinuate but not sharply pointed (fig. 36).
Some males with prominent posteriorly directed lateral expansions of abdominal sternites V
and VI which appear as large spines.

Variation. — Specimens vary in extent of orange color on abdominal segment VII and VIII.
This was used by Portevin to establish several aberrations. There is also variation in the extent
to which the elytral costae are effaced. We have not been able to observe a geographic pattern

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263

in this variation.

Natural history and distribution. — The species occurs over much of northern and central
South America. It is most frequently collected in lower to middle elevation rain forest habitats
and is active in all months. We have seen 268 specimens representing the following records:

BOLIVIA. Beni. Ivon, II. Rosario, Lago Rogagua, X. Santa Cruz. Buena Vista, Ichilo, II, 3; XII: no date, 2. Rio
Japacani, Santa Cruz de la Sierra, 450 m, 34. CASC, CMNH, MZUSP, USNM.

BRAZIL. Amapa. Rio Amapani, VII. Rio Amapai, VIII, 2. Rio Branco, Boa Vista, I. Serra do Navio, X, 5.
Amazonas. Manaus, INPA Station, VII; X. Maues, II, 3. Reserva Ducke, 26 km ex Manaus, I; III; V, 45. Rio Purus,
Hyutanaha. Mato Grosso. Reserva Humboldt, Bento Mascarenhas (not located). Serra do Norte (not located). III.
Xavantina, gallery forest. Minas Gerais. Bello Horizonte, IV. Miscosa (not locaated), XI. Para. Cachimbo. Rio de
Janeiro. Rio de Janeiro. Rondonia. V. Rondonia, 378 km S P. Velho, 387 km S P. Velho, I. SZo Paulo. Boraceia, Casa
Grande, I, 2. USNM, SBP, INPA, MZUSP, MCZC, UFPB.

COLOMBIA. Amazonas. Leticia, rainforest, II. Cundinamarca. Bogota, no date, 4. Norte de Santandar. 35 km S
Cucuta, Quebrada Honda, 700 m, V. 4. SBPC, USNM.

ECUADOR. Manabi. 78 km NE Chone, 450 m„ VI, 2. Napo. 12 km SW Tena, 500 m, VII, 21. Pastaza. Puyo, II, 8.
22 km SW Puyo, 900 m, VII, 5; 22 km W Puyo, II. Pichincha. Rio Palenque Station, 47 km S Santo Domingo, rainforest,

V, 7. Tinalandia, 16 km SE Santo Domingo, lower montane rainforest 680 m, II, 2; VI, 61. SBPC, USNM.

FRENCH GUIANA. Marioni River, Duserre. Mana River, V. CMNH, USNM.

GUYANA. Bartica. Essequibo River, Morabaldi Creek; Monkey Jump. Kartabo, VIII, 2. Membaro Creek, upper
Mazaruni River. Oronoque and New River Heads. BMNH, FMNH, MCZC.

PERU. Loreto. Estiron, Rio Ampiyachu, XI, 2. Junin. La Merced. Valle Chanchamayo, 800 m, I, 2. BMNH, FMLC,
FMNH.

VENEZUELA. Amazonas. Mt. Marahuaca, N slopes, V, 9. Peraitepuy, 4. Aragua. Rancho Grande, N of Maracay,

VI. Zulia. Kunana, Perija, 1 100 m, Rio Negro, XII, 8. IZAV, USNM.

Oxelytrum discicolle (Brulle)

Figures 1, 3, 15, 37, 38; Map 9

Silpha discicollis Brulle, 1840: 75, in Brulle and Blanchard, 1840. Type locality: Altamachi River, near Cochabamba,
Bolivia. Type in: MNHN?, not seen.

Hyponecrodes discicollis (Brulle), Portevin, 1905: 50.

Hyponecrodes ( Hyponecrodes ) discicollis (Brulle), Portevin, 1921: 85.

Silpha (Oxelytrum) discicollis (Brulle), Hatch, 1928: 115.

Necrodes analis Chevrolat, 1843: 26. Type locality: Orizaba, Mexico. Type in: MNHN?, not seen.

Hyponecrodes analis (Chevrolat), Kraatz, 1876: 376. Matthews, 1888: 95.

Hyponecrodes (Hyponecrodes) discicollis var. elongatus Portevin, 1921: 84. Type locality: not given. Type in: MNHN?,
not seen.

Hyponecrodes (Hyponecrodes) discicollis var. discretus Portevin, 1921: 84. Type locality: not given. Type in: MNHN?,
not seen.

Silpha (Oxelytrum) discicollis (Brulle), Hatch, 1928: 115.

Diagnosis. — Length 11-19 mm. Head with eyes prominent, separated by distance about
twice width of an eye in dorsal view (fig. 3); frontal depressions shallow; occipital-frontal crest
acute. Antennae black. Pronotum with margins orange-red, disc black; transverse, about
0.6-0.65 times as long as wide; posterior angles obtusely angulate; margins very slightly
reflexed upwards in some specimens; pronotal costae present but indistinct. Pronotal postcoxal
lobe black. Elytra black; inner two costae effaced at base in most specimens; middle costa
partially to completely effaced from midlength to basal one-third in most specimens, otherwise
costae elevated; elytral humeri each with a single tooth (fig. 15). Males and females with
abdominal segment VIII entirely orange; tergum of abdominal segment VII with orange-red
spot of variable size at apical margin, otherwise black. Males with elytral hind angles evenly
rounded, sutural angles very slightly prolonged to a sharp point (fig. 37); apices in females
slightly prolonged and with sutural spines very slightly longer than in males (fig. 38). Some
larger males with lateral margins of abdominal sterna V and VI very slightly produced
laterally.

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Variation. — The extent of the orange coloration of abdominal segment VII varies within
and between sexes of a single population. This variation was used by Portevin to establish
aberrations. There is also variation in the extent to which the elytral costae are effaced.

Natural history and distribution. — This is the most commonly collected species of silphid
in Latin America. Many adults are attracted to carrion baits, and come commonly at night to
ultraviolet and other light traps. The species is distributed from southern Brazil and Paraguay,
through much of central and northern South America (but not the lowlands of the Amazon
Basin), through Central America, to Mexico and extreme south Texas. Adults have been
collected in every month of the year, in habitats ranging from rainforest to montane cloud
forest from near sea level to more than 3000 m elevation, and in open semi-arid thorn-scrub

vegetation. We have seen 3096 specimens representing the following records:

ARGENTINA. Misiones. Cataracas de Iguazu, XI. Eldorado, XI, 2. Iguazu, III, X. FMLC, USNM.

BELIZE. Belmopan, 50 m, rainforest, VIII, 2. 6 km S. Belmopan, VIII, rainforest. SBPC.

BOLIVIA. Cochabamba. Alto Palmar, 800 m, X. Chapare, 1000 m. Cochabamba. Incachaca, 2500 m, 30. La Paz. La
Paz. Yan(aca)chi (?). Santa Cruz. Carahuasi, 250 km E Cochabamba, 3000 m, VII. Santa Cruz, 300 m. Parapeti, X, 4.
Yanchi. CMNH, FMLC, MZUSP, USNM.

BRAZIL. Amazonas. Tucano, 1500 m, IV, 2. Bahia. Salvador, VIII. Distrito Federal. Brasilia. Foresta da Tijuca,
VIII. Espiritu Santo. No data. Guarapari, 3. Minas Gerais. Bello Horizonte, IV. Lambari, XI. Laveras, III. Vicosa, III.
Serra do Caraca, XI, 13; XII, 8. Sapucai-mirim, Cda. de Azul, 1400 m, XI, 2. Parana. Banhados (Curitiba to Paranagua),
800 m, II, 34. Fozo do Iguazu, IX, 2; XII, 4. Guaixa (not located), XII. Jaguariaiva, I. Marumbi, II, VI. Quatro Barras,
III, 3. Sao Joao, Guayra. S.J. Pinhais (not located), I, 3. Rio de Janeiro. Angra dos Reis, IX, 6. Rio de Janeiro, X; no date,
3. Itatiaia, II, 3. Ouro Preto. Petropolis, Km 50 Estrada Contorno, 900 m, XI. Teresopolis, XII. Rio Grande do Sul.
Cochoeira (not located), X. Santa Catarina. Nova Teutonia, III, 4; no date, 11. Rio dos Autos Camayo, I. Rio dos Reis, 3.
Sfto Paulo. Alto da Serra, IX; 3. Anbembi (not located), 7. Barreiro, Serra do Bocaira, II. Bariero (not located). III.
Barueri, I; II; III; VI; XI, 4; XII, 3. Boraceia Station, Salesopolis, 850 m, I, 2; II, 2; III; IV, 4; XII, 2; no date, 14. Campos
de Jordao, 13, no date; XI. Jaba-quara, X. Consobacao (not located), II. Faz Campinas, Mogi Guacu, I, 13. Mogi das
Cruzes. Iguap, 2. Jpiranga, XI. Pampeia (not located), VIII. Parana Macaba, III. Pindamonhangaba, I, 16; III; IX; X, 9.
Porto Cabral, X. Paranapicaba. Piraciacaba, XII, 9. S. Bernando, 2. Santana, II, 6; VI; X; XI; XII, 4. San Jose dos
Campos, IX. Opasco, I. Sao Paulo, 3. Tremembe, III. Ypiranaga, XI; XII, 2. CNCI, FSCA, GMNH, IZAV, MZUSP,
UFPB, USNM.

COLOMBIA. Cundinamarca. Bogota, 6. Cajica, I. Sasaima, IV. Cesar. Valledupar, 1300 m, 2. Guajira. Sierra de
Perija, Socorpo Mission, 1400 m, IX, 5. Magdalena. Campana, 26 km S Santa Marta, 1050 m, V. El Libano, 1800 m.
Meta. E of Villavicencio, no date. 23 km NW Villavicencio, Quebrada Suumuco, 1000 m. III, 3. Narino. Mallama. Norte
de Santandar. 2 mi N Chinacota, 900 m, V. 32 km S Cucuta, Quebrada Honda, 600 m, V, 2. Putomayo. Santa Rosa, Rio
San Miguel. Quindio. No data. Valle. Km 18 Buenaventura Hwy, I. BMNH, CASC, CMNH, FMNH, FSCA, SBPC,
USNM.

COSTA RICA. Alajuela. Poasito, Volcan Poas, 1840 m, VII, 14. Cartago. Irazu, 1650 m, II; 2200 m, II; 3000 m, IX,
2. Turrialba, III, 2; VIII, 2; IX, 2. Puntarenas. Coronado, VI; VII, 4. La Palma (1500 m?), VI. Monteverde, 1400 m, V, 4;

VI, 4; VII, 2; 1500 m, II; VI, 2. 21 km NE Potrero Grande, IX, 3. 6 km N Santa Elena, 1400 m, V. San Jose. Carillo. San
Jose, 1 172 m, VIII, 6; 1200 m, I; no date, 2. 14 km N San Isidro; 1600 m, VI. CMNH, CNCI, FSLC, LACM, SBPC.

ECUADOR. Guayas. Guayaquil. Loja , 2220 m, 7. San Ramon, 27 km WSW, V, 7. Napo. Baeza, 2000 m, III, 20. 6
km N Baeza, 2000 m, II, 29. 17 km NE Baeza, 4 km SW Chaco, III, 32. 7 km S Baeza, 2000 m, II, 61. 125 km NW
Baeza, 2000 m, III, 2. 24 km NE Baeza, 1200 m, III, 4. Pastaza. 1 km E Mera, 1100 m, VII. Pichincha. 28 km NE
Alluriquin, Chiriboga Road, 1600 m, VI, 8. 3 km E Tandapi, 1300 m, VI, 25. 18 km E Tandapi, 1800 m, VI, 24. 24 km E
Tandapi, 2300 m, VI, 60. 16 km E Tandapi, 2000 m, VI, 3. Tungurahua. Banos, 1200 m, Mera Trail, IX; 1800 m, IX, 7.
39 km E Banos, I. 6 km E Rio Negro, 1400 m, VII. 8 km E Rio Negro, 1500 m, VII, 13. CASC, SBPC.

EL SALVADOR. 16 km N Metapan, Montecristo, 1760 m, V. SBPC.

GUATEMALA. Alta Verapaz. Coban, VI, 2. Patal, 5 km S Tactic, 1350 m, VIII. Chimaltenango. Yepocapa, VIII.
Peten. Pacomon (not located), VI, 2. SBPC.

MEXICO. Chiapas. 32 km N Bochil, 1700 m, VIII, 3. Cerro Tres Picos, 2000 m, 5. 8 km SW El Bosque, VI.
Montebello Lagunas, VIII. Ocozocuautla, 800 m, 2. Rosario Izapa. San Cristobal de las Casas, V, 2; VIII, 5. Santa Rosa,
VIII, 3. Distrito Federal. No data. Durango. 66 km SW Ciudad de Durango, 2300 m, VI, 15. Guerrero. 6.5 km W
Miatlan, 1450 m, IX, 3. 12 km W Maystlan, Microondas, 2150 m, IX. Hidalgo. 4.5 km N Tlanchinol, 1600 m, VIII, 2.
Jalisco. 9.5 km W Atenquique, 1700 m, IX, 2. 15 km SW Autlan, 1300 m, IX, 2. 19 km SW Cocula, 1750 m, IX, 2. Los
Volcanes, 1650 m, near El Rincon. Mexico. 5 km S Temascaltepec, 2000 m, IX, 2; 9.6 km NE, 2150 m, IX, 4. 8 km SW
Tenancingo, 2200 m, IX. Valle de Bravo, 1830 m, XI, 2. Michoacan. Morelia. Patzcuaro, IX, 2. San Jose Purua, II.
Tacambaro. Tancitaro, 1850 m, VIII, 4. Urapan, VIII. Morelos. Cuernavaca, I; VII. No data, 2. 12 km E Cuernavaca,

VII, 2. Nayarit. Tepic, VII, 3. Nuevo Leon. Allende. 29 km W Linares, Santo Rosa Canyon, 700 m, oak-thorn forest, VI,

Carrion Beetles of Latin America

265

2. Monterrey, Chipinque Mesa, 1350 m, V; VI; VII. Monte Peila, V. 21 km W Montemorelos, Cueva de Chorros de Agua,

VI, 5; Chorros de Agua, VI, 291. Rayones, III, 2. Oaxaca. 14.5 km E El Camaron 1300 m, IX. Juquila Mixes, 1450 m, VI;

VII. Oaxaca, VI; VIII. 14.5 km NE Oaxaca, 1900 m, VII, 24. 147 km N Oaxaca, V. Sierra Madre del Sur, Escondido
Road Crest, VI, 28. 24 km S Sola de Vega, 1850 m, V, 5. 13.5 km S Suchixtepec, IV. 5 km N Suchixtepec, 2900 m, VI. 12
km S Valle Nacional, 900 m, V, 18. Temascal, VI. Peubla. Cholula. 7 km SW Huauchinango, 1700 m, VII. Villa Juarez,

X. Queretaro. 29 km E Landa de Matamoros, 1600 m, VI, 2. 32 km W Xilitla, 1600 m, VI, 9. San Luis Potosi. 24 km W
El Naranjo, 1 100 m, VI, 2. Cueva de la Puente, 20 km S San Francisco, 3000 m, V. Sinaloa. 62 km NE Concordia, 1900
m, IX. El Palmito, VII, 3; 2200 m, VIII. 7 km NE La Capilla del Taxte (not located), VI, 2. Tamaulipas. 7 km W El
Encino, III, 6. Gomez Farias, Rancho del Cielo, cloud forest, 1000 m, 1150 m, VI; VII, 19. Sotano de las Salas, I.
Veracruz. Catemaco, V, 2. Chocaman, VII. Cordoba, I, 2; II, 3; III; VII, 3; VIII, 3; IX, 6; XII; no date, 3. 7 km N
Huatusco, 1300 m, VII; VIII, 3. Jalapa, V, 2; VI, 2; VII, 2; no date, 8. Orizaba, I; XI, 3; XII, 6. Perote, V, 2. Presidio, X.
Rio Metlac, near Fortin, 1000 m, VI, 2; VII, 40. Sumidero, near Fortin, 750 m, V, 6. 22.5 km S Tlapacoyan, 120 m, VII.
Cueva de Tlilapan, 5 km S Orizaba, VIII. BMNH, CASC, CNCI, FMNH, ITMM, LACM MHNM, RDCC, SBPC,
TMMC, USNM, UTDZ.

NICARAGUA. Chontales. BMNH.

PANAMA. Bocas del Toro. 20 km SE Chiriqui Grande, 900 m, VI, 7. Chiriqui. Boquete, IX, UV Light, 5. Boquete,
Alto Lindo, VII, 2; IX, UV light, 8. 5 km W Boquete, El Salto Road, 1610 m, VI, 3. 10 km W Cerro Pando, V, 39. 2 km
W Cerro Punta, 1760 m, V, 2; VI, 159. 2 km E Cerro Punta, 2200 m, VI, 37. La Fortuna Dam, 800 m, VII, 1 1; 1000 m,
VII, 4. 4 km W Hato del Volcan, 1360 m, VI, 301. Lagunas, 5 km SW Hato del Volcan, 1360 m, VI, 273. 2 km N Santa
Clara, Hartman Finca, 1200-1500 m, V, 375; VI, 41. Volcan de Chiriqui, 1200-1900 m, 4. BMNH, GMNH, SPBC,
USNM.

PARAGUAY. Alto Parana. Puente Stroessner, VIII, 8. Caaguazu. Paso Yobai (not located), X. Itapua. Trinidad, X.
Misiones. Loreto, VI, 5; X, 2. No location, 2. Paraquari. Parque Nacional Ybycui, I, 4. Parana. Iguazu, X. San Pedro.
Carumbe (not located), II. FMLC, GMNH, LPMCN, RDCC.

PERU. Cuzco. Cuzco, VIII. Marcapata, Hacienda Cadena, VIII. Huanuco. N side Cerro Carpish, nr. Chinchao, 1920
m, montane rainforest, I, 2. Junin. Perene, 600-900 m. Valle Chanchamayo, 1400 m, I; II. La Libertad. Samne, 1500 m,
VII. Pasco. Huancabamba, 3000 m, 2. Ucayali. Sinchono ( = Fundo Chinchona), 1300 m, V. La Divisorio, 1400 m, V.
AFNC, FMLC, USNM.

VENEZUELA. Aragua. Camp Rangel, VII. Choroni, V; Km 27, VI, 2; Km 25, III, 2; 9000 m, X, 10. Rancho
Grande, N of Maracay, 1100 m, I, 2; II, 78; III, 7: IV, 4; V, 9; VI, 27; VII, 5; VIII, 26; IX, 1; X, 2; XI, 2; XII, 5; 1500 m,
II, 6; 1700 m, V. Bolivar. El Dorado - Santa Elena, Km 107, 520 m, VIII, 10; Km 109, VIII, 3; Km 38, 160 m, VIII; Km
125, 1 100 m, IX, 3. Carabobo. Borburato, III, 2. Districto Federal. Caracas, Quebrada de Catuche, VI. Caracas. El Avila,
1400 m, X. El Junquito, 1900 m, VI, 7. El Limon, 1350 m, VI, 14. Macizo, Naiguanta, IX. Lara. Terepaima (not located),

XI. Merida. Merida, 5; La Pedregosa, 1800 m, IX, 4. Mucay (not located), IX. Santa Rosa, X, 2. Miranda. Fila de
Mariche (not located), 1200 m. III. San Antonio de los Altos, 1300 m, IV; V; VII; VIII, 28; IX, 31. Sucre. Carripana (not
located), VI. Tachira. San Cristobal, 1200 m, VIII, 7. 20 km NE San Cristobal, 1200 m, V, 1 1. 38 km NE San Cristobal,
2150 m, V, 12. Trujillo. Bocano, VIII, 16. Yaracuy. San Felipe. CMNH, CBMV, IZAV, MZUSP, SBPC, USNM.

UNITED STATES. Texas. Willacy County, Lyford, VI (record in Davis, 1980). This is the only known United States
record.

SUBFAMILY NICROPHORINAE
NICROPHORUS FABRICIUS 1775

These large insects are commonly called burying or sexton beetles. About 85 species are
known in the genus throughout the world. Most live in Europe and Asia. In the Old World, they
occur only as far south as Ethiopia in Africa and in southern Asia to New Guinea and the
Solomon Islands. Fifteen species live in the United States and Canada. Nine species are known
in Latin America, from Mexico southward to southern Chile and Argentina.

Key to species of Nicrophorus in Latin America

1 Posterior lobe of metepimeron (fig. 39) with many long golden hairs 2

V Posterior lobe of metepimeron with only a few dark hairs or glabrous 3

2 (1) Anterior face of procoxa with short hairs; elytral maculations large (figs. 2,

45, 48) N. marginatus Fabricius, p. 266

Quaest. Ent., 1985, 21 (3)

266

Peck and Anderson

2 ' Anterior face of procoxa with long hairs; elytral maculations large, reduced

or absent (figs. 40-44) N. guttula Motschulsky, p. 267

3 (1') Dorsal ridge of elytral epipleuron long, extending anteriorly to or past level

of apex of scutellum (fig. 46)

3' Dorsal ridge of elytral epipleuron short, not extending to level of apex of

scutellum (fig. 47)

4 (3) Elytron fasciate (fig. 54b); elytral epipleuron orange-red with pre-basal

black spot (fig. 54a) N. mexicanus Matthews, p. 267

4' Elytron and elytral epipleuron wholly black

N. nigrita Mannerheim, p. 268

5 (3') Elytral epipleuron predominantly or wholly black

5' Elytral epipleuron from about half to predominantly or wholly orange-red

6 (5) Elytral epipleuron wholly black; dorsal surface of elytra with abundant

long hairs; each elytral fascia entire, four elytral spots in total (fig. 49) . .

N. quadrimaculatus Matthews, p. 268

6 ' Elytral epipleuron black with orange-red spot at base (fig. 50a); dorsal

surface of elytra with few long hairs; each elytral fascia of most specimens
divided into two spots, thus eight elytral and two epipleural spots in total
(fig. 50b) N. didymus Brulle, p. 268

7 (50 Apical three antennomeres black; elytral epipleuron orange-red with black

spot at base (fig. 51a); elytra with fasciae confluent, dorsal surface largely

orange-red (fig. 51b) N. chilensis Philippi, p. 269

T Apical three antennomeres orange-red; elytral epipleuron wholly

orange-red; elytral pattern various

8 (70 Anterior and posterior elytral fasciae confluent laterally (fig. 52b); elytra

with dorsal surface with abundant hairs N. olidus Mathews, p. 269

8' Anterior and posterior elytral fasciae not confluent (fig. 53b); elytra with

dorsal surface with but few long hairs, most confined to lateral margins
N. scrutator Blanchard, p. 270

Nicrophorus marginatus Fabricius
Figures 2, 4, 45, 48; Map 10

Necrophorus marginatus Fabricius, 1801: 334. Type locality: “North America”. Type: location unknown, not
seen. Miller and Peck, 1979; Anderson and Peck, 1985.

Necrophorus montezumae Matthews, 1888: 92. Type locality: Mexico. Type in: BMNH, seen.

Diagnosis. — Length 15-22 mm. Pronotum markedly cordate, with narrow lateral margins
and wide basal margin (fig. 2). Anterior face of procoxa with short black hairs. Apical four
antennomeres orange-red. Metasternal pubescence dense, yellow; metepimeron with dense
yellow pubescence. Hind tibiae slightly curved. Elytron with epipleural ridge long, extending to
about level of base of scutellum (figs. 45, 46); dorsal surface lacking hairs; elytron with pattern
as in figures 2, 45, 48.

Natural history and distribution. — The species is widespread in open grassland, old field,
and shrubby habitats from southern Canada and most of the United States into northern
Mexico (Anderson and Peck, 1985). Adults have been collected throughout the summer
months. We have seen the following 17 records:

Carrion Beetles of Latin America

267

MEXICO. Coahuila. 14 km NW Saltillo, VII. Parras. Rancho Encantada, Sierra de la Encantada, VII. Distrito
Federal. Mexico City. Durango. Villa Lerdo. 32 km E El Salto, 2400 m, VI, 9. Puebla. Cholula, Esperanza. Veracruz.
Jalapa. BMNH, CNCI, SBPC.

Nicrophorus guttula Motschulsky
Figures 40-44; Map 10

Necrophorus guttula Motschulsky, 1845: 53. Type locality: Sitka, Alaska. Type: in Leningrad?, not seen. Miller and Peck,

1979; Peck and Miller, 1982; Anderson and Peck, 1985

Diagnosis. — Length 14-20 mm. Pronotum markedly cordate, with narrow lateral margins
and wide basal margin. Anterior face of procoxa with long black hairs. Apical three
antennomeres orange-red or black. Metasternal pubescence dense, yellow; metepimeron with
dense yellow pubescecne. Hind tibiae slightly curved. Elytron with epipleural ridge long,
extending almost to level of base of scutellum (figs. 40a, 44a). Elytral pattern various, as in
figures 40-44.

Taxonomic notes and distribution. — The species is widespread in dry forests and
grasslands in southwestern Canada and the western United States. The more conspicuously
maculate individuals were formerly called N. hecate Bland (Peck and Miller, 1982). Border
records indicate that the species probably occurs in northwestern Mexico. We know only of the
following record:

MEXICO. Baja California. Valle de Trinidad, Aguajito Spring, III, 3, SDMC.

Nicrophorus mexicanus Matthews
Figures 46, 54; Map 1 1

Necrophorus mexicanus Matthews, 1888: 91. Type locality: Mexico. Type in: BMNH, seen.

Diagnosis. — Length 14-18 mm. Pronotum quadrate, with wide lateral and basal margins.
Apical three antennomeres orange-red. Metasternal pubescense dense, dark brown;
metepimeron with small tuft of dark brown hairs. Hind tibiae straight. Elytron with epipleural
ridge long, extending almost to level of base of scutellum (fig. 46); dorsal surface lacking hairs.
Elytron with pattern as in figure 54.

Natural history and distribution. — The species occurs in habitats ranging from semi-arid
and open thorn scrub to moist closed-canopy cloud forests in the southwestern United States,
through Mexico, to Guatemala and El Salvador. Over its southern range, adults have been
collected in all months of the year. Zaragoza and Perez (1975) give a morphometric and
seasonal analysis based on 436 specimens collected in black light traps over three years near
Mexico City. They report N. mexicanus to be most abundant in October. Reproductive
behavior has been studied by Halffter et al.( 1982). We have seen 127 specimens representing
the following records:

EL SALVADOR. Montecristo, 23 km N Metapan, 2300 m, cloud forest, V, 8. SBPC.

GUATEMALA. Zacapa. Jabah, S slope Sierra Minas, VII. FMNH.

MEXICO. Chiapas. 5 km W San Cristobal de las Casas, 2440 m, IX, 2. Chihuahua. Mesa del Huracan, 2557 m, VII,
4. Nuevo Casas Grandes, 20 km SE, Hwy 10, 1700 m, VIII. Sierra de la Catarina, 30 km SW Buenaventura, 2600 m,
VIII, 9. Sierra de Choreachic, Microwave Sta. Hwy 16, 30 km W Cuauhtemoc, 2500 m, VIII. Sierra Huachinera, 30 km
SW Colonia Juarez, 2200 m, VIII, 3. Distrito Federal. Lomas, V. Mexico City, V. No locality, III. Durango. 5 km W El
Salto, 2745 m, VI, 15; VII, 8. 16 km W El Salto, 2745 m, VI; VII, 16; VIII. 66 km SW Durango. 2250 m, VI, 2. Ciudad
de Durango, 1800 m, IV. 38 km W La Ciudad, VII, 20. 32 km E El Salto, 2400 m, VI, 5. 54 km E El Salto, 2100 m, IX, 5.
Guerrero. 12 km W Mazatlan, 2130 m, IX, 3. Hidalgo. Guerrero Mills. 10 km S Tenango de Doria, 3000 m, VII, 5.
Mexico. Ayolta. Nuevo Leon. Galeana, 2217 m. Iturbide, 1800 m, VII, 3. Oaxaca. La Parada. Sierra Madre del Sur,
Escondida Road Crest, VI. 5 km N Suchixtepec, 2850 m, VI. Yolotepec. Road to Yuvila, 2430 m, VIII, oak-pine forest, 2.
Tamaulipas. Gomez Farias, Rancho del Cielo, 2000 m, XII. Tlaxcala. 3 km S Apizaco, pine forest, VI, 3. BMNH, CNCI,

Quaest. Ent., 1985,21 (3)

268

Peck and Anderson

FMNH, LACM, MCZC, MHNM, OSUC, SBPC.

Nicrophorus nigrita Mannerheim
Map 10

Necrophorus nigrita Mannerheim, 1843: 251. Type locality: California. Type: in Helsinki?, not seen. Miller and Peck,

1979; Anderson and Peck, 1985.

Diagnosis. — Length 13-18 mm. Pronotum quadrate, with wide lateral and basal margins.
Apical three antennomeres orange-red. Metasternal pubescense dense, dark brown;
metepimeron glabrous. Hind tibiae straight. Elytron with epipleural ridge long, extending
almost to level of base of scutellum; dorsal surface lacking hairs. Elytron wholly black.

Natural history and distribution. — The species occurs in drier forests on the Pacific coast
from British Columbia to southern California, including the Channel Islands (Miller and Peck,
1979). Horn (1876, 1880) lists the species from Guadalupe Island, Baja California. We know
of only the following Mexican records:

MEXICO. Baja California. Guadalupe Island, 5 (not shown on map). 10 km E El Rosario, uv light. III. MCZC,
UICM.

Nicrophorus quadrimaculatus Matthews
Figures 49, 55; Map 12

Necrophorus quadrimaculatus Matthews, 1888: 93. Type locality: Guatemala. Type in: BMNH, seen.

Diagnosis. — Length 9-16 mm. Pronotum orbicular with wide lateral and basal margins
(fig. 55). Apical three antennomeres orange-red. Metasternal pubescense moderately dense,
dark brown; metepimeron with long dark brown hairs. Hind tibiae very slightly curved. Elytral
epipleural ridge short (as in fig. 47); dorsal surface of elytron with long, dense hairs. Elytron
with pattern as in figure 49. Metatrochanter with sharp spine.

Natural history and distribution. — The species ranges from southern Mexico to western
Panama. Adults have been collected only from June to September, and only in montane pine or

cloud forests. We have seen 52 specimens representing the following records:

COSTA RICA. Puntarenas. Monteverde, 1400 m, VI; VII, 2; IX; 1520 m, VII; 1700 m, V. No data, 2. BMNH,
CMNH, LACM, SBPC.

EL SALVADOR. 16 m N Metapan, Montecristo, 1760 m, mixed pine forest, V, 2. SBPC.

GUATEMALA. Alta Verapaz. 6 km S Coban, 1373 m, VIII, 5. Patal, 5 km S Tactic, 1373 m, VIII. Senahu, 1098 m,
VIII. Baja Verapaz. San Jeronimo. Quezaltenango. Volcan Zunil. BMNH, SBPC.

MEXICO. Chiapas. Lagunas de Montebello, 1373 m, VIII. 1 km SW Rizo de Oro, 834 m, VIII. SBPC.

PANAMA. Chiriqui. 4 km N Santa Clara, Cerro Pelota, Hartman Finca, 1200 m, V; 1500 m, V, 20; VII, 6; VIII, 5.
SBPC.

Nicrophorus didymus Brulle
Figure 50; Map 13

Nicrophorus didymus Brulle, 1840: 73. in Brulle and Blanchard, 1840. Type locality: Altamachi River, eastern mountain
slopes, Cochabamba, Bolivia. Type in: MNHN?, not seen. Berg, 1901: 326; Portevin, 1903: 331.

Nicrophorus didymus var. peruvianus Pic, 1917: 2. Type locality: Peru. Type in: MNHN?, not seen.

Nicrophorus flexuosus Portevin, 1924: 191. Type locality: not given. Type in: MNHN?, not seen. Hatch, 1928: 128.
Nicrophorus flexuosus var. portevini Pic, 1933: 6. Type locality: Merida, Venzuela. Type in: MNHN?, not seen.

Diagnosis. — Length 13-16 mm. Pronotum orbicular with wide lateral and basal margins.
Apical three antennomeres orange-red. Metasternal pubescense moderately dense, dark brown;
metepimeron with few short brown hairs. Hind tibia very slightly curved. Elytron with
epipleural ridge short, not extending to base of scutellum; dorsal surface with long and

Carrion Beetles of Latin America

269

moderately dense hairs. Metatrochanter with spine reduced or absent. Elytra with pattern as in
figures 50a, 50b.

Variation. — Many elytral patterns have received varietal and aberrational names. The
elytral maculations vary from two distinct bars to four distinct spots (a pair of spots forming
from a color bar) on each elytron. We have not seen a pattern of geographic variation in such
coloration.

Natural history and distribution. — The species occurs in middle to upper elevation Andean
forests from Venezuela through Colombia, Ecuador, and Peru to Bolivia. We believe literature
records for the species from Mexico, Central America, and Argentina pertain to other species.
Adults have been collected during eight months of the year. We have seen 76 specimens

representing the following records:

BOLIVIA. Cochabamba. Incachaca, 2300 m. MCNH.

COLOMBIA. Antioquia. Medellin. Magdalena. Rio Don Amo, 600 m, VII. Rio Don Diego, 36 m, VII. No other data.
BMNH, CMNH.

ECUADOR. Napo. 7 km S Baeza, 2000 m, II, 2. El Chaco, 2000 m, II. Province Unknown. Mangosia River (not
located), 650 m. BMNH, SBPC.

PERU. Huanuco. N side Cerro Carpish, near Chinchao, cloud forest, 2300 m, I, 24; 2400 m, I. Pasco. Oxapampa,
1800 m, I. Pozuzo (10°4'S 75°32'W). AFNC, FMLC, SBPC, USNM.

VENEZUELA. Aragua. Cerro Choroni, 1600 m, II. Rancho Grande, N of Maracay, 1500 m, II, 8; V; VIII, 2; XII.
Maracay to Choroni, 1000 m, XII; 1300 m, XII. Districto Federal. Caracas, no data. Caracas, Rio Caurimare, 1000 m, V,
7. El Junquito, VI; X. El Limon, 1350 m, VI, 5. Lara. Cabudare, Terepaima Creek, 1200 m, I. Tachira. San Cristobal,
1200 m, VIII, 17. Trujillo. Bocono, VIII. Zulia. Sierra de Peria, Kunana, 1 100 m, XII. BMNH, CBMV, IZAV, SBPC.

Nicrophorus chilensis Philippi
Figure 51; Map 14

Necrophorus chilensis Philippi, 1871: 293. Type locality: Santa Cruz, Curico, Chile. Type in: MNSC, holotype no. 171,

not seen.

Diagnosis. — Length 13-16 mm. Pronotum subquadrate with wide lateral and basal
margins. Apical three antennomeres black. Metasternal pubescence dense, dark brown;
metepimeron glabrous. Hind tibiae straight. Elytron with epipleural ridge short, not extending
to level of apex of scutellum; dorsal surface lacking hairs. Metatrochanter with sharp spine.
Elytron with fasciae confluent and large, pattern as in fig. 51.

Variation. — The anterior and posterior elytral maculations of some specimens are joined
and may be so large that the black area is reduced to only the extreme anterior and posterior
sutural margins of the elytra.

Natural history and distribution. — The species apparently occurs in open and semi-arid
areas of central Chile and adjacent Argentina (Pena, 1981), as well as in Nothofagus and
Arucaria forests of Chile. Adults are seemingly active from November to March. We have seen

only 19 specimens representing the following records:

ARGENTINA. Neuquen. Lago Tramen, 1000 m. III; XI; XII. San Martin de los Andes, XII. Tucuman. Tucuman
(questionable location). Locations Unknown: “Patagonia”, no data, 4. “Pampas”, no data. “Salinas Chicas”, no data (in
Berg, 1901). LPMCN.

CHILE. Curico. Cordillera de Teno. Malleco. 6.5 km E Malalcahuello, 1080 m, Nothofagus forest, XII. 14 km E
Malalcahuello, 1570 m, Nothofagus and Aurucaria forest, XII. Laguna Jesus-Maria and Pino Hachado (records of Pena,
1981). Maule , No data, 3. Valdivia. No data. No locality, 2. AFNC, BMNH, MNSC, SBPC.

Nicrophorus olidus Matthews
Figure 52; Map 15

Necrophorus olidus Matthews, 1888:92. Type locality: Mexico. Type in: BMNH, seen.

Quaest. Ent., 1985, 21 (3)

270

Peck and Anderson

Diagnosis. — Length 10-14 mm. Pronotum orbicular with wide lateral and basal margins.
Apical three antennomeres orange-red. Metasternal pubescense moderately dense, dark brown;
metepimeron with a few short brown hairs. Hind tibia very slightly curved. Elytron with
epipleural ridge short, not extending to apex of scutellum; dorsal surface with long dense hairs.
Metatrochanter with sharp spine. Elytra with pattern as in figure 52.

Natural history and distribution. — The species is known only from Mexico north of the
Isthmus of Tehuantepec. Adults have been collected from May to November. It occupies open
forests, cloud forests, and rainforests. We do not accept Portevin’s (1926) statement that the
species occurs in Central America and Colombia. We have seen 267 specimens representing the
following records:

MEXICO. Durango. 66 km SW La Ciudad de Durango, 2250 m, VI, 6. Revolcaderos, VII, 3 (location unknown).
Guerrero. Omilteme. Xucamanatlan. Hidalgo. 10 km S Tenango de Doria, 3000 m, VII, 2. Jalisco. Ajijic, 1567 m, V, 3;
XI, 2. 9.5 km W Atenquique, 1677 m, XI, 12. 13 km W Atenquique, 1799 m, IX. 15 km SW Autlan, 1312 m, IX, 30. 19
km SW Cocula, IX, 6. El Rincon, Los Volcanes, 1647 m. Cd. Guzman (L. de Zapotlan). Mexico. 5 km NE
Temascaltepec, 1922 m, IX, 5. 9.6 km. NE Temascaltepec, 2135 m, IX, 2. Tenancingo, 2165 m, IX. Morelia. 12 km E
Cuernavaca, VII. Nuevo Leon. Iturbide, 1800 m, VII, 2. 26 km W Linares, 671 m, V. Oaxaca. 1 1 km E Hautla. Juquila
Mixes, XI, 4. 14.5 km NE Oaxaca, 1891 m, VIII, 7. 84 km S Oaxaca, V, 2. Sierra Madre del Sur, Escondido Road Crest,
VI, 14. 24 km from Sola de Vega, 1830 m, V. Suchixtepec. 21 km S Valle Nacional, 1098 m, VII, 16; VIII, 8. 24 km S
Valle Nacional, 1220 m, V. Puebla. 7 km SW Huachinango, 1700 m, VII, 2. Nuevo Necaxa, VII, 3. Queretaro. 29 km E
Landa de Matamoros, 1617 m, VI, 3. 32 km W Xilitla, VI. San Luis Potosi. 17.5 km W El Naranjo, 960 m, VI, 25. 20 km
W Xilitla, 1600 m, cloud forest, VI-VII, 3. 22.5 km W Xilitla, 1312 m, VI, 3. Tamaulipas. 10 km W El Encino, 2000 m,
cloud forest, VII. Gomez Farias, 300 m, tropical deciduous forest, VI-VIII, 34; Rancho del Cielo, cloud forest, 1000 m,
VI- VIII, 25; 1129 m, VII, 27. Veracruz. Cordoba. Tuxtla. 1.9 km S Huatusco, 1344 m, VIII, 2. 7 km N Huatusco, 1281
m, VIII, 3. 8 km W San Andres Tuxtla, VII.

Nicrophorus scrutator Blanchard
Figure 53; Map 14

Necrophorus scrutator Blanchard, 1840: 74, in Brulle and Blanchard, 1840. Type locality: Bolivia. Type in: MNMN?, not
seen.

Diagnosis. — Length 17-22 mm. Pronotum orbicular, with wide lateral and basal margins.
Apical three antennomeres orange-red. Metasternal pubescence dense, dark brown;
metepimeron with a few short brown hairs. Hind tibia very slightly curved. Elytron with
epipleural ridge short, not extending to level of scutellar apex; dorsal surface with only a few
long hairs, majority confined to lateral margins. Metatrochanter with spine reduced, broadly
rounded. Elytron with pattern as in figure 53.

Variation. — A single specimen from Machu Pichu, Peru (in MZUSP) has an epipleuron
with slightly more than the anterior half orange-red and the two fasciae on each elytron are
confluent along the epipleural margin. This specimen also has typical brownish metasternal
pubsecence and the last three antennal segments are orange-red.

Natural history and distribution. — The species occurs in Peru, Bolivia and northwestern
Argentina, seemingly in both open semi-arid and moist forested habitats. It is active from

October to April. We have seen 40 specimens representing the following records:

ARGENTINA. Catamarca. Andalgala, 4. Cuesta Mina Capillas (not located), 3200 m, II. Las Estancias (not
located), I. San Angelo (not located), II. Jujuy. Jujuy, II. Volcan. Misiones. No data, questionable record. Salta. Anta,
XII. 20 km N La Caldera, El Ucumar, 780 m, I; II. Cerillos, 1200 m, X. Tucuman. Ciudad Universitaria, San Javier,
Tucuman, II. Horco Molle, 12 km W Tucuman, 700 m, I. Infiernillo, III. Mala-Mala, 2000 m, IV. Parque Aconquija, IV.
Quebrada de Lules, III, 3; XII. Rio Pueblo Viejo, 1000 m, humid forest. San Pablo, 1200 m, 2. Siambon, II; VII. Tafi
Viejo. Tucuman, VI; no date, 2. Villa Nouques, V; XII. Villa P. Mont, Burrayacu. BMNH, FMLC, LPMCN, SBPC.

BOLIVIA. Chuquisaca. La Laguna, Neubo Mundo Mountains, XII (Brulle and Blanchard, 1840). Tiguipa, IV.
Pando. Rio Negro, II, doubtful record. Santa Cruz. Valle Grande (between Chilan and Tasajos, X (Brulle and Blanchard,
1840: 74). No data, 2. BMNH, MNHN.

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PERU. Cuzco. Machu Pichu, 2600-2800 m, VII. MZUSP.

PHYLOGENY AND ZOOGEOGRAPHY

In this section we discuss the phylogenetic and geographic relationships of the Latin
American silphid fauna and propose hypotheses about its origin.

RECONSTRUCTED PHYLOGENY OF OXELYTRUM

Ranges of all silphines of Latin America (listed in table 1), except Oxelytrum , extend into
Mexico from the north and terminate at or before the southern edge of the Mexican
Neo-Volcanic Plateau. Of these genera, only Heterosilpha is endemic to North America. All
other genera have the majority of their species and ranges in Eurasia. A phylogenetic analysis
of these genera can best be accomplished by including the Palearactic and Oriental species and
will not be attempted here.

We do, however, present a reconstructed phylogeny for all members of the wholly New
World genus Oxelytrum (fig. 56). Oxelytrum is identified as a monophyletic group on the basis
of possession of the derived character state of presence of coxal spines or tubercles. Two
lineages, each including four species, are recognized within Oxelytrum. One lineage, the
lineatocolle group, is characterized by the derived character state of a black pronotum. It is
associated with western coastal lowland and Andean montane habitats. Members of the other
lineage, the emarginatum group, share the derived character state of a pronotum with reflexed
margins, and a generally very similar overall habitus. It is associated with eastern and northern
montane and lowland habitats.

The character analysis uses only adult characters (table 3) and is based on out-group
comparison with the silphine genus Ptomaphila of Australia and New Guinea. These two
genera are considered to compose a monophyletic group based on their shared possession of the
derived character states of long hairs on the underside of the elytra near the apical callus, and a
pronotum with elevated costae. These character states appear in no other Silphinae. We
interpret these two genera as otherwise comparatively primitive, of great antiquity and derived
from a common Gondwanaland ancestor.

Oxelytrum characters. — Characters used are those of taxonomic value. Whether all such
characters are valuable as indicators of phyogenetic relationships is questionable. Characters
have not been objectively weighted. As virtually all characters have unknown biological
significance, it is not known how prone they are selective pressures promoting homoplasy.
Nevertheless, gross inferences have been made concerning the degree of homoplasy expected in
each character (Table 3).

Character 1, pronotum-dorsal surface. Two character states have been identified. Because
elevated costae are not known in other Silphinae aside from Oxelytrum and Ptomaphila this
state is interpreted as apotypic and of high weight.

Character 2, elytra-undersurface. Two character states have been identified. Because long
hairs in the region of the apical callus are unknown in other Silphinae this state is interpreted as
apotypic and of high weight.

Character 3, elytra-apex. Two character states have been identified. An internal flange is
well-developed in members of Ptomaphila and lacking in Oxelytrum species. No other
Silphinae possess such a feature, considered here as apotypic, although development of similar
structures do occur in other Coleoptera probably as means of locking the elytra together at the

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apex.

Character 4, head-frons. Three character states have been identified although there is
variation in the degree to which the frons is swollen. In most other Silphinae, the frons is
uniformly swollen while in Oxelytrum species a “v-shaped” swollen area (chevron) is present.
Ptomaphila species lack any degree of swelling on the frons. Because the frons is swollen in
members of Nicrophorinae, lack of or reduction in swelling is interpreted as apotypic. Tendency
towards reduction in swelling could also be interpreted as further evidence of sister-group
relationship between Oxelytrum and Ptomaphila.

Character 5, elytra-dorsal surface. Two character states have been identified although
within species of Oxelytrum and other Silphinae, there is variation in the degree of elevation of
costae. Presence of tubercles arranged linearly in place of costae (latter are only vaguely
defined) is interpreted as apotypic. Linearly placed tubercles are lacking in all other Silphinae
although tubercles interspersed between costae are known in species of Thanatophilus.

Character 6, elytral epipleuron-width. Two character states have been identified. Because a
narrow epipleuron is found in most Silphidae this state is considered plesiotypic.

Character 7, coxae-ornamentation. Two character states have been identified: lack of any
ornamentation and, presence of ornamentation as tubercles or spines. Because spines and
tubercles are not known in other Silphinae, presence of these structures is considered apotypic.

Character 8, pronotum-color. Four character states have been identified. Because members
of Ptomaphilia possess a pronotum with the margins orange-red and disc black, presence of this
state in Oxelytrum species is considered plesiotypic. An entirely black pronotum is interpreted
as apotypic. Presence of a black pronotum with the posterolateral corners orange-red is
interpreted as autapotypic and derived from an entirely black pronotum. A fourth character
state is represented by a reduction in the size of the central black spot on the disc. Pronota with
colored margins are known in species of other silphine genera but it is not known if they
represent symplesiomorphy or secondary apotypic developments. Although the sole basis for
recognizing monophyly of the lineatocolle group, it should be emphasized that this is a
character likely prone to convergence and should thus be considered accordingly.

Character 9, eyes-size. Two states have been identified. Because large eyes are found in
species of Ptomaphila , this state is considered plesiotypic. Eye size is undoubtedly correlated
with diel activity patterns. Nocturnal species have large eyes; diurnal species small eyes.
Accordingly this character is extremely prone to homoplasy and should be weighted
accordingly.

Character 10, pronotum-posterior angles. Two states have been identified. Because obtuse
angles are known in species of Ptomaphila and most other Silphinae, this state is considered
plesiotypic.

Character 11, female genitalia-stylus. Two states have been identified. Because aberrant
scoop-like styli are unknown in any other Silphinae, they are considered apotypic. Whether the
modification of the styli represents a change in oviposition habits is not known.

Character 12, coxae-ornamentation. Two states have been identified. Spines and tubercles
on the coxae are not known in other Silphinae. Presence of spines is considered apotypic
because they represent a likely progression from an ancestor which possessed tubercles, the
plesiotypic state. Consideration of spines as apotypic is also compatible with distribution of
states of character 1 1 and with overall similarity of members of the group being defined.

Character 13, pronotum-margins. Three states have been identified. Because flat or
deflexed pronotal margins are known in other Silphinae, including Ptomaphila , reflexed

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273

margins are considered apotypic. States of this character represent apparently simple changes
and are probably prone to homoplasy. Distribution of states of this character is the sole basis
for recognizing the monophyly of the emarginatum group, and the latter should be considered
accordingly.

Character 14, abdominal segments of male-lateral margins. Two states have been identified
although there is variation in species of Oxelytrum in the prominence of the lateral projections.
Because lateral projections are unknown in other Silphinae, their occurrence in species of
Oxelytrum is considered apotypic. The reduced prominence of the projections in O. discicolle
are interpreted as secondarily reduced and autapotypic.

Character 15, elytra-humeri. Two states have been identified. Toothed humeri are known in
some species of Oiceoptoma. However, they are not known in Ptomaphila or other Silphinae
and thus likely represent independent developments. Presence of a tooth on the humerus in
species of Oxelytrum is therefore considered apotypic but possibly homoplasous.

Character 16, pronotum-color. Four states have been identified. Two of these have been
considered previously. A pronotum with the central black spot markedly reduced in size is
considered apotypic, derived from a similarly colored pronotum with the black spot larger in
size.

RECONSTRUCTED PHYLOGENY OF NICROPHORUS

We present here a phylogenetic analysis of the relationships of all species of Nicrophorus in
the New World (listed in table 2), based upon both adult and, where available, larval
characters (tables 4-7 and figures 57-60). Larval characters and interpretations of their
polarity are from Anderson (1982a). The adult characters, habitats, and distributions of the
Nearctic species are from Anderson and Peck (1985), and are interpreted for the first time
here. Polarization of all larval and adult character states in Nicrophorus are based on
out-group comparison with the Asian genus Ptomascopus, the only other genus in the
subfamily Nicrophorinae. We also make predictions about phylogenetic affinities of some
Nicrophorus species which are testable by the discovery and description of their larvae.

Since only New World species of Nicrophorus are considered, the cladograms may require
subsequent modification when Old World species are included. This will be especially so if the
New World component of a particular species group is found not to be monophyletic. Hatch
(1927) gives a start at an evolutionary analysis, but his assignment of Nicrophorus species into
groups was based on shared ancestral characters (symplesiomorphies) and on characters which
we believe are subject to convergence. We agree with his placement of some species, but dispute
others. Since we have not carefully studied many Palearctic species, we refrain from including
any of these in our delimited species groups although we think that at least some Palearctic
species are easily placed in our groups. We do not attempt to demonstrate relationships between
species groups. This can only be reliably accomplished following examination of all
Nicrophorus species.

The orbicollis group

This species group is characterized by the uniquely derived adult character states of a short
elytral epipleural ridge and by most members having prominent hairs on the dorsal surface of
the elytra. We place six New World species in this group and propose phylogenetic
relationships as in table 4 and figure 57. Larvae of all Latin American species are undescribed.

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orbicollis group characters. — Character 1, elytral epipleuron-length. Two character states
have been identified. Because a long elytral epipleuron occurs in Ptomascopus species and
Silphinae, this state is considered plesiotypic. A short epipleuron is also known in some
southeast Asian species of Nicrophorus, however the relationships of these species to members
of the orbicollis group have yet to be assessed.

Character 2, pronotum-shape. Four character states have been identified in Nicrophorus
species. For Nicrophorus , a cordate pronotum is considered plesiotypic because this is the state
which occurs in species of Ptomascopus. Among subquadrate, quadrate and orbicular pronta,
we hypothesize the subquadrate condition as plesiotypic, directly derived from a cordate
pronotum. We believe that quadrate and orbicular pronota each represent unique apomorphic
states derived directly from the subquadrate state. Alternative interpretations are perhaps
equally likely concerning the position in the transformation series of the orbicular pronotum.
Alternatively, this state could be directly derived from the primitive cordate state. We find the
former alternative to be most compatible with distribution of states of other characters.

Characters 3,5,6 elytron-vestiture of dorsal surface. Five character states concerning the
nature ofthe dorsal vestiture have been identified in Nicrophorus species. A dorsal surface with
very few or no hairs is considered plesiotypic because this state also occurs in species of
Ptomascopus. We hypothesize increasing density of the hairs as increasing degrees of apotypy.
Hairy elytra are also known in some southeast Asian species of Nicrophorus. However, the
relationships of these species to members of the orbicollis group have yet to be assessed.

Character 4, metatrochanter-ornamentation. Two states have been identified in
Nicrophorus species. Because a metatrochanter with a sharp, well-developed spine is known in
other Nicrophorus and Ptomascopus a reduced, blunt spine is considered apotypic.

Character 7, elytral epipleuron-vestiture. Three character states have been identified in
Nicrophorus species. Because an epipleuron with a few short hairs is known in Ptomascopus
species, this state is considered plesiotypic. Occurrence of a densely hairy epipleuron is
probably correlated with a densely hairy elytral dorsal surface, and is considered apotypic.

New World members of this group are associated with forested or open habitats in both
North and Latin America. This species group appears to have its center of diversity in Latin
America, with only one New World species being distributed north of Mexico. All species of
Nicrophorus known from southern Central and South America belong to this group. This may
indicate that the group is endemic to the New World and that the species evolved from an early
lineage within Nicrophorus. This latter suggestion is supported by the plesiomorphic condition
of larval character states of TV. orbicollis (Anderson, 1982a).

According to Portevin (1920a, 1926), the derived character states of short epipleural ridges
and hairy elytra are also known to occur (at least in part) in TV. distinctus Grouvelle
(Sulawesi( = Celebes) Islands), P. heurni Portevin (New Guinea), and TV. podagricus Portevin
(Borneo and Sulawesi) and also in TV. kieticus Mroczkowski (1959) from the Solomon Islands.
The relationship of these southern Indo-Malayan species to our orbicollis species group may be
of importance for subsequent biogeographic interpretations to be discussed later.

The defodiens group

This species group can be defined by the uniquely derived larval character states of narrowly
separated labial palpi with the basal segment ventrally unsclerotized.

defodiens group characters. — Character 1, larval labial palpi-relative position of bases.
Two states have been identified. Because labial palpi with widely separated bases are known in

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Ptomascopus morio , this state is considered plesiotypic.

Character 2, larval labial palpi-sclerotization of basal segment. Two states have been
identified. Because labial palpi with a sclerotized ventral surface of the basal segment are
known in Ptomascopus morio, this state is considered plesiotypic.

Character 3, pronotum-shape. See discussion of Character 2 in the orbicollis group.

Character 4, adult antennomeres 9,10-setosity. Two character states have been identified.
Because white setae on the ventral surfaces of antennomeres 9 and 10 are not known in
Ptomascopus species or other Nicrophorus species, this state is considered apotypic. Dense
setae, arranged in a “figure eight” pattern are autapotypic in N. vespilloides.

We place three New World species in this group and their relationships are indicated in
table 5 and figure 58. New World members are associated with northern Nearctic forested and
swampy habitats. No species in the group is known or suspected to occur in Latin America.

The investigator group

This species group can presently best be defined by the uniquely derived character state of a
prepupal overwintering stage. In N. mexicanus and N. nigrita there are no known
overwintering stages but the species are most active in fall, winter and spring seasons
suggesting descent from an ancestor with a prepupal overwintering stage. Both N. nigrita and
N. investigator share the derived larval character of a sclerotized ventral apex of abdominal
segment 10.

investigator group characters. — Character 1, overwintering stage. Two stages have been
identified based on studies of populations at northerly latitudes. We hypothesize overwintering
as an adult as plesiotypic and expect that it occurs in Ptomascopus. Southerly species that are
fall-through-spring active are considered derived from a northern ancestor which had a
prepupal overwintering stage.

Character 2, adult metasternum-vestiture. Two states have been identified. Because a
uniformly pubescent metasternum is known in Ptomascopus species and all other Nicrophorus
species, this state is considered plesiotypic.

Character 3, larval abdominal segment 10-ventral apex. Two states have been identified.
Because an unsclerotized apex is known in Ptomascopus species and all other Nicrophorus
species, this state is considered plesiotypic.

Character 4, pronotum-shape. See discussion of Character 2 in the orbicollis group.

Character 5, adult metasternum-color of vestiture. Two states have been identified. Because
yellow pubescence is found in primitive Nicrophorus species, this state is deemed plesiotypic
within the investigator group. Yellow pubescence may be apotypic for the genus Nicrophorus
because Ptomascopus species possess brown pubescence. Distribution of states of this character
in all Nicrophorus species suggests a high degree of homoplasy and cautions against its
overemphasis. Distributional data on N. mexicanus and N. nigrita, and uniformity of habitus,
also support a sister-species relationship between these two species.

We place five New World species in this group and their relationships are indicated in table
6 and figure 59. Larvae of N. mexicanus are undescribed but are expected to possess the
derived character states in table 6 based upon our interpretation of its phylogenetic position
with respect to other members of this group. New World members of the group are associated
with open, sparsely forested, and densely forested habitats throughout North America. Within
this group only N. mexicanus and N. nigrita have ranges extending into Mexico. N. nigrita
occurs not only on the mainland of Baja California, but has dispersed 250 km to Guadelupe

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Island. It has also crossed smaller water gaps to the California Channel Islands. N. mexicanus
extends throughout the Mexican Plateau and into Guatemala. It and N. marginatus are the
only species of Nicrophorus with an extensive distribution in both the United States and Latin
America.

The marginatus group

This species group is defined by the uniquely derived adult character state of dense yellow
hairs on the posterior lobe of the metepimeron and the derived larval character state of an
unsclerotized base of the venter of abdominal segment 10.

marginatus group characters. — Character 1, larval abdominal segment 10-ventral base.
Two character states have been identified. Because a sclerotized base occurs in Ptomascopus
species, this state is considered plesiotypic.

Character 2, adult metepisternum, posterior lobe-pubescence. Three states have been
identified in Nicrophorus species. Because a metepimeral lobe with a few sparse hairs is known
in Ptomascopus species, this state is considered plesiotypic.

Character 3, larval abdominal segment 9-sternite. Two states have been identified. Because
a sternite with the outer angles acute is known in Ptomascopus morio , this state is considered
plesiotypic.

We place three New World species in this group and their relationships are shown in table 7
and figure 60. A fourth, N. carolinus , is tentatively placed within this group although adults
lack the above derived characters and larvae are undescribed. However, we suspect that N.
carolinus is phylogentically close to the marginatus group based on its overall habitus and
retention of some ancestral character states, shared with members of the marginatus group.
We predict that larvae, when described, will support these suspicions (fig. 60). New World
members of the group (excluding N. carolinus) are primarily associated with open habitats
throughout western North America. Within the group, only N. marginatus has a distribution
which extends into the arid regions of the northern half of the Mexican Plateau.

Incertae Sedis

At present we are unable to assign the North American N. americanus and N. pustulatus to
definable New World species groups. This is partly due to the fact that larvae are undescribed
for both species, and that adults retain primitive states for all characters used to define the
above groups. Our inability to assign these species to groups may also be because they have no
other relatives in the New World, as has already been suggested for N. americanus by
Anderson (1982c).

ZOOGEOGRAPHY AND SPECIES ORIGINS

Silphinae other than Oxelytrum. — Genera and their number of included species in North
America north of Mexico are as follows: Necrodes (1), Thanatophilus (5), Aclypea (2),
Oiceoptoma (3), Heterosilpha (2), and Necrophila (1). Of these, only Heterosilpha is endemic.
We assume all but Heterosilpha to have originated in the Palearctic region because this is
where their highest species diversity is, and where several additional related genera occur. We
assume that members of these genera independently invaded North America at least five
different times in the Tertiary, probably across the Bering Land Bridge, but alternatively
across North Atlantic land bridges, before the opening of this ocean in the early Tertiary

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(Matthews, 1979). There is no evidence that any of these silphine genera occurred in Latin
America any farther south than the edge of the Mexican Plateau. In these genera, the sole
endemic Latin American species, T. graniger , is readily interpreted as a southern isolate of T.
lapponicus or its ancestor, which reached the uplands of the Mexican Plateau and the
Transverse (or Neo) Volcanic Sierra in a cooler glacial climate, and became isolated there in a
warmer interglacial.

Latin American silphid fossils are not known. Hatch (1927) reviewed the known Mesozoic
and Tertiary fossils known to that time and attributed to Silphidae. They tell little about the
origin of extant North American or Latin American genera. This is also true of the recent
review of Russian Mesozoic beetles (Arnoldi et al., 1977).

Oxelytrum. — The sister group relationship of a Latin American genus ( Oxelytrum ) and an
Australian genus ( Ptomaphila ) is a pattern known in many insect and other groups (reviewed
in Keast, 1973). This pattern can be best interpreted as resulting from separation of an
ancestral distribution on at least part of the southern Mesozoic supercontinent of
Gondwanaland following its breakup during the Cretaceous. Temperate lands remained in
proximity between Australasia and South America into the Eocene (50 million years BP), some
40 million years after their separation from Africa, and 30 million years after the separation of
New Zealand (Raven and Axelrod, 1975). No other closely related silphine genera occur on the
other present or formerly southern main land masses of New Zealand, Africa, Madagascar, or
India. We suggest that Oxelytrum diversified and speciated after the separation and isolation
of South America from other southern land masses.

We suggest initial divergence of this Oxelytrum stock into two lineages; the lineatocolle
group in more western coastal lowland habitats, and, the emarginatum group in more northern
and eastern lowland habitats.

The lineatocolle group probably had an ancestral species possessing many character states
similar to those of O. lineatocolle and may have originally occupied temperate habitats such as
Nothofagus forests along the western coast of South America. We hypothesize that the first
phase of the Andean orogeny during the late Cretaceous may have provided the earliest set of
barriers allowing for the divergence of this lineage into two descendant forms. One of these is
currently represented by O. lineatocolle in the south-central Chilean coastal lowlands and
Andean slopes. The other, perhaps a more inland and higher elevation form, representing the
ancestor of the remaining three species in this group, underwent subsequent divergence into
(1), a more southerly cold-temperate, but lower-elevation montane form, and (2), a more
northerly cold-adapted high-elevation montane form. This perhaps occurred during the second
phase of Andean orogenic activity and formation of high elevation grassland habitats. The first
is presently represented by O. biguttatum in extreme southern Chile and Argentina. The second
represents the ancestor of O. apicale and O. anticola which probably inhabited the high
elevation grasslands and steppes of Argentina, Bolivia, Ecuador and Peru. A possible early
Pliocene or Pleistocene isolation of northern and southern forms, perhaps as a result of glacial
events (Noonan, 1981), is indicatedby the descendant species, the more northerly O. anticola
and the more southerly O. apicale , allopatrically distributed in these high Andean open
habitats.

In the emarginatum group, the ancestral species probably possessed many character states
similar to those of O. emarginatum and may have occupied the lowland forests of northern,
central and eastern South America. We hypothesize isolation of a more upland form in the
southern Brazilian Highlands, presently represented by O. emarginatum , and a widespread

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northern lowland form representing the ancestor of the remaining three species in this group.
This latter form diverged into southerly lowland northern lowland to middle elevation montane
forms perhaps during the second phase of Andean orogeny in the mid-Tertiary. The southern
lowland form is presently represented by the allopatric O. erythrurum and the other form
represents the ancestor of the widespread, but largely allopatric, O. discicolle and O.
cayennense. Divergence into these latter two species may also be due to continued Andean
orogeny in the Pliocene or Pleistocene. O. cayennense is apparently limited to the lower
montane and lowland Amazon Basin forests, while O. discicolle is distributed in forests of the
surrounding regions at higher elevations. O. discicolle is the only species of Oxelytrum ranging
into Central America and Mexico. This is probably the result of Pleistocene dispersal from
montane areas of northern South America along the island-like montane habitats of Central
America to Mexico. Although many recent interpretations of the evolutionary histories of Latin
American taxa have emphasized the role of Pleistocene forest refugia caused by climatic
changes in South and Central America (reviewed by Simpson and Haffer, 1978; Prance, 1982;
Whitehead, 1976; but for an alternative view see Endler 1982) in promoting speciation we
believe that species origins of all Latin American Silphidae, with the possible exception of
Thanatophilus graniger, predate the Pleistocene.

Nicrophorinae. — The full biogeographic history of Nicrophorus can be presented only after
the Eurasian fauna has been extensively studied. The sister genus of Nicrophorus is
Ptomascopus of eastern Asia whose adults exhibit more primitive states of structural
characters and have not evolved advanced parental care of the larvae, as is found in members of
Nicrophorus (Peck, 1982). The genus Nicrophorus seems to be Eurasian in origin because this
is where the sister genus occurs, and because more species of Nicrophorus occur in Eurasia
(about 60) than in the New World (20). No species are known to occur in Australia or
sub-Saharan Africa.

We suggest that each of the four species groups, plus TV. americanus and TV. pustulatus, may
represent one or more ancestral invasions of North America via the Bering or North Atlantic
Land Bridges during the Tertiary or Pleistocene. Only two species, TV. vespilloides and TV.
investigator , are in both North America and Eurasia, occupying far northern localities. Both
probably occurred on and moved freely across the Bering Land Bridge during low sea stands in
the Pleistocene.

Somewhat more than half of the North American species live in deciduous forests of the
eastern and southeastern United States. We interpret this to be suggestive of the ancestral
habit. Occupation of North America by species ancestral to these can therefore date to the
early Tertiary when such forests were continuous from Asia, across Beringia, to North America
or alternatively from Europe, directly to eastern North America (see Matthews, 1979, 1980 for
review).

Grassland and open shrub habitats seemingly started to become abundant in North America
in the Miocene, as a result of the formation of large rain shadows caused by the uplift of the
Rocky Mountains. We suggest that species occupying these more open and semi-arid
environments are younger and more derived, or represent later ancestral invasions. Except for
the orbicollis group, only these species of more open and arid habitats have entered northern
Mexico, and most of them range no farther south than the edge of the Mexican Plateau. We
suggest only a Pleistocene or Recent occupation of Mexico by these species. The single
exception, TV. mexicanus, reaches Guatemala and El Salvador.

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Only the orbicollis group of Nicrophorus contains species endemic to Latin America. This
group may be the earliest to enter North America from Eurasia due to its apparent primitive
position with respect to other species groups, and its association primarily with hardwood
forested habitats.

The basal stock of the orbicollis group would seem to possess character states similar to
those of N. chilensis (fig. 57). This stock probably reached South America from North and
Central American ancestors in the date Cretaceous or early Tertiary while the two areas were
still connected, or by over-water dispersal as is proposed for some island-hopping mammals in
the early or mid-Tertiary (Darlington, 1957; Simpson, 1980). Alternatively, arrival of the
orbicollis group of Nicrophorus in South America may date from the early Pliocene, and be
contemporaneous with the start of the “Great American Interchange” of faunas. At this time
the Andes were considerably uplifted, but not to their present height. By late Pliocene the
Panama seaway was closed and additional uplift of the Andes formed a continuous temperate
Andean dispersal corridor (Haffer, 1974; Simpson, 1980).

Diversification and endemicity of Central and South American Nicrophorus species argues
for an early rather than late entry into Central and South America. There is no evidence to
suggest that Nicrophorus reached South America from the south, when it was part of the
Gondwanaland supercontinent, although Melville (1981) discusses plant taxa with South
American and Indo-Malayan affinities and interprets them as parts of a fragmented
hypothesized supercontinent called Pacifica. We have not closely examined Old World
relationships of the orbicollis group. They may lie with some southeast Asian species and may
be another example of this type of distribution pattern. Alternatively, such relationships may be
taken as evidence of a more widespread distribution of the members of this species group than
initially thought.

After reaching South America, the Nicrophorus ancestor seemingly remained in forested
habitats and spread down the rising Andes chain to Chile. Here, perhaps due to the later
development of an arid barrier across the Andes, an isolated population now represented as N.
chilensis was formed. The remaining northern South American ancestal stock was again split
with the development of two isolated forest regions separated partly by the high Andes. We
suggest that this produced N. scrutator on the eastern flanks, in Peru, Argentina and Bolivia,
and N. didymus with a range generally in more northerly Andean forests.

We finally suggest that the ancestor of the remaining three species in this group was
distributed throughout warm-termperate or subtropical humid forests from eastern North
America, through Mexico to at least Guatemala, if not Panama. Many organisms, and
especially species or species pairs in tree genera such as Fagus (Beech), Liquidambar
(Sweetgum), Cercis (Redbud), Carpinus (Blue Beech), and Ostrya (Ironwood), show this
former Tertiary distributional connection across the present broad and arid barrier formed by
the Rio Grande depression (see Martin and Harrell, 1957; Rosen, 1978; Allen and Ball, 1980).
The formation of this arid barrier in the late Tertiary allowed for the concurrent isolations of
N. orbicollis in the United States, and populations in Mexico through to Panama which
subsequently gave rise to the allopatric N. olidus in humid upland forests in Mexico north of
the arid lowland barrier of the Isthmus of Tehuantepec, and N. quadrimaculatus in the upland
forests between the Chiapas highlands and western Panama.

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BIOGEOGRAPHIC SYNTHESIS

Latin American silphid species can be grouped into four categories based upon distribution
(modified from Savage, 1982): 1), Widespread-A single species found throughout Middle and
South America; 2), South American-Ten species endemic to South America; 3), Middle
America-One species endemic to Central America, two endemic to Mexico; and 4),
Extratropical North American-Ten species found largely in North America but whose ranges
extend at least in part into Mexico or northern Central America.

Many entomologists do not realize that silphids are present in Latin America. This is
perhaps because the necrophagous niche has been extensively exploited by Scarabaeidae
possibly due to a comparative lack of large herbivores which supply the fecal material for these
beetles in other areas of the world. That the silphid presence in Latin America is of great
antiquity is indicated by the endemic South American assemblage of species. Of the ten South
American endemics, seven belong to the genus Oxelytrum, and three to the orbicollis group of
Nicrophorus. The presence of these two groups in South America, however, is the result of two
different historical pathways.

Oxelytrum , like many South American taxa, appears to owe its presence in South America
to fragmentation of the supercontinent of Gondwanaland during the Cretaceous period. Species
diversification of Oxelytrum occurred during the Tertiary while South America was in
isolation. On the other hand, endemic species of Nicrophorus in South America appear to be
the result of southerly movement from North America into South America perhaps during the
late Cretaceous or early Tertiary while the two land areas were still connected. Alternatively,
dispersal to South America could have taken place during the Tertiary over the island chain
that is now Central America, as proposed for angiosperms by Raven and Axelrod (1974). The
former appears preferrable since the two Middle American endemic Nicrophorus are highly
derived and have a North American sister-group and thus probably originated at some later
time in the Tertiary through a second inter-island dispersal from a northern and not southern
source.

Only a single species of Oxelytrum has dispersed from South America north into Middle
America, but because no divergence has taken place between populations, it is likely that this
was a Pleistocene event. No South American Nicrophorus have reached Middle America. The
other Mexican endemic, Thanatophilus graniger, is readily interpreted as resulting from a
Pleistocene isolation of northern ancestral form.

The ten remaining species are all extratropical North American with the greater part of
their range in temperate North America, and in most instances they just range into the arid
desert lands of northern Mexico. Only two species occur extensively in these arid areas,
reaching as far south as the Neo-Volcanic Sierra. A third species, Nicrophorus mexicanus,
ranges to El Salvador. Most of these species seem to be of recent origin, and probably evolved in
situ in response to increasing aridity and cooling trends in the late Tertiary.

Thus the Latin American silphid fauna originated from a variety of sources during various
time periods since the late Cretaceous. As with many Latin American taxa, South America
possesses a characteristically more primitive and largely endemic assemblage of species.
Central America is largely transitional with species found there either being widespread
tropical or montane endemics of probable Tertiary origin and either direct North or South
American ancestry. Mexico not only possesses endemics of this latter kind, but also species of
more recent Pleistocene origin. A final significant portion of the fauna of Mexico is due to the

Carrion Beetles of Latin America

281

widespread nature of the distributions of species found in the arid southwestern United States.
SUGGESTIONS FOR FUTURE WORK

The present review has attempted to clarify understanding of the classification, phylogeny
and zoogeography of Latin American Silphidae. We hope we have, at least in part, succeeded.
During our work however, we soon came to realize that many aspects of Latin American
silphids have not been well studied and warrant further attention. We think it important to
outline some of the more interesting and potentially useful of these topics here in the hope that
someone will find them stimulating enough to undertake.

First, we think life history studies of species of Latin American Nicrophorus and Oxelytrum
should be undertaken. Not only will this provide missing basic biological information, but also
other life stages including larvae, which can be subsequently used to test reconstructed
phylogenies presented here by adding more characters for analysis. Second, comparative
ecological studies should be undertaken to find out how silphids are interacting with other
necrophagous arthropods in tropical, subtropical and south-temperate habitats and if their roles
in carrion communities are similar regardless of locality. Third, patterns of color variation in
some species of Oxelytrum and Nicrophorus should be examined, and the results considered
within the framework of the Pleistocene forest refugium theory. These species represent needed
further examples that could be used to support or discredit this now highly controversial theory.
Finally, attempts should be made to provide more specimens, particularly of South American
species. This will lead to increased resolution of species distributions and species chorological
relationships. The latter especially, may play an important role in determining species
geographic limits.

ACKNOWLEDGMENTS

We thank all the curators and collectors who responded to our requests for loans of material,
even when they had no specimens. We are especially indebted to those who actually collected
the specimens upon which this review is based, even though space has not allowed us to cite
them individually. Field work and museum visits of S.B. Peck, and field work of R.S. Anderson,
have been partially supported by operating grants from the Natural Sciences and Engineering
Research Council of Canada to S.B. Peck and G.E. Ball. We are indebted to Dr. Gonzalo
Halffter, Dr. Pedro Reyes Castillo, the Instituto de Ecologia de Mexico, and the Instituto de
Biologia de la Universidad Nacional Autonoma de Mexico for facilitating field work in Mexico.

The manuscript and our work with Silphidae has been helped generously by Dr. Alfred
Newton and Dr. Ronald Madge. The manuscript was also improved by data or comments from
G.E. Ball, and S.E. Miller. The distribution maps were partially prepared by Michael Kaulbars
and J.S. Scott.

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Table 1. Classification of known or possible Latin American Silphinae.

Oxelytrum Gistel
lineatocolle group

O. lineatocolle (Laporte)

O. biguttatum (Philippi)

O. apicale (Brulle)

O. anticola Guerin-Meneville
emarginatum group

O. emarginatum (Portevin)

O. erythrurum (Blanchard)

O. cayennensis (Sturm)

O. discicolle (Brulle)

Necrodes Leach

N. surinamensis (Fabricius)

Thanatophilus Leach

T. graniger (Chevrolat)

T. lapponica (Herbst)

T. truncata (Say)

Heterosilpha Portevin
H. aenescens (Casey)

H. ramosa (Say)

Oiceoptoma Leach

O. rugulosum Portevin

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Table 2. Classification of New World Nicrophorus species.

Nicrophorus Fabricius
orbicollis group

TV. chilensis Philippi
TV. scrutator Blanchard
TV. didymus Brulle
TV. orbicollis Say
TV. olidus Matthews
TV. quadrimaculatus Matthews
defodiens group

TV. sayi Laporte
TV. defodiens Mannerheim
TV. vespilloides Herbst
investigator group

TV. tomentosus Weber
TV. hybridus Hatch and Angell
TV. investigator Zetterstedt
TV. nigrita Mannerheim
TV. mexicanus Matthews
marginatus group

TV. marginatus Fabricius
TV. obscurus Kirby
TV. guttula Motschulsky
TV. carolinus (Linnaeus)

Incertae sedis

TV. americanus Olivier
TV. pustulatus Herschel

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Table 3. Analysis of character transformations in Oxelytrum - Ptomaphila lineage of
Silphinae. Relationships only of species of Oxelytrum are shown in fig. 56. Species of
Ptomaphila are not considered. Superscripts on characters indicate that we consider them to be
homoplasious (1) or simple and unique (2). *Terms of Arnett 1944.

character

plesiotypic character state

apotypic character state

pronotum2

without elevated costae

with costae

elytral

undersurface2

lacking long hairs

with long hairs near apical callus

elytral apex2

lacking flange

with flange

head2

with v-shaped swollen area
(chevron) on frons

lacking frontal chevron

elytron2

tricostate

tuberculate

elytral

epipleuron2

narrow

extremely wide

male coxae2

without tubercles or spines

with tubercles or spines

pronotum1

margin orange red, disc black

all black

eyes1

large size

small size

10.

pronotum1

posterior angles obtuse

posterior angles rounded

11.

female genital with styli small, unmodified
coxites2*

with styli large, scooplike

12.

male coxae2

with tubercles

with spines

13.

pronotum1

margins flat or deflexed

margin reflexed

14.

abdominal
segments 4, 51
(of some
males)

without lateral projections

with lateral projections

15.

elytra1

humeri not toothed

humeri each with single tooth

16.

pronotum1

black spot on disc large

black spot on disc reduced

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Table 4. Analysis of character transformations in orbicollis group of species in Nicrophorus.
Relationships of species are shown in fig. 57.

character plesiotypic character state

apotypic character state

1 . elytral long

epipleuron

short

2. pronotum subquadrate

orbicular

3. elytron dorsal surface glabrous, or with

few short hairs

with long hairs

4. metatrochanteiwith spine sharp, well developed

with spine reduced, rounded

5. elytron hairs sparse

hairs dense

6. elytron hairs dense

hairs extremely dense, short

7. elytral glabrous or with few short hairs

epipleuron

densely hairy

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Table 5. Analysis of character transformations in the defodiens group of species in
Nicrophorus. Relationships of species are shown in fig. 58.

character

plesiotypic character state

apotypic character state

larval labial
palpi

bases widely separated

bases narrowly separated

larval labial
palpi

ventral surface of basal segment
sclerotized

ventral surface of basal segment
unsclerotized

adult

pronotum

subquadrate

quadrate

adult

antennomeres
9, 10

lacking white setae on ventral
surfaces

possessing white setae on ventral
surfaces

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Table 6. Analysis of character transformations in the investigator group of species in
Nicrophorus. Relationships of species are shown in fig. 59.

character

plesiotypic character state

apotypic character state

overwintering

stage

adult

prepupa

adult

metasternum

lacking bald spot

with bald spot immediately
posterior to mesocoxae

ventral apex
larval
abdominal
segment 10

unsclerotized

sclerotized

adult

pronotum

sub-quadrate to cordate

quadrate

adult

metasternal

pubescence

yellow

brown

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Table

. Analysis of character transformations in the marginatus group of species in
Nicrophorus. Relationships of species are shown in fig. 60.

character plesiotypic character state apotypic character state

ventral base of sclerotized unsclerotized

larval

abdominal

segment 10

pubescence of glabrous or with few hairs dense yellow hairs

adult

metepimeral
posterior lobe

sternite of outer angles acute outer angles truncate

larval

abdominal

segment 9

hairs on short long

anterior face
of adult
procoxae

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Plate 1. Figures 1, 2. Fig. 1. Habitus of Oxelytrum discicolle , body length 15 mm. Fig. 2. Habitus of Nicrophorus
marginatus , body length 19 mm.

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Plate 2. Figures 3-15. Fig. 3. Head of Oxelytrum discicolle : 1, labrum; c, clypeus; f, frons. Mandibles not drawn. Fig. 4.
Head of Nicrophorus marginatus\ symbols as in fig. 3; Note nicrophorine character of very small second antennal
segment. Fig. 5. Right elytra of Necrodes surinamensis with apical reddish markings. Fig. 6. Short and broadly rounded
pronotal postcoxal lobe (p) of Necrodes surinamensis. Fig. 7. Longer pronotal postcoxal lobe (p) of Thanatophilus
graniger. Fig. 8. Pointed elytral apices of female Heterosilpha ramosa. Fig. 9. Reticulate sculpturing and rounded elytral
apices of male Heterosilpha ramosa. Fig. 10. Rounded elytral apices of male and female Heterosilpha aenescens. Fig. 1 1.
Head of Oiceoptoma rugulosum. Fig. 12. Elytron of Thanatophilus truncatus. Fig. 13. Elytron of female Thanatophilus
lapponicus. Fig. 14. Elytron of female Thantophilus graniger. Fig. 15. Elytron of female Oxelytrum discicolle

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Plate 3. Figures 16-32. Fig. 16. Elytral shoulder of Oiceoptoma rugulosum. Fig. 17. Pronotum of Necrodes surinamensis.
Fig. 18. Dorsal view aedeagus Heterosilpha ramosa. Fig. 19. Right lateral view aedeagus Heterosilpha rantosa. Fig. 20.
Dorsal view aedeagus Heterosilpha aenescens. Fig. 21. Right lateral view aedeagus Heterosilpha aenescens. Fig. 22.
Elytral epipleuron of Oiceoptoma inaequale. Fig. 23. Elytral epipleuron of Oiceoptoma rugulosum. Fig. 24. Plain posterior
margin of female Fifth visible abdominal sternite of Thanatophilus lapponicus. Fig. 25. Crenulate posterior margin of
female fifth visible abdominal sternite of Thanatophilus graniger from (a) Cerro Potosi, Nuevo Leon, and (b) Guerrero
Mills, Hidalgo. Fig. 26a. Head of Oxelytrum apicale Fig. 26b. Head of Oxelytrum biguttatum. Fig. 27. Elytral shoulder
of Oxelytrum erythrurum. Fig. 28. Elytral shoulder of Oxelytrum cayennense. Fig. 29. Elytral apex male Oxelytrum
biguttatum. Fig. 30. Elytral apex of Oxelytrum apicale ; (a) male, (b) female. Fig. 31. Elytral apex male Oxelytrum
anticola. Fig. 32. Elytral apex female Oxelytrum emarginatum.

Plate 4. Figures 33-47. Fig. 33. Elytral apex male Oxelytrum lineatocolle. Fig. 34. Elytral apex female Oxelytrum
lineatocolle. Fig. 35. Elytral apex male Oxelytrum cayennense. Fig. 36. Elytral apex female Oxelytrum cayennense. Fig.
37. Elytral apex male Oxelytrum discicolle. Fig. 38. Elytral apex female Oxelytrum discicolle Fig. 39. Left lateral view of
elytra, epipleuron, and part of thorax of Nicrophorus : mes, metepisternum; meml, metepimeral lobe; ms, metasternum
with anterior area which may be glabrous. Figs. 40-44. Dorsal view of elytron and lateral view of left elytral epipleuron of
Nicrophorus guttula showing variation in size of orange-red fasciae. Fig. 45. Elytron and left epipleuron of Nicrophorus
marginatus. Fig. 46. Long epipleural ridge, Nicrophorus mexicanus. Fig. 47. Short epipleural ridge, Nicrophorus
orbicollis.

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Plate 5. Figures 48-55. Fig. 48. Elytron Nicrophorus marginatus, Coahuila, Mexico, Fig. 49. Epipleuron and elytron,
Nicrophorus quadrimaculatus, Chiriqui, Panama. Fig. 50. Epipleuron and elytral fasciae variation Nicrophorus didymus ,
Cerro Carpish, Huanuco, Peru. Fig. 51. Epipleuron and elytron Nicrophorus chilensis, Malleco, Chile. Fig. 52. Epipleuron
and elytron Nicrophorus olidus, Jalisco, Mexico. Fig. 53. Epipleuron and elytron Nicrophorus scrutator, Tucuman,
Argentina. Fig. 54. Epipleuron and elytron Nicrophorus mexicanus, Durango, Mexico. Fig. 55. Pronotum Nicrophorus
quadrimaculatus, Chiiiriqui, Panama.

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Ptomaphila (Aust., N.Guin.)

O. anticola (5. A.)

O. apicaje (S. A.)

O. biguttatum (S.A.)

O. Ijneatocojle (S.A.)

O. emarginatvm (S.A.)

O. erythrurum (S.A.)

O. cayennensis (S.A.)

O. djscicojje (S.A.,C.A.,
Mex.,Tex.)

Figure 56. Reconstructed phylogeny of Oxelytrum - Ptomaphila lineage of Silphidae. Numbers refer to characters in
Table 3; closed circles indicate apotypic character state.

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N. chijensis (S.A.)

N. scrutator (S.A.)

N. didymus (S. A.)
N. orbicollis (N. A.)

N. ohdus (Mex.)

N. quadrimaculatus

(cX)

Figure 57. Reconstructed phylogeny of species of orbicollis group of Nicrophorus. Numbers refer to characters in Table 4;
closed circles indicate apotypic character state.

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301

N. sayi (Nearc.)

N. vespilloicles (Holarc.)

N. defocliens (Nearc.)

Figure 58. Reconstructed phylogeny of species of defodiens group of Nicrophorus. Numbers refer to characters in Table 5;
closed circles indicate apotypic character state.

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N. tomentosus (Nearc.)

N. hybrialus (Nearc.)

N. investigator (Holarc.)

N. nigrita (Nearc.)

N. mexicanus (Nearc.,
Mex.,C. A.)

Figure 59. Reconstructed phylogeny of species of investigator group of Nicrophorus. Numbers refer to characters in Table
6; closed circles indicate apotypic character state.

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Figure 60. Reconstructed phytogeny of species of marginatus group of Nicrophorus. Numbers refer to characters in Table
7; closed circles indicate apotypic character state; dotted line indicates uncertain placement.

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115° 110° 105° 100°

Map 1 . Distribution of Thanatophilus graniger (black dots) and Heterosilpha ramosa (black squares) in Mexico.

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Map 2. Distribution of Thanatophilus truncatus (black dots) and Thanatophilus lapponicus (square) in Mexico.

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Map 3. Distribution of Oxelytrum biguttatum in southern South America. Map 4. Distribution of Oxelytrum apicale in
Bolivia and northern Argentina. Map 5. Distribution of Oxelytrum anticola in the central and northern Andes.

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307

Map 6. Distribution of Oxelytrum lineatocolle in Chile and Argentina. Map 7. Distribution of Oxelytrum erythrurum
from Bolivia to southern Brazil and Argentina (black dots) and Oxelytrum emarginatum in southeastern Brazil (black
squares).

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Map 8. Distribution of Oxelytrum cayennense in northern South America.

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Map 9a. Distribution of Oxelytrum discicolle in Texas and Middle America.

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Map 9b. Distribution of Oxelytrum discicolle in South America.

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Map 10. Distribution of Nicrophorus marginatus (black dots), Nicrophorus gut tula (open dot), and Nicrophorus nigrita
(black square, Guadelupe Island record not shown) in Mexico.

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Map 1 1 . Distribution of Nicrophorus mexicanus in Mexico to El Salvador.

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Map 12. Distribution of Nicrophorus quadrimaculatus in Chiapas, Mexico and Central America.

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Map 13. Distribution of Nicrophorus didymus in northern Andean South America. Map 14. Distribution of Nicrophorus
chilensis (Black dots) and Nicrophorus scrutator (black squares) in Bolivia, Argentina, and Chile. Question mark
indicates anomalous record.

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Map 15. Distribution of Nicrophorus olidus in Mexico.

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INDEX TO NAMES OF TAXA
(Synonyms in italics)

FAMILY GROUP TAXA
Nicrophorinae, 265, 278
Silphidae, 248-249
Silphinae, 249, 253, 272, 276

GENERA AND SUBGENERA
Aclypea Reitter, 276

Heterosilpha Portevin, 252-253, 271, 276
Necrodes Leach, 252-253, 276
Necrophila Kirby and Spence, 276
Nicrophorus Fabricius, 249, 252, 265,
273-275, 278-280
Oiceoptoma Leach, 252, 254, 276
Oxelytrum Gistel, 252, 257, 271-272, 277,
280

Ptomaphila Portevin, 271-272
Ptomascopus Kraatz, 273, 275, 278
Thanatophilus Leach, 252, 255, 276

SPECIES AND SUBSPECIES
aenescens (Casey), Heterosilpha, 253-254
aenescens Casey, Silpha, 254
aequinoctiale Gistel, Oxelytrum, 262
americanus Olivier, Nicrophorus, 276, 278
analis (Chevrolat), Hyponecrodes, 263
analis Chevrolat, Necrodes , 263
andicola (Guerin-Meneville),
Hyponecrodes , 259
anticola (Guerin-Meneville),
Hyponecrodes , 259

anticola (Guerin-Meneville), Oxelytrum,
257, 259, 277

anticola Guerin-Meneville, Silpha , 259
apicale (Brulle), Oxelytrum, 257-258, 277
apicalis (Brulle), Hyponecrodes , 258
apicalis Brulle, Silpha , 258
biguttata (Philippi), Silpha , 258
biguttatum (Philippi), Oxelytrum,
257-258, 277

biguttatus (Philippi), Hyponecrodes , 258
biguttatus (Philippi), Paranecrodes , 258
biguttatus Philippi, Necrodes , 258

biguttula Fairmaire and Germain, Silpha ,
258

biguttulus (Fairmaire and Germain),
Necrodes , 258

californicus Mannerheim, Thanatophilus,
256

carolinus (Linnaeus), Nicrophorus, 276
cayennense (Striirm), Oxelytrum, 258,
261-262, 278

cayennensis (Sturm), Hyponeceodes, 262
cayennensis Sturm, Silpha , 262
chilensis Philippi, Nicrophorus, 266, 269,
279

didymus Brulle, Nicrophorus, 266, 268,
279

didymus peruvianus Pic, Nicrophorus,

268

discicolle (Brulle), Oxelytrum, 257-258,
263, 273, 278

discicollis (Brulle), Hyponecrodes, 263
discicollis Brulle, Silpha, 263
discicollis discretus Portevin,
Hyponecrodes, 263
discicollis elongatus Portevin,
Hyponecrodes, 263

distinctus Grouvelle, Nicrophorus, 274
emarginata (Portevin), Silpha, 260
emarginatum (Portevin), Oxelytrum, 257,
260, 277

emarginatus Portevin, Hyponecrodes, 260
erythrura (Blanchard), Hyponecrodes,

261

erythrura (Blanchard), Silpha, 261
erythrurum (Blanchard), Oxelytrum, 257,
261-262, 278

erythrurus (Blanchard), Hyponecrodes,
261

erythrurus melancholicus Portevin,
Hyponecrodes, 26 1
erythrurus melanurus Portevin,
Hyponecrodes, 26 1
erythrurus pygialis Portevin,
Hyponecrodes, 26 1

flexuosus Portevin, Nicrophorus, 268

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gayi Solier, Necrodes, 260
graniger (Chevrolat), Thanatophilus,
255-256, 277-278, 280
granigera Chevrolat, Oiceoptoma, 256
guttula Motschulsky, Nicrophorus,
266-267

heurni Portevin, Ptomascopus, 274
inaequale , Oiceoptoma, 255
inaequalis rugulosa Portevin, Silpha , 254
investigator Zetterstedt, Nicrophorus, 275,

278

kieticus Mroczkowski, Nicrophorus, 274
lapponica Herbst, Silpha , 256
lapponicus (Herbst), Thanatophilus,
255-257, 277

lineaticollis (Laporte), Hyponecrodes, 260
lineatocolle (Laporte), Oxelytrum, 257,
260, 277

lineatocollis (Laporte), Hyponecrodes ,

260

lineatocollis Laporte, Silpha , 260
marginatus Fabricius, Nicrophorus,

265- 266, 276

mexicanus Matthews, Nicrophorus,

266- 267, 275-276, 278, 280
microps Sharp, Silpha , 259
montezumae Matthews, Necrophorus, 266
nigrita Mannerheim, Nicrophorus, 266,

268, 275

occidentale Gistel, Oxelytrum , 262
olidus Matthews, Nicrophorus, 266, 269,

279

opaca (Portevin), Silpha , 261
opacum , Oxelytrum, 262
opacus Portevin, Hyponecrodes , 261
opacus tristis Portevin, Hyponecrodes ,

261

orbicollis , Nicrophorus, 274, 279
podagricus Portevin, Nicrophorus, 274
pronotus Gistel, Necrodes, 250
pustulatus Herschel, Nicrophorus, 276,

278

quadricollis Gistel, Nicrophorus, 250, 279
quadrimaculatus Matthews, Nicrophorus,
266, 268

ramosa (Say), Heterosilpha, 253-254

ramosa Say, Silpha , 254
rugulosum (Portevin), Oiceoptoma,
254-255

scrutator Blanchard, Nicrophorus, 266,
270, 279

surinamensis (Fabricius), Necrodes, 253
surinamensis Fabricius, Silpha , 253
truncata (Say), Philas, 255
truncata Say, Silpha , 255
truncatus (Say), Thanatophilus, 255
vespilloides Herbst, Nicrophorus, 275, 278

Quaest. Ent., 1985, 21 (3)

THE TIGER BEETLES OF ALBERTA (COLEOPTERA: CARABIDAE, CICINDELINI)1

Gerald J. Hilchie
Department of Entomology
University of Alberta
Edmonton, Alberta T6G 2E3.

Quaestiones Entomologicae
21:319-347 1985

ABSTRACT

In Alberta there are 19 species of tiger beetles ( Cicindela ). These are found in a wide
variety of habitats from sand dunes and riverbanks to construction sites. Each species has a
unique distribution resulting from complex interactions of adult site selection, life history,
competition, predation and historical factors. Post-pleistocene dispersal of tiger beetles into
Alberta came predominantly from the south with a few species entering Alberta from the north
and west.

INTRODUCTION

Wallis (1961) recognized 26 species of Cicindela in Canada, of which 19 occur in Alberta.
Most species of tiger beetle in North America are polytypic but, in Alberta most are
represented by a single subspecies. Two species are represented each by two subspecies and two
others hybridize and might better be described as a single species with distinct subspecies.
When a single subspecies is present in the province morphs normally attributed to other
subspecies may also be present, in which case the most common morph (over 80% of a
population) is used for subspecies designation.

Tiger beetles have always been popular with collectors. Bright colours and quick flight make
these beetles a sporting and delightful challenge to collect.

The purpose of this paper is to provide a guide to the tiger beetles occurring in the province
of Alberta. Information on life history, species recognition, habitat preference, collecting sites
and a brief synopsis of biogeographical considerations for interpretation of present distribution
patterns have been included.

LIFE HISTORY

Adults

Tiger beetles are capable fliers and quick on their legs, being able to escape rapidly when
disturbed. Most tiger beetles are diurnal, preferring bright sun, but, some are active at night.
Adults of one species in Alberta, Cicindela lepida, are normally active during the day, but, on
warm nights, they will resume hunting activities shortly after sunset. Most tiger beetles at the
onset of night or inclement weather dig shallow burrows for refuge.

Alberta Cicindela can also be divided into two categories based on the life span of the adult:
those species having long lived adults which overwinter (spring-fall) and those in which adults

‘Portions of the text were published in Alberta Naturalist 14: 105-1 1 1, 1984. Tiger Beetles of
Alberta- life history and key.

320

Hilchie

live for a single summer (summer). In spring-fall species, adults freshly emerged from pupae
prepare for winter in late summer or early autumn by excavating a deep burrow. Wintering
quarters vary in depth, depending on the species and the nature of the soil. Cicindela repanda
adults for example will dig burrows 15 to 21 cm deep in clay soils and up to 56 cm deep in
sandy soils, while those of C. formosa have been recorded digging burrows down to 109 cm in
sand (Criddle 1907).

Spring-fall-adults are sexually immature in the fall (Willis 1967) and appear to need winter
chilling to break reproductive diapause. Wintered adults lay eggs the following spring.

Summer-adults do not diapause and are reproductively active within days of emerging from
the pupa.

Tiger beetles can be grouped ecologically. In Alberta, adults of species found along streams
tend to have dark elytra with well defined maculations (C. repanda, C. hirticollis , C.
duodecimguttata, C. oregona ), those found on dark soils often have reduced maculation and
are black (C. nebraskana, C. purpurea purpurea, C. longilabris ), those found along margins of
sand dunes tend to be brightly coloured (C. formosa formosa, C. lengi, C. scutellaris ) and
those found in the open on drifting sand are very pale and blend in with the sand (C. limbata
nympha, C. lepida, C. formosa gibsoni ).

Food for adults consists of other insects. Almost any insect will be taken with the exception
of certain bugs and of prey too large or small to handle. Large tiger beetles will prey
opportunistically on members of other smaller species.

There are few effective predators of tiger beetles. Dragon flies and robber flies have been
observed catching tiger beetles in flight (Graves 1962, Lavingne 1972) and birds occasionally
prey on them. For example, droppings of Ringbilled Gulls (at Gull Lake, Alberta) contained
pieces of elytra from C. repanda C. hirticollis (personal observation, 1973). Normally the quick
movements, rapid flight and cryptic colouration of tiger beetles make their capture by people
and other predators difficult.

Eggs and Larvae

Eggs are deposited in spring through summer, depending on species. The female makes a
small hole in the ground with her ovipositor and deposits a single egg. The egg hatches a few
weeks later into a first instar larva. This larva first enlarges its hole and then positions its head
and thorax at the burrow entrance and waits for its first meal (Fig. 1). If prey is readily
available, feeding will last for a few weeks, then the larva plugs its burrow entrance and
remains dormant until late summer. Feeding then resumes for a few weeks or until the onset of
cold weather. The larva diapauses until spring at which time feeding resumes (Hamilton 1925,
Willis 1967).

Depending on the species, larvae may reach maturity (third instar), pupate and emerge as
adults by the end of June (summer-adult species). Alternatively, larvae may stop feeding
during mid-summer, resume feeding in late summer, then pupate and emerge as adults in late
August or early September (spring-fall-adult species). In all species at least one larval instar
passes through winter, but only in spring-fall species do adults survive a winter (Criddle 1907,
1910, Hamilton 1925, Shelford 1908, Willis 1967).

Tiger beetle larvae, like adults, are carnivorous. Large mandibles provide an effective means
of subduing prey (Fig. 2). Larvae of C. formosa may dig a pit at the burrow entrance, that may
be used to trap ants and other small insects (Criddle 1910). Larvae of most species do not dig a
trap but lunge and seize prey near the burrow entrance. Prey is pulled into the burrow and the

Tiger beetles of Alberta

321

larva then consumes it in relative safety. Strong abdominal spines anchor the larva to the
burrow wall (Fig. 1) preventing accidental dislodgement by large prey. If the prey is too large,
the larva releases the would be victim and retreats by dropping to safety at the bottom of its
burrow.

Larval burrows vary in depth according to species, soil type and larval age. Depths range
from a few cm for larvae of Cicindela repanda and C. duodecimguttata along stream margins
to depths in excess of 3 m for those of C. lepida and C.formosa on sand dunes.

Few predators attack tiger beetle larva. Some Beeflies ( Anthrax spp., Bombiliidae) lay eggs
near burrow entrances. The hatching fly maggot may then locate and parasitize the tiger beetle
larva. Parasitized larvae fail to complete development, dying during or just before pupation.
Swan (1975) found up to 7% of a population of Cicindela scutellaris larvae to be parasitized.

Other larval parasites are members of the tiphiid wasp genus Methocha Latreille. These
small wingless wasps wander in areas occupied by tiger beetle larvae. When siezed, the wasp
immediately stings the beetle larva in the soft gular cuticle under its head, causing temporary
paralysis; an egg is then deposited and the wasp wanders off to be grabbed by another victim.
The narrow, elongate shape and armored cuticle of the wasp prevents it from being pierced by
the larva’s mandibles and allow the wasp to manoeuvre and sting the larva. The wasp larva
hatches and remains attached as an external parasitoid. When the host larva pupates, the wasp
larva then begins to actively feed, consuming and killing the pupa from inside out.

Pupae

Pupation occurs in a specially prepared chamber opening into the side of the larval burrow.
In summer-adult species, the larvae feed in spring before molting into pupae. The larvae of
spring-fall adult species feed for an extended period in the spring and may feed in late summer
before molting into pupae (August). Pupation lasts for a few weeks. Summer-adult species
emerge as adults in late June and early July, while spring-fall-adult species emerge as adults in
late August through early September. The spring-fall-adults feed for a short time before
entering winter diapause, with breeding occurring the following spring.

Possible factors influencing selection for summer or spring-fall species

Maintainence of summer and spring-fall species involves a combination of past- and
presently acting selective pressures. Habitat, an obvious feature, may at first glance be
implicated as a factor influencing life history. However, the four summer species in Alberta
occupy different habitats. Members of C. lepida live on sand dunes, those of C. nevadica on
alkaline soil, those of C. punctulata on gravelly prairie soil and those of C. terricola on clay or
dark loamy grassland soils. Similar diversity of habitats are found in spring-fall species.

Interactions between tiger beetle species appear important. Interspecfic contact may involve
competitive interactions for adult hunting sites, food, larval burrow sites, oviposition sites,
wintering sites and predation on or by other tiger beetles. For example, the spring-fall species,
C. formosa, C. lengi , C. scutellaris and C. limbata nympha, and the summer species, C. lepida
adults share overlapping habitat on the Empress sand dunes of Alberta. Habitat partitioning
appears to be expressed as adult hunting sites. Food selection is similar for all species, the
beetles feed on any insect of suitable size (see Willis (1967) for diet of saline habitat tiger
beetles). Adults of C. lengi , C. lepida and C. scutellaris live in the margins of sand dunes and
adjacent grasslands, while those of C. /. nympha live in the margins and out onto open sand.
Beetles of C. /. nympha appear to reduce detrimental interaction (predation) by moving away

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Hilchie

from dune margins when tiger beetles of other species are present. Temporal partitioning of
habitat also effectively reduces intraspecfic interactions for beetles of C. lepida. Adults of C.
lepida occupy the same habitat as those of C. /. nympha, but at a different time of year, mid
July versus May, June and August. This temporal isolation effectively removes these beetles
from direct competition for food and predation with larger spring-fall tiger beetles. In regions
south of Alberta, several summer species may occur in similar habitats, When this occurs
additional temporal shifts may occur in population abundance (Willis, 1967).

The other, summer-adult species in Alberta do not appear to have as complex interactions
with other tiger beetle species as does C. lepida. It is possible that being a summer-adult species
has supplied a competitive advantage to members of the species either in the past or in a portion
of the species present range. Following post pleistocene dispersal, these interactions may no
longer occur in Alberta.

There must also be some advantage to be gained from being a spring-fall species. Upon
examining distribution maps, one notices immediatlely that spring-fall species have ranges
which extend north into cooler climatic zones. Prolonged larval development with a long
feeding period allows for greater success in obtaining adequate nutrition for completion of
development. In some spring-fall species, larvae may take several seasons to complete
development. Interactions between other tiger beetle species are limited with one or two species
occupying similar but not identical habitats.

Each tiger beetle species has its own unique history. Variations do occur, however, with
some species not fitting neatly into a defined pattern. Further investigations are required to
elucidate precisely what factors are involved in maintaining these two life history types in
Alberta.

KEY TO THE TIGER BEETLES OF ALBERTA

This key is adapted from Wallis (1961), Freitag (1965) and Willis (1968). When two
subspecies are present in Alberta, information regarding their separate recognition is included.
See Fig. 3 for details on nomenclature of elytral maculation. When examining for characters
such as presence of microserrations, a magnification of 40X may be required. Most of the other
characters can be seen with the unaided eye or with a 10X hand lens.

1 Frons glabrous or with two supraorbital setae (Fig. 6) 2

T Head covered with hairs or with clusters of hair on the inner margin of

each eye (Fig. 5) 6

2 ( 1 ) Elytral apices serrate with a row of blue or green foveae (Fig. 31)

C. punctulata, p. 331

2' Elytra apices not serrulate, without a row of metallic blue to green foveae 3

3 (2) Small beetles, less than 1 5 mm, labrum short (length parallel to long axis

of body less than one half its width) C. terricola 4

Larger beetles, greater than 15 mm, labrum long 5

4 (3) Marginal band complete (Fig. 25) C. terricola cinctipennis, p. 332

4' Marginal band reduced (Fig. 26) C. terricola imperfecta , p. 332

5 (4) Elytra shallowly punctate or sculptured into waves, shiny between

punctures or on crests of waves, abdominal sternites dull black (Fig. 9)
C. nebraskana, p. 328

Tiger beetles of Alberta

323

6 (1)
6'

7 (6)

8 (7)
8'

9 (8)
9'

10 (9)
10'

11 (7)
11'

12 (11)
12'

13 (12)
13'

14 (12)
14'

15 (11)
15'

16 (15)
16'

17 (16)
17'

Elytra granulate, dull or with slight sheen near base, abdominal sternites

metallic green to violet (Fig. 10) C. longilabris, p. 327

Head more or less hairy (Fig. 4), at least a few hairs on frons, if abraded

punctures mark former location of hair

Head with only clusters of hair on anterior inner margin of each eye (Fig.

Hair on head, thorax and abdomen decumbent

Hair more or less erect on at least part of body (beetles killed in liquid may

have matted hair), markings usually well defined

Dark elytral markings not sharply defined, legs pale, on sand dunes (Fig.

32) C. lepida, p. 333

Dark elytral markings more sharply defined, legs dark

Elytral dark markings reduced, mostly pale (Fig. 13)

C. limbata nympha, p. 326

Elytral dark markings not greatly reduced, more typical banding pattern .
Middle band straight and oblique, not “hooked” at end, pale markings

heavy, wide (Fig. 12) C. limbata hyperborea, p. 326

Middle band sinous, curved, hooked at end, pale markings not heavy; alkali

washes (Fig. 30) C. nevadica, p. 332

Marginal line joined or touching humeral lunule

Marginal line separated, not touching humeral lunule, often greatly

reduced

Humeral lunule “c” shaped

Humeral lunule oblique, pale markings wide

Genae glabrous (Fig. 7), posterior tip of humeral lunule (when present)
with slight anterior hook (marginal line may touch apical lunule, Fig. 11)

C. hirticollis, p. 326

Genae setose (if hairs abraded, punctures mark their former position, Fig.
8), marginal line usually separated from apical lunule (Fig. 14)

C. repanda , p. 324

Length greater than 1 5 mm, line of humeral lunule obliterated totally or in

part by marginal band (Figs. 21 & 22) C.formosa, p. 328

Length less than 15 mm, humeral lunule long, spur may almost touch

middle band (Fig. 23) C. lengi, p. 330

Marginal line greatly reduced or absent, humeral lunule absent or reduced

to spots (Figs. 27, 28 & 29)

Marginal line present (Figs. 18, 19 & 20), obvious spur, humeral lunule

present

Middle band wide long, apical end not markedly curved, color green to

violet (Fig. 28) C. decemnotata , p. 329

Elytral markings thin, light, middle band more strongly curved, shorter
than width of elytron

Post humeral spot usually absent, middle band widely separated from
margin, color green or black (Fig. 27) C. purpurea , p. 328

Post humeral spot usually present, middle band narrowly separated or
touching margin, color red or greenish, middle band transverse, often

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324

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strongly curved at apical end (Fig. 29) C. splendida limbalis , p. 329

18 (15) Genae glabrous (Fig. 7), elytra with greasy appearance, pale markings

heavy, on alkali soils and washes (Fig. 20) C.fulgida, p. 330

18' Genae with hairs or or setigerous punctures (Fig. 8) 19

19 (18) Humeral lunule “c” shaped, scape (basal segment) of antenna with few

hairs (Fig. 18) C. duodecimguttata, p. 325

19' Humeral lunule oblique, scape of antenna hairy (more than 10 hairs) (Fig.

19) C. tranquebarica, p. 331

20 (7) Elytra not serrulate, non punctate, in Alberta red/green elytra without pale

maculations (Fig. 24) C. scute llaris, p. 330

20' Elytra punctate, serrulate, color brown, blue, olive, maculations typical to

reduced C. oregona 21

21 (20) Pleura of thorax blue/purple, elytra brown, green, blue, maculations

narrow, pronotum brown (Fig. 15) C. oregona oregona, p. 325

21' Thoracic pleura coppery, elytra dark brown, maculations narrow (Fig. 16)
C. oregona guttifera, p. 325

TIGER BEETLES OF ALBERTA

1. Cicindela repanda Dejean (Figs. 14 & 33)

C. repanda repanda Dejean

Recognition. — (Fig. 14) These beetles resemble members of C. duodecimguttata.
Separation between them is based on configuration of the marginal band: in C. repanda the
band is continous or narrowly separated from the humeral lunule (Fig. 14), whereas in C.
duodecimguttata the band has a wide gap (Fig. 18). For positive identification the male
genitalia must be examined (see Freitag (1965) for method). Cicindela duodecimguttata, C.
repanda, C. hirticollis, C. limbata and C. oregona form part of the Maritima group and many
species in this group resemble each other.

Habitat. — These beetles inhabit sand, gravel and clay soils with sparse vegetation adjacent
to streams and rivers. Adults may be found running near the water over patches of mud.
Wintering grounds for the adults may be some distance from their summer haunts. Winter
burrows are made in bare dry hillsides (Criddle 1907). Larvae may be found scattered through
vegetation near stream or pond margins. This species has a two year life cycle, with the third
instar larva passing through the first winter and the adult the second (Hamilton 1925).

Localities. — (Fig. 33) Athabasca River (5 km east, Chain Lakes), Barker Lake, Barrier
Reservoir, Brazeau River (near Lodgepole), Calgary, Chin (4.8 km south), Crimson Lake,
Clyde (6.5 km east), Deadwood (banks of Peace River), Devon, Dilberry, Drayton Valley,
Dunvegan, Edmonton, Empress (11 km south), Fawcett, Flatbush (Pembina River), Fort
MacKay, Fort McMurray, Garth, Gem, Gibbons, Green Island (sic!, = Verte Island), Gull
Lake, House River, Jenner, Lesser Slave Lake, Little Smoky River, Lethbridge, Medicine Hat,
McGrath, Meikle River (Mackenzie Highway), Milk River (junction with Lost River), North
Saskatchewan River (near Rocky Mountain House), Patricia (near), Peace River, Pembina
River (near Lodgepole), Red Deer, Red Deer River (near Bindloss), Saunders, Smoky River,
Wainwright, Wapiti River (south of Grande Prairie).

Tiger beetles of Alberta

325

2. Cicindela duodecimguttata Dejean (Figs. 8, 18 & 34)

Recognition. — (Figs. 8 & 18) This species is similar to C. repanda but is more closely
related to C. oregona and hybridizes with it along the Rocky Mountain Foothills of Alberta
(Freitag, 1965). Populations of C. duodecimguttata occur east of the foothills and populations
of C. oregona to the west along mountain valleys. Hybrid populations have markings
intermediate to those of C. oregona and C. duodecimguttata (Freitag 1965) (Fig. 17). The
humeral lunule may be narrowly broken or expressed as a spot and the marginal line is of
variable length.

Habitat. — These beetles live close to pond and stream margins. When in association with C.
repanda , beetles of C. duodecimguttata move away from the water’s edge reducing habitat
overlap. Adults and larvae winter in burrows 1.2 to 2 m back from the stream or pond margin.
If the water rises in spring before beetles are active, large numbers may perish (Criddle 1907).
Members of this species have a two year life cycle similar to that of C. duodecimguttata.

Localities. — (Fig. 34) Andrew, Beaverhill Lake, Bilby, Brazeau River (near Lodgepole),
Calgary, Chin, Clyde (6.5 km east), Cooking Lake, Crimson Lake, Cypress Hills, Doussal,
Drayton Valley, Edmonton, Fallea, Flatbush, Fort Chipewyan, Fort McMurray, Fort MacKay,
Gull Lake, Happy Valley (Porcupine Hills), Halfwayhouse, Jenner, Lake Cardinal, Lesser
Slave Lake, Lethbridge, Louis Bull Reservation, Medicine Hat, North Saskatchewan River
(near Rocky Mountain House), Police Lake, Redwater, Saunders, Smith-Fitzgerald Road (km
11), Stirling Lake, Tilley, Tofield, Vilna, Wabamum.

3. Cicindela oregona LeConte (Figs. 15, 16 & 35)

C. oregona oregona LeConte (Fig. 15)

C. oregona guttifera LeConte (Fig. 16)

Recognition. — (Figs. 15 & 16) These beetles are distinguished from those of related species
by the presence of small groups of hairs on the inner margin of each eye. Markings are similar
to those of C. duodecimguttata and C. repanda. This species hybridizes with C.
duodecimguttata in the foothill region (Freitag 1965) (see discussion under C.
duodecimguttata) and in the Northwest Territories.

Wallis (1961) called the Albertan populations C. oregona guttifera. On examination of
Albertan material I found these beetles to be intermediate between C. oregona oregona and C.
oregona guttifera which agrees with Freitag (1965). Members of the subspecies guttifera are
characterized by a humeral lobe represented by two large spots, the dorsal surface is more or
less olive with a metallic lustre, thoracic pleura are coppery and the ventral surface is bicolored.
The elytral spine is small and serrulations of the apex are weak. Representatives of the
subspecies oregona are similar to guttifera but the elytral spine and serrulations are well
developed and the thoracic pleura are metallic blue like the ventral surface. Imprecise
definition of Albertan populations is due to hybridization of C. o. oregona with C. o. guttifera
and C. duodecimguttata. The specimen collected in the Peace River area was clearly C. o.
guttifera.

Habitat. — Individuals of Cicindela oregona live along margins of streams and lakes on clay
or sandy soils with little vegetation cover. The habitat may be shared with members of C.
repanda , C. duodecimguttata , and C. hirticollis.

Localities. — (Fig. 35) C. o. oregona X C. o. guttifera: Athabasca Falls, Banff, Carbondale,
Castle River, Highwood River, Hillcrest, Kootenay Plains, Laggan ( = Lake Louise), New
Dayton (1.6 km east), North Saskatchewan River (near Nordegg), Waterton.

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C. o. guttifera : Green Island (sic!, = Verte Island).

4. Cicindela hirticollis Say (Figs. 11 & 36)

Recognition. — (Fig. 11) Members of this species are distinguished from others of the
Maritima group by the strongly “C” shaped humeral lunule on the elytra. In other respects they
are similar to members of C. repanda.

C. hirticollis is represented by one form throughout most of Canada. Subspecies recognition
is based on width of elytral maculations. Most specimens have wide markings and are called C.
h. ponderosa Thoms. The problem is that, within local populations, many individuals may have
narrow maculations and could be assigned to another subspecies. Dull brown elytra are the
norm, but, on the Athabasca drainage, some blue beetles have been collected. Due to the wide
geographic range and nature of variation I do not feel that assigning the Albertan populations
to a particular subspecies is appropriate.

Habitat. — These tiger beetles are found on light colored beach sands with little or no
vegetation. The habitat may be shared with other members of the Maritima group. This species
appears to have summer-adults.

Localities. — (Fig. 36) Athabasca River, Calgary, Crimson Lake, Dunvegan, Empress, Fort
McMurray, Gull Lake, Jasper, Jasper Lake, Lac La Biche, Lesser Slave Lake, Medicine Hat,
Red Deer, Red Deer River (near Bindloss), Snaring River (Jasper National Park).

5. Cicindela limbata Say (Figs. 4, 12, 13, 37 & 38)

Cicindela limbata nympha Casey (Figs. 4, 12 & 37)

Cicindela limbata hyperborea LeConte (Figs. 13 & 38)

Recognition. — (Figs. 4, 12 & 13) Two subspecies, very different in appearance, occur in
Alberta.

Individuals of Cicindela limbata nympha are distinguished by the pale elytra with reduced
dark markings. The marginal band is expanded to cover most of the elytra leaving a narrow
dark band down the centre. A brownband identifies the subspecies nympha , a green band C. /.
limbata. Nearly all of the specimens examined were marked with the brown; hence the
Albertan populations are assigned to C. /. nympha.

Specimens of Cicindela limbata hyperborea do not look like those of the southern
subspecies. Instead they resemble other members of the Maritima group of which C. limbata is
a member. Markings are of typical tiger beetle design with a brown elytral ground color. The
greatly thickened elytral bands distinguish members of this subspecies from those of all other
Maritima group species found in Alberta.

Habitat. — These beetles prefer sandy blowouts or sand dunes which are sparsely vegetated.
When the species is present it is usually very abundant. Adults are known to winter in loose
sand. Beetles of Cicindela limbata nympha occur on prairie sand dunes and blowouts, whereas
those of C. /. hyperborea are found in similar habitats in the boreal forest. Individuals of the
northern subspecies also occur in disturbed sandy areas along roadways. In Manitoba C. /.
nympha has a three year life cycle (Criddle 1907, Hamilton 1925).

Localities. — C. /. nympha-. (Fig. 37) Blindman River, Bruderheim, Calgary, Chauvin,
Claysmore, Clyde, Clyde (6.5 km east), Crimson Lake, Czar, Dilberry, Egerton, Edmonton,
Empress, Empress (1 1 km south), Gull Lake, Hondo, Lesser Slave Lake, Nestow, Orion, Opal,
Pakowki Lake, Ponoka (5 km south), Red Deer, Ribstone, Rochester, Stauffer, Tawatinaw,
Winterburn.

Tiger beetles of Alberta

327

C. /. hyperborea : (Fig. 38) Barber Lake, Fort Chipewyan, Fort MacKay, Fort MacKay (10
km south), Fort Mackay (8.5 km east, north of Athabasca River Bridge), Fort McMurray,
Fort Smith (N.W.T., northern border of Alberta), Gregorie Lake.

6. Cicindela longilabris Say (Figs. 10 & 39)

Cicindela longilabris longilabris Say

Recognition. — (Fig. 10) This tiger beetle of the foothill and boreal regions is closely related
to C. nebraskana. Correct identification is often difficult. Species determination is based on
microstructure of the elytra and habitat preference when known.

Adults of are characterized by smooth or very slightly waved sculpture on the elytra. The
surface is covered with minute granules giving the beetle a dull lustre. If shiny areas do occur,
these are small and restricted to the crests of the waves. The lustre may be due to wear on the
elytra in older beetles.

Cicindela nebraskana adults have a larger shiny area, giving the elytra more lustre. A series
of granulate punctures surrounded by a glossy mesh or well developed series of waves and
ridges are characteristic of this species. Members of Cicindela nebraskana live in prairie
grasslands and of C. longilabris in forest clearings and meadows.

In the Alberta foothills, prairie grassland extends along valley bottoms and on
southwest-facing hill sides into the mountains in several places (e.g. Bow River Valley,
Crowsnest Valley). In these areas the ranges of these two species overlap. Hybridization has not
been observed to occur between them. In localities of overlap, they appear to partition the
habitat. Individuals of Cicindela longilabris stay near clearings on sandy soils with conifer trees
and of C. nebraskana on clay soils of the valley grasslands.

Hatch (1953) did not recognize C. montana LeConte ( = C. nebraskana) as a valid species
but rather as a variation (abberation) of C. longilabris. My experience with these species
indicates that they are morphologically very similar but can be distinguished and that they have
different habitat preferences. Populations of C. longilabris in Alberta are composed primarily
of black beetles, with slender feeble markings, hence assignment to the subspecies C.
longilabris longilabris. However, there are individuals of other phenotypes present. Beetles
with heavy markings and a bronzed color are of the laurenti Schaupp phenotype, and those
with slender markings with a vivid green are of the perviridis Schaupp phenotype.

Habitat. — Cicindela longilabris adults prefer sandy areas such as ridges and blowouts in
conifer forests. Adults are found along sandy forest paths and road sides, and over winter.

Localities. — (Fig. 39) Banff, Barber Lake, Barrier Reservoir, Beaverlodge, Beauvallon,
Calling River Ranger Station, Canmore, Cline River, Crimson Lake, Coleman, Crownest Lake,
Drayton Valley, Edmonton, Exshaw, Fairview (10 km southeast), Fedora, Fortress Mountain,
Fort MacKay, Fort MacKay (8.5 km east, north of Athabasca River Bridge), Fort McMurray
(22.4 km north), George Lake, Gorge Creek, Green Island (sic!, = Verte Island), Hargwen,
Hinton, Kananaskis Lakes, Kootenay Plains, Marlboro, Millarville, North Saskatchewan River
(near Nordegg), Opal, Peace River, Pembina River (near Fawcett), Poachers Landing (Tp.69
Rge.19 W.4), Prairie Bluff Mountain, Robb, Rocky Mountain House, Sand Hill Lake,
Saunders, Waterton, Wapiti River (south of Grande Prairie), Whirlpool River, Winterbum.

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7. Cicindela nebraskana LeConte (Figs. 6, 9 & 40)

Cicindela nebraskana nebraskana LeConte

Recognition. — (Fig 6 & 9) Adults of this species are slightly smaller than those of C.
longilabris and have reduced elytral maculations. Ground color of the elytra is black to slightly
bronzed. Features for separating specimens of C. nebraskana from those of C. longilabris are
discussed under C. longilabris.

The Alberta subspecies is C. nebraskana nebraskana. Many subspecfic names have been
created to describe minor differences in various populations. In Alberta the beetles examined
were within the normal range of variation for the subspecies nebraskana. These are smooth
black to slightly bronzed beetles from prairie grasslands.

Habitat. — These beetles prefer open bare areas between clumps of grass and earth mounds
made by ground squirrels. Cicindela nebraskana occurs throughout the prairie grasslands on
heavy clay soils, and overwinter as adults.

Localities. — (Fig. 40) Barrier Reservoir, Calgary, Carbondale River (junction of Lost
Creek), Chin (5 km south), Coronation, Del Bonita, Dorothy, Drumheller, Evans-Thomas
Creek, Empress (11 km south), Exshaw, Kananaskis River (Ribbon Creek), Frank, Gorge
Creek, Ghost Dam, Highwood River, Hilda, Jenner, Lethbridge, Manyberries, Medicine Hat,
Milk River (16 km north of Aden), Milk River (junction with Lost River), Oyen (4.8 km
south), Pincher, Porcupine Hills, Prairie Bluff Mountain, Scandia, Seebe, Strathmore,
Standard, Stettler, Steveville, Taber.

8. Cicindela formosa Say (Figs. 21 & 41)

Cicindela formosa formosa Say

Recognition. — (Fig. 21) This is the largest of the Albertan tiger beetles. The shape of the
humeral lunule is distincitive. Superficially, adults of C. formosa resemble adults of C. lengi. In
Alberta the species is represented by the typical subspecies C. formosa formosa. Adults have a
bright metallic lustre on the ventral surface, with the pigmented areas of the elytra red to
red-purple.

Another subspecies, C. formosa gibsoni Brown (Figs. 22 & 42), occurs a few kilometers east
of the Alberta boundary, in the Great Sand Hills of Saskatchewan. Characteristics of these
beetles are: a metallic venter and reduced elytral pigmentation with the marginal band
expanded to cover most of the elytra. This subspecies should be watched for in sandy areas near
the Saskatchewan boundary.

Habitat. — Beetles of Cicindela formosa inhabit sandy blowouts and marginal areas of
active sand dunes in areas of sparse vegetation. Adults spend the winter in deep burrows dug
into the side of sand dunes. The beetles are sometimes slow to appear in spring because of the
slow warming of deeper sand.

Localities. — (Fig. 41) Empress, Empress (11 km south). Fort Macleod, Gem, Medicine
Hat, Sandy Point, South Saskatchewan River (junction with Red Deer River).

9. Cicindela purpurea LeConte (Figs. 27 & 43)

Cicindela purpurea purpurea X C. p. auduboni LeConte

Recognition. — (Fig. 27) Beetles of this species come in two color forms, black or green. In
Alberta most specimens lack the subhumeral spot, and this readily distinguishes them from
adults of C. splendida limbalis

Tiger beetles of Alberta

329

The name C. purpurea purpurea refers to populations of black beetles, and C. purpurea
auduboni to populations of green beetles. Populations in Alberta are similar to those across the
Great Plains, being comprised of a mixture of green and black beetles. Subspecies designation
is normally written as C. p. purpurea X auduboni to reflect the mixture of forms. The
subspecies in Alberta is C. p. purpurea X auduboni.

Habitat. — These beetles are found on patches of bare clay soil interspersed with clumps of
grass and other plants. This habitat occurs frequently in prairie grasslands. Cicindela purpurea
winters as an adult. Members of this spring-fall species require at least two years for larval
development.

Localities. — (Fig. 43) Aden, Bassano, Brooks, Burdett, Calgary, Castor, Cessford, Cypress
Hills, Dillberry, Edmonton, Empress (11 km south), Etzikom, Fort Macleod, Ghost River,
Gleichen, Hanna, Hussar, Jenner, Lethbridge, Lost River (5 km north, junction with Milk
River), Magrath, Manyberries, Medicine Hat, Merid, Milk River, Orion, Pincher Creek, Ross
Creek, Taber, Tilly, Walling.

10. Cicindela splendida Hentz (Figs. 29 & 44)

Cicindela splendida limbalis Klug

Recognition. — (Fig. 29) Elytral maculations of these beetles are similar to those of adult C.
purpurea. The reddish tinge and presence of a subhumeral spot on the elytra serve as
distinguishing features. Wallis (1961) considered limbalis as a separate species, without
distinct forms. Johnson (in prep.) includes limbalis as a subspecies of Cicindela splendida.
Beetles of the limbalis phenotype are characterized by a coppery to brown head and prothorax
with moderately wide elytral maculations. The denverensis phenotype is characterized by
blue-green colors and more variable elytral maculations. In Alberta, beetles with coppery
greenish to coppery brown colors occur. Johnson considers these beetles to be a blend of
limbalis and denverensis phenotypes. The majority of specimens in Alberta can be assigned to
the limbalis phenotype, hence the subspecies designation, C. splendida limbalis.

Habitat. — These beetles prefer steep clay banks for breeding purposes but adults may be
found almost anywhere. I have collected them in the reedy margin of a slough in Calgary. The
usual habitat is on bare clay banks of streams. These beetles have a two year life cycle with
overwintering adults.

Localities. — (Fig. 44) Bilby, Brocket, Calgary, Calling River Ranger Station, Devon,
Edmonton, Elk Island National Park, Fairview, Fawcett, Fort MacKay, Fort McMurray, Fox
Creek, George Lake, Gleichen, Golden Spike, Grande Prairie, Happy Valley (Porcupine Hills),
Heatherdown, Lausand, Leduc, Nestow, Nevis, Pembina River, Pincher, Pincher Creek, Pouce
Coupe (B.C., east in Alberta), Prairie Bluff Mountain, Red Deer, Redwater, Stauffer, Smoky
Lake, Stettler, Sundance, Trochu, Wabamum, Wapiti River (south of Grande Prairie),
Wetaskwin.

11. Cicindela decemnotata Klug (Figs. 28 & 45)

Recognition. — (Fig. 28) This green or violaceous tiger beetle is characterized by reduction
of the humeral lunule and a long, descending arm of the middle band of the elytron. There are
no recognized subspecies.

Notes. — The violet form of this species is common in the Peace River district and a
population of this beetle occurs in grasslands surrounding Whitehorse, Yukon Territory. Adults
of this species should be watched for in grassland areas along northern rivers.

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Habitat. — These beetles occur on clay, sandy, or gravel soils often along cowpaths and
roads. Adults are frequently captured on clay alluvium in badlands and are known to
overwinter.

Localities. — (Fig. 45) Deadwood, Dorothy, Drumheller, Empress (11 km south), Fairview
(16 km southeast), Fort Macleod, Green Island (sic!, = Verte Island), Happy Valley (Porcupine
Hills), Lethbridge, Lost River (5 km north, junction with Milk River), Manyberries (8 km
south), Manyberries (32 km south), Majestic, Medicine Hat, Milk River (junction with Lost
River), Onefour, Peace River, Pincher, Taber.

12. Cicindela fulgida Say (Figs. 7, 20 & 46)

Cicindela fulgida fulgida Say

Recognition. — (Figs. 7 & 20) These beetles differ from all other Albertan cicindelids in
their heavy maculation pattern and greasy appearance. Adults of the subspecies C. fulgida
fulgida are 12 mm in length and have a brilliant coppery to coppery green lustre. The humeral
lunule is widely separated from the middle band at its tip.

Habitat. — Members of this species occur on alkaline soils along streams, badlands, coulees
and some sloughs on the prairies. Adults are found in areas devoid of vegetation. Beetles can be
collected early in spring and late in fall, implying a wintering adult.

Localities. — (Fig. 46) Chappice Lake, Grassy Lake (2.5 km south), Jenner Ferry (now
Jenner Bridge), Onefour, Lost River, Medicine Hat, Sandy Point.

13. Cicindela scutellaris Say (Figs. 5, 24 & 47)

Cicindela scutellaris scutellaris Say

Recognition. — (Figs. 5 & 24) Adults of this species cannot be mistaken for those of any
other species of Albertan tiger beetle. Bright red elytra lacking all maculations characterize
this species in Alberta. Subspecies recognition is also based on absence of maculations and on
bright red color; other subspecies have reduced maculations along the elytral margins.

Habitat. — Beetles of Cicindela scutellaris occur on blowouts and sand dunes in southern
Alberta and inhabit the sparsely vegetated edge zone dominated by Scurf Pea ( Psorealea
lanceolata Pursh). The sand dune habitat is shared in part with representatives of Cicindela
formosa, C. lengi and C. limbata nympha. Adults overwinter.

Localities. — (Fig. 47) Chappice Lake, Drumheller, Empress, Empress (11 km south),
Medicine Hat.

14. Cicindela lengi Horn (Figs. 23 & 48)

Cicindela lengi versuta Casey

Recognition.— (Fig. 23) Adults of this species appear similar to those of Cicindela formosa
but are distinguished by their smaller size, and long, straight humeral lunule. Most Albertan
specimens have a reddish elytral ground color. A few beetles examined were green and one was
bright metallic blue. Since the majority of specimens have reddish elytra and coppery thoracic
sclerites the subspecfic name C. /. versuta applies. The name C. /. lengi refers to blue to
blue-green populations. Other color variations also occur; a deep purple specimen was taken at
Opal and Wallis(1961) reported a black specimen from Saskatchewan.

Habitat. — Members of this species inhabit dry open sandy areas in the grasslands of
Alberta and may be found on prairie sand dunes and boreal forest sand ridges. These spring-fall
beetles may take up to three years to complete larval development. Adults winter in burrows

Tiger beetles of Alberta

331

dug in sandy soil.

Localities. — (Fig. 48) Barber Lake, Blackfalds, Chappice Lake, Claysmore, Clyde (6.5 km
east), Edgerton, Edmonton, Empress, Empress (11 km south), Fort Macleod, Milk River (16
km north of Aden), Milk River (junction of Lost River), Opal, Pakowki Lake, Rolling Hills,
Sand Hill Lake, Writing on Stone Provincial Park (32 km east).

15. Cicindela tranquebarica Herbst (Figs. 19 & 49)

Cicindela tranquebarica kirbyi LeConte

Recognition. — (Fig. 19) The long, obliquely-directed, descending arm of the humeral
lunule is a distinguishing character. Cicindela tranquebarica is common and widespread.

Geographical variation in C. tranquebarica is complex and poorly known. Many names have
been given to local varieties. The dominant phenotype found in Alberta is that of C. t. kirbyi
LeConte. The markings are broad with a bronzy-green ground color. North of Alberta, adults
of C. t. borealis Harrington, can be recognized by a broken humeral lunule band or by the ends
of the band narrowly joined in the middle. I have not seen material from Alberta representing
the borealis phenotype. However specimens from north of Wandering River had reduced band
widths although further north, at Fort MacKay, elytral band widths reverted back to the wide
state. Specimens of Cicindela tranquebarica borealis should be watched for in northern
Alberta.

Habitat. — Representatives of this species occur in almost any tiger beetle habitat, ranging
from alkaline mud flats, sandy blowouts, and prairie grasslands to boreal forest trails.
Disturbed areas are readily colonized. Areas with reduced vegetation cover are preferred. These
beetles overwinter as adults.

Localities. — (Fig. 49) Aden (16 km west), Barber Lake, Barnwell, Barons, Bilby, Brazeau
River (near Lodgepole), Calgary, Calling Lake Ranger Station, Castor, Chappice Lake, Chin,
Claresholm, Clyde (6.5 km east), Clyde (10 km north), Clymont, Consort, Crimson Lake,
Deadwood, Drayton Valley, Drumheller, Dunvegan, Edmonton, Empress (11 km south),
Fairview (16 km southeast), Fawcett, Fort MacKay, Fort Macleod, Fort McMurray (22.4 km
north), Garth, Golden Spike, Gorge Creek, Grande Prairie, Gull Lake, High River, Jenner,
Jenner Ferry (Jenner Bridge), Kootenay Plains, Lac La Biche, Lake Cardinal, Lethbridge,
Lethbridge (8 km south), Lesser Slave Lake, Lundbreck, Medicine Hat, Milk River (junction
of Lost River), Nanton, Nestow, New Dayton (1.6 km east), North Saskatchewan River (near
Nordegg), Opal, Peace River, Pincher, Police Lake, Ranfurly, Red Deer, Rosedale, Sand Hill
Lake, Smoky River, Snaring River (Jasper National Park), Saint Mary’s Reservoir, Simpson,
Soda Lake, Stavely, Stauffer, Tofield, Vilna, Wandering River (64 km north), Wetaskiwin,
Winterburn, Writing on Stone Provincial Park (32 km east).

16. Cicindela punctulata Oliver (Figs. 31 & 50)

Cicindela punctulata punctulata Oliver

Recognition. — (Fig. 31) Adults of Cicindela punctulata are readily distinguished by a row
of metallic blue or green dots running down the length of each elytron. The other elytral
maculations are quite variable, ranging from immaculate to well marked. Usually the
maculations consist of a few white spots. Cicindela puntulata punctulata is the only known
subspecies occurring in Canada.

Habitat. — Cicindela punctulata occurs in the southern prairie regions. Thin grass with bare
patches of sandy loam is preferred. Adults survive a single summer with the larvae being the

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only wintering stage. Members of this species are reported to have a one year life cycle
(Shelford 1908), however Hamilton (1925) speculated that it took two years to reach maturity.

Localities. — (Fig. 50) Bassano (junction of Highways 1 & 550), Burdett, Comrey, Empress
(11 km south), Grassy Lake, Happy Valley (Porcupine Hills), Jenner, Lethbridge, Medicine
Hat.

17. Cicindela terricola Say (Figs. 25, 26, 51 & 52)

Cicindela terricola cinctipennis LeConte (Figs. 25 & 51)

Cicindela terricola imperfecta LeConte (Figs. 26 & 52)

Recognition. — (Figs. 25 & 26) Adults of Cicindela terricola cinctipennis are small slender
beetles with an unbroken marginal band. The middle band of the elytra may be distinct or
reduced.

Adults of Cicindela terricola imperfecta are slightly larger. The marginal band is greatly
reduced or at most the humeral lunule is represented by a small spur which joins up with the
middle band.

Notes. — Some believe that these subspecies should be elevated to full species status. This
recognition is supported by their distinct appearance, habitat preference and geographic
distribution.

Cicindela terricola imperfecta ranges through British Columbia into western Alberta, on
the Kootenay Plains near Nordegg. C. oregona (also primarily found in B.C.) is found here as
well. To the west of the plains are low mountain passes into British Columbia. It appears that
individuals of this subspecies dispersed eastward over the mountains along river valleys into
Alberta, colonizing the grasslands of Kootenay Plains. To the east, extensive montane forest
appears to have prevented further dispersal onto the prairies. Specimens of Cicindela terricola
imperfecta should be watched for in other mountain grasslands along the foothills. It would be
very informative if mixed populations of C. t. imperfecta and C. t. cinctipennis could be found.
This would help in determining the species or subspecies status of the two forms.

Adults of Cicindela terricola cinctipennis have been taken in the grasslands around
Whitehorse, Yukon Territory. This species should be watched for in all native grassland areas
in Alberta.

Habitat. — Adults of Cicindela terricola cinctipennis prefer sparse grass on clay soils in the
prairie grasslands. Wallis (1961) reported that these beetles may also be found on saline and
alkaline soils with sparse vegetation. Adults of C. t. imperfecta have been taken on sandy clay
soils on river banks, the type of habitat on which the beetles were captured at Kootenay Plains
(Ball, pers. com. 1975). Adults of both subspecies are active during midsummer.

Localities. — Cicindela terricola cinctipennis : (Fig. 51) Calgary, Dorothy, Drumhelier,
Dunvegan, Empress, Fairview (16 km southeast), Fort Macleod (junction Highways 1 & 2),
Green Island (sic! = Verte Island), Hussar, Jenner, Lethbridge, Manyberries, Medicine Hat,
Milk River (junction of Lost River), Munson, Patricia (near), Redcliff.

Cicindela terricola imperfecta-. (Fig. 52) Kootenay Plains.

18. Cicindela nevadica Leconte (Figs. 30 & 53)

Cicindela nevadica knausi Leng

Recognition. — (Fig. 30) These tiger beetles resemble those of C. cuprescens, a nonresident
species. Adults of Cicindela nevadica knausi are bronze with off-white elytral maculations. The
humeral lunule is slightly recurved toward the base and body hairs are decumbent. Only one

Tiger beetles of Alberta

333

subspecies is known from Canada, C. n. knausi.

Habitat. — These beetles may be found along margins of streams and lakes on alkaline soil.
Adults prefer open areas with sparse vegetation and are active during midsummer.

Localities. — (Fig. 53) Jenner Ferry (Jenner Bridge), Lost River (near junction with Milk
River), Sandy Point.

19. Cicindela lepida Dejean (Figs. 32 & 54)

Recognition. — (Fig. 32) Adults of this species are the palest tiger beetles in Alberta and
have obscure markings and pale legs offering them excellent cryptic protection on pale sand. It
is often easier to see the beetle’s shadow on the ground than it is to see the beetle.

Notes. — Adults of Cicindela lepida may become inactive when ground surface
temperatures exceed 48° C during midafternoon. They burrow beneath the lethal temperature
zone and resume activity when the temperature drops later in the day. On warm evenings
activity will continue through the night, with short pauses at dusk and dawn. When night time
temperatures exceed 25° C (rarely in Alberta), adults may disperse. In Nebraska I have
collected flying adults in a black light trap many kilometers from the nearest sand dune.

Habitat. — Representatives of Cicindela lepida occur on pale yellow to white sand, usually
on sand dunes. No vegetation or other protective cover is present. Adults prefer wind-swept
dune crests and larvae are found in sheltered bowl areas on sand dunes. Members of this
summer species take two years to complete their life cycle (Criddle 1910, Hamilton 1925,
Shelford 1908). Adults are present from late June to early August in Alberta.

Localities. — (Fig. 54) Empress (1 1 km south).

Tiger beetles of the species Cicindela lepida occur in the Great Sandhills of Saskatchewan,
just east of the Alberta boundary. Specimens should be watched for in the sand dunes and
blowouts of the Middle Sand Hills of Alberta.

20. Other species.

In Vaurie’s paper (1950, p 153) Cicindela togata La Ferte is reported to occur in Alberta.
This report is a typographical error and should read C. tranquebarica as per species references
later on the page. No additional records or specimens have been located. Wallis (1961) did not
include this species as part of the Canadian tiger beetle fauna. The known range of Cicindela
togata does not extend north of Nebraska (Willis 1967). This species is not expected to be
found in Alberta.

Members of Cicindela togata inhabit alkali mud flats a type of habitat abundant in the
south eastern corner of the province. Adults are pale with reduced pigment areas of the elytra.
For details on appearance and habitat see Willis (1967).

Another species not yet recorded but to be watched for is Cicindela cuprescens LeConte.
This species occurs in Manitoba and along the lower reaches of the Milk River in Montana.
Adults are characterized by a peculiarly shaped humeral lunule, which resembles the
maculations found on beetles of C. nevadica knausi. The strongly embossed maculations
distinguish specimens of C. cuprescens from those of C. n. knausi. In Manitoba, adults of C.
cuprescens have been collected on the sandy crest of a ravine. Willis (1967) characterized these
beetles as inhabitants of fluvial mesic and saline habitats. This species may occur along the
Milk River drainage in southern Alberta.

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BIOGEOGRAPHIC CONSIDERATIONS

The Albertan tiger beetle fauna is of relatively recent origin. The displacement of biota by
the Wisconsian glaciation is well documented (e.g. Wright and Frey 1965, Flint 1971,
Matthews, 1979, etc). Movement of biota back into glaciated regions is not as well understood.

Faunal and floral elements survived in well-identified refugia during this glaciation but
which elements of the biota survived in which refugia? Workers (e.g. Ross 1970, Martin 1958,
Frenzel 1973, etc.) have variously interpreted where these elements went and which factors
influenced their distribution.

A poor fossil record for beetles (Morgan et al. 1983) necessitates the use of distribution
patterns to interpret faunal source regions for Albertan tiger beetles. Modern
species/subspecies distribution maps (Freitag 1965, Wallis 1961, Willis 1967) provide data on
possible source regions. Source areas for Albertan tiger beetles can, potentially, be any of the
following; 1) nunataks: refugia surrounded by glacial ice usually harbouring endemic forms; 2)
north of the ice: unglaciated areas of Alaska and the Yukon; and 3) south of the ice:
unglaciated continental North America. Region 3 can be subdivided into three major
subregions, a) western: the Pacific Northwest, west of the Rocky Mountains, b) central: the
Great Plains region, and c) eastern, including the boreal forest.

Tiger beetles are sensitive to low temperature and short growing season as shown by their
absence from extreme northern, alpine and subalpine regions. It is therefore unlikely that any
species survived on nunataks within glacial areas or in regions close to ice margins during
glacial maxima.

A northern source area for Albertan tiger beetles is suggested by present distribution
patterns (Fig. 56). One species, C. oregona , has several subspecies. Freitag (1965) showed that
C. oregona guttifera ranges from Alaska and the Yukon through northern British Columbia.
From central British Columbia and south, C. oregona guttifera hybridizes with C. oregona
oregona in a zone which extends south along the Rocky Mountains into Utah. From Colorado
and south into New Mexico C. oregona guttifera hybridizes with C. oregona navajoensis Van
Dyke and with C. oregona maricopa Leng in southwest Utah. “Pure” populations of C. oregona
guttifera occur in Colorado and New Mexico in the south and in northern British Columbia,
Yukon and Alaska. These widely separated population loci and extensive zones of hybridization
suggest that the ancestors of C. oregona guttifera were isolated in the unglaciated regions of
Alaska and the Yukon in the north and in the mountainous areas of New Mexico and Colorado
east of the Great Divide during the last glacial period. Populations of these beetles dispersed
north and south following deglaciation whereby contact was renewed with other
subspecies/sibling species. The hybrid zones reported by Freitag (1965) are these areas of
contact. Thus it appears that at least one tiger beetle species may have dispersed into Alberta
from a northern refugium.

The remaining tiger beetle species probably originated from source areas south of the
Wisconsinan ice margin. One subspecies, C. limbata hyperborea may be tentatively attributed
to a southeastern origin. These tiger beetles are restricted to the boreal forest in northern
Alberta and Saskatchewan (Fig. 56). A problem with attributing an eastern origin to the
subspecies is that there are no extant populations known from the east. Rates of subspeciation
have been proposed for montane carabids (Kavanaugh 1979) but it is not known how quickly
tiger beetles can subspeciate. In most tiger beetle species there is considerable individual
variation. It is possible that such rates are rapid and that C. /. hyperborea evolved to subspecies

Tiger beetles of Alberta

335

status while isolated on the jack pine sand plains of northern Alberta and Saskatchewan in the
past 7000 years. Another explanation is that ancestral populations survived on ’’boreal” sand
hills south of glacial ice and are now absent from these areas. Additional research is required to
solve this problem.

A southwestern source region for Cicindela terricola imperfecta and C. oregona
oregona(F\%. 56) is readily supported by populations found along mountain passes and valleys
of western Alberta. Populations of these beetles appear to be in the process of dispersing and
colonizing Alberta. Both subspecies occur in British Columbia and the U.S.A., west of the
Great Divide.

The remaining Albertan tiger beetles probably originated on the central Great Plains.
Populations of these species in Alberta are simply northern extensions of these ranges (Fig. 55).
Habitat and climate appear to limit dispersal. Ranges of summer species do not extend north of
the prairie grasslands, whereas some spring-fall species have ranges extending north into the
Northwest Territories along streams and river banks (C. splendida limbalis, C. tranquebarica,
C. duodecimguttata, C. repanda ). A number of ’southern grassland’ tiger beetle species (C.
decemnotata, C. lengi versuta , and C. terricola cinctipennis ) occur in the prairie regions of the
Peace River district and two species (C. decemnotata , and C. t. cinctipennis ) in the grasslands
of the Yukon. This distribution parallels that of many plant species (Moss 1952).

Following deglaciation the fauna moved around, adjusting to changes in climate. About
7000 years B.P. a prolonged warm period, the hypsithermal occurred. During this time, prairie
grasslands probably expanded north in Alberta at the expense of the forested regions. The
grasslands of the Peace River district and the southern prairies were continuous, with a
resulting exchange of floral and faunal elements. Since the hypsithermal, the climate has
cooled and the forests have reclaimed much of these grasslands. This has resulted in the
reduction and isolation of remnants of northern grasslands with their relict prairie flora and
fauna.

Other tiger beetle species (C. lepida, C. formosa) may have dispersed north into Alberta
during the hypsithermal when dune habitats were in abundance. Riparian species (C. repanda ,
C. duodecimguttata , C. hirticollis ) followed the changing water sheds, losing habitat in times
of drought and flood. Species of alkaline mud flats ( C.fulgida , C. nevadica ) would lose habitat
during pluvial periods and gain it back during periods of drought. The tiger beetle fauna is thus
in a constant state of flux. Some species are still colonizing the province, some are represented
by relict populations, and others are adapting and flourishing in the wake of man’s activities:
colonizing and dispersing along roadways, and breeding in construction sites. Agriculture has
destroyed some habitats and created others.

ACKNOWLEDGEMENTS

I wish to thank my many friends who assisted in the development of this paper: B.F. and
J.L. Carr, C. van Nidek, J.H. Acorn, E.M. Pike, and F.A.H. Sperling for providing collection
data and companionship in the field; C.D. Bird the mentor of my first term paper on Alberta
tiger beetles; and G.E. Ball for opening his home and the U. of Alberta collection to me in 1975
to begin research on the tiger beetles of Alberta. Special thanks are extended to J.H. Acorn and
F.A.H. Sperling for critically reviewing and suggesting improvements to this manuscript.

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REFERENCES

Criddle, N. 1907. Habits of some Manitoba tiger beetles (Cicindelidae). The Canadian
Entomologist. 39: 105-114.

Criddle, N. 1910. Habits of some Manitoba tiger beetles, No. 2 (Cicindelidae). The Canadian
Entomologist. 42: 9-15.

Flint, B. 1971. Glacial and Quaternary geology. J. Wiley and Sons Inc. 892 pp.

Freitag, R. 1965. A revision of the North American species in the Cicindela maritima group
with a study of hybridization between Cicindela duodecimguttata and oregona. Quaestiones
Entomologicae. 1: 87-170.

Frenzel, B. 1973. Climatic fluctuations of the Ice Ages. The Press of Western Reserve
University, Cleveland and London. 306 pp.

Graves, R.C. 1962. Predation on Cicindela by a dragonfly. The Canadian Entomologist. 94:
1231.

Hatch, M.H. 1953. The beetles of the Pacfic Northwest. Part 1. Introduction and Adephaga.
University of Washington Press, Seattle, Washington. 340 pp.

Hamilton, C.C. 1925. Studies on the morphology, taxonomy and ecology of the larvae of
Holarctic tiger-beetles (family Cicindelidae). No. 2530 Proceedings U. S. National
Museum. 65. Art. 17, 1-87, pis. 1-12.

Johnson, W. (in prep). Notes on Cicindela splendida Hentz - limbalis Klug. - devenerensis
Casey and ludoviciana Leng. Cicindela.

Kavanaugh, D.H. 1979. Rates of taxonomically significant differentiation in relation to
geographic isolation and habitat: examples from a study of the nearctic Nebria fauna, p
35-57. In Carabid beetles: their evolution, natural history, and classification. Eds. Erwin,
Ball, Whitehead, and Halpern. Dr. W. Junk bv Publishers. The Hague 1979, 635 pp.

Lavingne, R.J. 1972. Cicindelids as prey of robberflies (Diptera: Asilidae). Cicindela 4(1): 1-7.

Martin, P.S. 1958. Pleistocene ecology and biogeography of North America, pp. 375-420. In
Hubbs, C.L. (Ed.). 1958. Zoogeography. American Association for Advancement of
Science, Symposium. Washington, D.C.

Matthews, J.V., jr. 1979. Tertiary and Quaternary environments: historical background for an
analysis of the Canadian insect fauna, pp. 31-86. In Danks, H.V. (Ed.). 1979. Canada and
its insect fauna. Memoirs of the Entomological Society of Canada. No. 108, 573 pp.

Morgan, A.V., A.C. Achworth and J.V. Matthews jr. 1983. Late Wisconsin fossil beetles in
North America. In Porter, S.S. (Ed.). 1983. Late Quaternary environments of the United
Sates, Vol. 1. University of Minnesota Press, Mineapolis. 407 pp.

Moss, E.H. 1952. Grassland of the Peace River region, western Canada. Canadian Journal of
Botany. 30:98-124.

Ross, H.E. 1970. The ecological history of the Great Plains, evidence from grassland insects,
pp. 225-240. In Dort, W. and J.K. Jones (Ed.). 1970. Pleistocene and recent environments
of the Central Great Plains. Department of Geology, University of Kansas Special
Publication 3, University of Kansas Press. 433 pp.

Shelford, V.E. 1908. Life histories and larval habits of the tiger beetles (Cicindelidae). The
Journal of the Linnaean Society, London. 30: 157-184, pis. 23-26.

Swan, L.A. 1975. Tiger Beetle. Insect World Digest. Jan/Feb. 1975: 1-5.

Vaurie, P. 1950. Notes on the habitats of some North American tiger beetles. Journal of the
New York Entomological Society. 58: 143-153.

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Wallis, J.B. 1961. The Cicindelidae of Canada. University of Toronto Press, Ontario. 74 pp.
Willis, H.L. 1967. Bionomics and zoogeography of the tiger beetles of saline habitats in the
central United States (Coleoptera: Cicindelidae). University of Kansas Science Bulletin.
47(5): 1 43—3 1 3.

Willis, H.L. 1968. Artificial key to the species of Cicindela of North America north of Mexico
(Coleoptera: Cicindelidae). Journal of the Kansas Entomological Society. 41: 303-317.
Wright, H.E. and D.G. Frey (Ed.). 1965. The Quaternary of the United States. Princeton
University Press, Princeton, N.J. 922 pp.

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Figures 1-8. Drawings of larva and character states on adults. Scale line = 5 mm. Fig. 1 . Habitus of generalized Cicindela
larva. Fig. 2 Larval head and prothorax of C. oregona oregona, Osoyoos, B.C. Fig. 3. Generalized maculation pattern with
nomenclature used. Fig. 4. Hairy head; C. limbata nympha. Fig. 5. Head with culstered hairs on inner margin of each eye;
C. scutellaris. Fig. 6. Glaborous head; C. nebraskana. Fig. 7. Glaborous genae; C. fulgida. Fig. 8. Hairy genae; C.
duodecimguttata.

Tiger beetles of Alberta

339

Figures 9-16. Adult tiger beetles, line drawings, color refers to normal range, locality refers to where the specimen was
collected. Scale line = 10 mm. Fig. 9. C. nebraskana\ black; Empress, Alta. Fig. 10. C. longilabris ; black to green; Fedora,
Alta. Fig. 11. C. hirticollis ; brown to olive; Gull Lake, Alta. Fig. 12. C. limbata hyperborea\ brown; Fort MacKay, Alta.
Fig. 13. C. limbata nympha\ brown to greenish; Crimson Lake, Alta. Fig. 14. C. repanda ; brown; Edmonton. Alta. Fig. 15.
C. oregona oregona-, brown, green, blue; Summerland, B.C. Fig. 16. C. oregona guttifera\ brown; Kootenay Plains, Alta.

Quaest. Ent., 1985,21 (3)

Figures 17-24. Adult tiger beetles, line drawings, color refers to normal range, locality refers to where the specimen was
collected. Scale line = 10 mm. Fig. 17. C. oregona X C. duodecimguttatcr, brown; Carbondale River, Alta. Fig. 18. C.
duodecimguttata brown; Stauffer, Alta. Fig. 19. C. tranquebarica\ grey to brown; Calgary, Alta. Fig. 20. C. fulgida\
coppery to metallic green; Lost River (near Onefour), Alta. Fig. 21. C. formosa formosa, red violet; Empress (11 km
south), Alta. Fig. 22. C. formosa gibsonr, red to violet; Maple Creek (16 km north), Sask. Fig. 23. C. lengi versuta; rust,
rarely green, blue, or black; Chappice Lake, Alta. Fig. 24. C. scutellaris scutellaris\ red-green; Empress ( 1 1 km south),
Alta.

Tiger beetles of Alberta

341

Figures 25-32. Adult tiger beetles, line drawings, color refers to normal range, locality refers to where the specimen was
collected. Scale line = 10 mm. Fig. 25. C. terricola cinctipennis\ green to olive; Calgary, Alta. Fig. 26. C. terricola
imperfecta ; green to olive; Kootenay Plains, Alta. Fig. 27. C. purpurea ; green or black; Lost River (near Onefour), Alta.
Fig. 28. C. decemnotata\ green; Lethbridge, Alta. Fig. 29. C. splendida limbalis\ red to green; Crimson Lake, Alta. Fig.
30. C. nevadica knausi\ copper to brown; Jenner Ferry, Alta. Fig. 31. C. punctulata ; grey brown; Empress (1 1 km south),
Alta. Fig. 32. C. lepida\ pale with brown; Empress (1 1 km south), Alta.

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Figures 33-36. Distribution maps. Fig. 33. C. repanda. Fig. 34. C. duodecimguttata. Fig. 35. C. oregona. Fig. 36. C.
hirticollis.

Tiger beetles of Alberta

343

Figures 37-40. Distribution maps. Fig. 37. C. limbata nympha. Fig. 38. C. limbata hyperborea. Fig. 39. C. longilabris.
Fig. 40. C. nebraskana.

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Figures 41-44. Distribution maps. Fig. 41. C.formosa formosa. Fig. 42. C.formosa gibsoni. Fig. 43. C. purpurea. Fig. 44.
C. splendida limbalis.

Tiger beetles of Alberta

345

Figures 45-48. Distribution maps. Fig. 45. C. decemnotata. Fig. 46. C.fulgida. Fig. 47. C. scutellaris sculellaris. Fig. 48.
C. lengi versuta.

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Figures 49-52. Distribution maps. Fig. 49. C. tranquebarica. Fig. 50. C. punctulata. Fig. 51. C. terricola cinctipennis. Fig.
52. C. terricola imperfecta.

Tiger beetles of Alberta

347

C. long ilabris

C. longi labris
C. hi rt icoll is
C. rapanda
C. t ra nquabarica

C scut •Maria
C. formosa
C. fulgida

Figures 53-56. Distribution maps. Fig. 53. C. nevadica knausi. Fig. 54. C. lepida. Fig. 55. Composite distribution map of
tiger beetle species believed to have dispersed into Alberta from the southern Great Plains. Fig. 56. Composite distribution
map of tiger beetle species believed to have dispersed into Alberta from the north, west and east.

Quaest. Ent., 1985, 21 (3)

CHARACTERISTICS AND EVOLUTION OF ELYTRAL SCULPTURE IN THE TRIBE
GALERITINI (COLEOPTERA: CARABIDAE)'

George E. Ball
Department of Entomology
University of Alberta

Edmonton, Alberta Quaestiones Entomologicae

Canada T6G 2E3 21: 349-367 1985

ABSTRACT

A reconstructed phytogeny, based primarily on structural features other than details of the
elytral cuticle, provides the basis for inferring evolution of macrosculpture and
microsculpture of the elytra of galeritine adults. Macrosculpture consists of a system of
alternating linear depressions ( interneurs ) and elevations (intervals). A transformation series
extends in the subtribe Galeritina from primary intervals that are broad and slightly convex
(or flat) to costate to carinate, with or without development of secondary intervals. In the
monobasic more plesiotypic subtribe Planetina, the elytra have developed carinate primary
and secondary intervals independently of these features in the more highly evolved groups of
subtribe Galeritina. Within some taxa of Galeritina, the secondary intervals (carinulae) have
been reduced or lost. The microsculpture system of microlines and included sculpticells has
undergone a complex series of changes in the Planetina and Galeritina. Independently, in both
subtribes, the plesiotypic microlines have been lost, and sculpticells are represented by
nodules, which are only parts of the original sculpticells. In the genus Eunostus Castelnau
(subtribe Galeritina), the plesiotypic microlines are evident, and sculpticells are transverse and
flat, but a few exhibit small nodules. Convergence is postulated between Planetina and
Galeritina with independent development of the same type of macrosculpture and
microsculpture, and also within the Galeritina, with independent reduction in different
lineages of the system of carinae and carinulae. The patterns of macrosculpture and
microsculpture are correlated to the extent that adults with carinate intervals exhibit elongate
sculpticells with transversely aligned nodules. This relationship may be the result of: 1)
selective forces acting similarly on different genes to produce a functional complex; or 2) there
may be a developmental constraint, such that ontogenetic development of carinae somehow
channels or influences development of the derived form of microsculpture. If alternative 1 is
correct, the derived, correlated forms of macrosculpture and microsculpture may be accepted
as discrete character states for evaluation of phylogenetic relationships; if alternative 2 is
correct, the derived pattern of macrosculpture and microsculpture must be regarded as a
single character state. The biological significance of these transformation series is unknown,
though the transverse form of sculpticells is generally correlated in other carabids with life in
tightly packed leaf litter. The system of longitudinal carinae and sculpticells is reminiscent of
a corrugated iron roof, and may be especially effective for shedding water and debris. Because
this latter form of sculpture is exhibited by related species that have strikingly different

'Based on the text of an address presented to the XVII International Congress of Entomology,
Hamburg, Federal Republic of Germany, August, 1984

350

Ball

ecological requirements, the additional inference is made that sculpture is not responding to
specific environmental factors, but rather to factors that are more general.

RESUME

Une reconstruction de la phylogenie des Galeritines, etablie principalement a partir de caracteres structuraux autres
que les details de la cuticule elytrale, nous sert de base pour deduire les etapes evolutives de la macrosculpture et de la
microsculpture des elytres chez les Galeritines adultes. La macrosculpture consiste en un ensemble de sillons
(interneures) alternant avec des elevations lineaires (intervalles). Dans la sous-tribu des Galeritina, il existe une serie de
transformations des intervalles primaires qui passent de larges et legerement convexes (ou aplatis ) d costes ou carenes,
avec ou sans apparition d’intervalles secondaires. Dans la sous-tribu monogenerique des Planetina, qui constitue un
groupe plus plesioty pique, les elytres ont developpe des carenes primaires et des intervalles secondaires independamment
de ceux qu’on retrouve dans les groupes plus evolues de la sous-tribu des Galeritina. Chez certains taxons des Galeritina.
il y a reduction ou perte des intervalles secondaires (carenules). La microsculpture, comprenant un ensemble de
microlignes et de «sculpticellules», a subi une serie complexe de changements chez les Planetina et les Galeritina.
Independamment dans les deux sous-tribus, les microlignes plesiotypiques ont disparu et les «sculpticellules» n’existent
plus qu'd I’etat de nodules qui correspondent d une partie des «sculpticellules» originelles. Chez le genre Eunostus
Castelnau (de la sous-tribu des Galeritina), les microlignes plesiotypiques sont evidentes et les «sculpticellules» sont
transverses et aplaties, mais certaines «sculpticellules» montrent de petits nodules. Nous postulons qu’il y a eu
convergence, d’une part entre les Planetina et les Galeritina lors du developpement d’un type semblable de
macrosculpture et de microsculpture, et d’autre part parmi les Galeritina oil il y a eu reduction du systeme de carenes et
de carenules de facon independante dans les differentes lignees. Les motifs de macrosculpture sont correles avec ceux de
microsculpture dans la mesure ou les adultes ayant des intervalles carenes possedent des «sculpticellules» allongees avec
des nodules alignes transversalement. Cette relation peut etre le resultat soit de forces selectives agissant similairement
sur des genes differents pour produire un ensemble fonctionnel, soit de contraintes de developpement qui font que, d’une
certaine faqon, I’ontogenese des carenes canalise ou influence le developpement du type derive de microsculpture. Si la
premiere alternative est correcte, nous pouvons accepter les types derives et correles de macrosculpture et de
microsculpture comme etant des etats de caracteres distincts pour revaluation des relations phylogenetiques; par contre,
si la seconde alternative est correcte, le motif derive de macrosculpture et de microsculpture doit etre considere comme
un seul etat de caractere. La signification biologique de ces series de transformations est inconnue, bien que la presence
de «sculpticellules» transverses est generalement correlee, chez d’autres Carabiques, avec un mode de vie dans la litiere
compacte. Le systeme de carenes et de «sculpticellules» longitudinales fait penser d un toit de tole ondulee et peut etre
particulierement efficace pour se debarasser de I’eau et des debris. Etant donne que ce dernier type de sculpture se
rencontre chez des especes apparentees qui possedent des exigences ecologiques fort differentes, nous deduisons par
surcroit que la sculpture n’est pas assujettie d des facteurs environmentaux specifiques, mais plutot d des facteurs plus
globaux.

INTRODUCTION

Most of what is known about galeritine carabids is summarized in various comparatively
recent taxonomic treatments: Basilewsky (1963), Afrotropical species, Reichardt (1965 and
1967), the Asian species of Galerita, and the species of Galeritini in the New World,
respectively; Lindroth (1969: 1091), and Ball and Nimmo (1983), species of the predominantly
Nearctic subgenus Progaleritina.

The tribe Galeritini is pan-tropical, with northern extensions into the Nearctic (northward
to southern Ontario and Quebec) and eastern Palaearctic (northward to the Japanese
Archipelago and southern Korea) Regions. Habitats occupied range from waterside stations
and the rain forest floor in the tropics to dry open forests and savannas. Most species live at low
altitudes, but in the American tropics, a number of species are known from montane forest.

Adults and larvae of all species are probably predators on other arthropods, though this has
been shown for only a few species. It seems reasonable to make the extrapolation, because of
general similarity in body form and details of the mouthparts among all taxa. Females of
Galerita ( Progaleritina ) bicolor Drury lay their eggs in mud balls which are then attached to
the undersides of leaves. This behavior is correlated with a peculiarly modified ovipositor.

Elytral Sculpture in the Tribe Galeritini

351

which is characteristic of the more highly evolved galeritines. By extrapolation, it seems likely
that all such taxa have similar habits, and that those with more plesiotypic ovipositors have
more plesiotypic habits, and probably lay eggs in cavities in the soil, as do most female
carabids. Adults of many of the macropterous species are found at lights, at night, showing that
they are nocturnal and that they fly. Little else is known about ecological aspects of galeritines.

Although knowledge of galeritines is markedly restricted, I was able to make a
reconstructed phylogeny, using previously studied features of adults, and adding analyses of
structure of the mandibles and ovipositor (Ball, in press). Macrosculpture of the elytra was
used to reconstruct the phylogeny of Galerita (sensu lato ), but microsculpture was not studied
in detail. Subsequently, I realized that elytral sculpture exhibited some interesting complexity,
so I asked if patterns of sculptural variation might be correlated with the reconstructed
phylogeny that I had made. Results are presented below.

MATERIAL AND METHODS

Material

The adults studied were those on hand that had been collected by me, were in the Strickland
Museum of my Department, or were borrowed from other institutions for the phylogenetic
study of the Galeritini. In aggregate, they represented a reasonably diverse cross-section of the
tribe, but not all species. Sculpture of the elytra was examined superficially using
representatives of the following taxa: Planetes bimaculatus MacLeay, P. ruficollis Nietner, P.
pendleburyi Andrewes, and Planetes species?; Eunostus herrarensis Alluaud, E. vuilloti
Alluaud, Eunostus new species; Ancystroglossus ovalipennis Reichardt, A. dimidiaticornis
Chaudoir, and Ancystroglossus new species; Trichognathus marginipennis Latreille; and all
seven species of Galerita, subgenus Progaleritina. From subgenus Galerita, I examined
specimens of G. perrieri Fairmaire, G. sulcipennis Reichardt, various members of eight New
World sub-groups: americana, carbonaria, costulata, gracilis, jelskii, occidentalis, striata, and
unicolor, and four species of the G. africana group.

Detailed examination of microsculpture was made for specimens of Planetes bimaculatus,
Eunostus herrarensis, Ancystroglossus ovalipennis, Trichognathus marginipennis, Galerita
mexicana Chaudoir, G. sulcipennis, G. perrieri, G. ruficollis Dejean, G. boucardi Chaudoir, G.
balli Reichardt, G. attelaboides Fabricius, and G. procera Gerstaecker.

Methods

Preparation and study of specimens.-Elytra of specimens chosen for superficial study were
cleaned initially with ammonia applied with a moistened bit of tissue paper held in forceps.
These specimens were examined with a Wild M5 Stereo-binocular microscope, at 50X
magnification. On the basis of such examination, major types of sculpture were identified and
specimens representing each type were selected for detailed examination.

For such study, except for the specimen of G. perrieri, the left elytron was removed, cleaned
in water using a sonicator, attached to a standard mount, and coated with gold using a sputter
coater. Specimens were examined and photographed, using a Cambridge S-250 “Stereoscan”
Scanning Electron Microscope. The specimen of the rare Madagascan G. perrieri, was
examined with its elytra attached to the body, uncoated, at relatively low magnifications of the
SEM.

Analytical procedures. — These concerned identification of ancestral features of sculpture
for each of the branching points of a tree that represented the reconstructed phylogeny of the

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suprageneric groups of Galeritini. The general method used was hypothesis of transformation
series (Figs. 2 and 5) polarized with the premises that flat (or slightly convex) elytral intervals
are plesiotypic features of macrosculpture, and an isodiametric pattern with flat, non-imbricate
sculpticells is the plesiotypic condition of the microsculpture. The latter premise is based on
conclusions reached by Hinton (1970: 41-42), and Lindroth (1974).

The sequence of stages proposed in evolution of macrosculpture and microsculpture follows
from the reconstructed phylogeny of Galeritini (Fig. 1, from Ball [in press]). For each pair of
sister groups, the sculpture pattern most like the proposed ancestral pattern was judged the
more plesiotypic, and accepted as the ancestral pattern for that pair of sister taxa.
Transformation series for macrosculpture and microsculpture were established separately. The
separate analyses are presented together on diagrams representing the reconstructed phylogeny
of Galeritini (Figs. 7 and 8).

SCULPTURE OF THE ELYTRA

For purposes of this presentation, the term “macrosculpture” refers to the alternating
system of longitudinal convexities (intervals) and concavities (interneurs) on the surface of a
typical elytron. Intervals mark the areas which are the courses of veins of the fore wing
(Jeannel, 1941: 30-31). “Microsculpture” refers to the network of fine lines and microscopic
sculpticells (Allen and Ball, 1980: 486) that cover the surface. This network, in its most
plesiotypic form, reflects the form of the cellular network of the underlying epidermis (Hinton,
1970: 41-42). Types of macrosculpture are designated by Roman numerals and capital letters;
microsculpture types are designated by Arabic numerals and capital letters.

Macrosculpture

Within the tribe Galeritini, intervals range in form from broad and flat (Fig. 2, Type I) to
broad and convex (costate, Fig. 2, Type III), to narrow and convex (carinate. Fig. 2, types
II-IV). An elytron exhibits a simple arrangement, with all intervals being equal in width and
convexity, or a complex arrangement, with a pair of secondary intervals (carinulae)
intercalated between adjacent broader, primary intervals (carinae, Fig. 2, Subtype IV A, and
Fig. 3). The number of carinae is either nine (Fig. 2, Subtype Ha, and Type IV), or five
(Subtype IIB).

Microsculpture

At magnifications of about 50X, the cuticle of most arthropods exhibits a mesh of fine lines,
like the lines of a fish net (Lindroth, 1974: 252, and Allen and Ball, 1980: 485-486). Meshes
are characterized as isodiametric, transverse, or longitudinal, depending upon their relative
lengths and widths. “Sculpticells” (Allen and Ball, 1980: 486) between microlines range in
form from flat to slightly or markedly convex, to carinate (Ball, 1975: Fig. 114).

Galeritines exhibit a variety of forms of microsculpture. At the base of an elytron,
sculpticells are flat, slightly imbricate (Harris, 1979: 19 and 30, Fig. 40). and nearly
isodiametric (Fig. 6), or transverse (Fig. 4). Most of the elytral surface is:

a. covered with a network of transverse meshes (some sculpticells with posterior nodules, Fig.
5, Type 1); or

b. with nodule-like swellings, either not arranged in a pattern (Subtypes 2A and B), or aligned
transversely (Types 3 and 4).

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353

RECONSTRUCTED PHYLOGENY OF TRIBE GALERITINI

PLANETINA

Planetes

(Progaleritma) Galerita (s sjricto)

perrieri americana

Complex Complex

GALERITINA

Eunostus

Ancystrogl.

Trichogn.

Galerita (s

lato)

perrieri africana sulcip americ.

Group Group Group Group

Fig. 1. Reconstructed phylogeny of Tribe Galeritini. Taxa are: Subtribe Planetina - Planetes MacLeay; Subtribe
Galeritina - Eunostus Castelnau; Ancystroglossus Chaudoir; Trichognathus Latreille; Galerita (sensu lato) - subgenus
Progaleritina Jeannel, and subgenus Galerita Fabricius, including the G. perrieri complex (with G. perrieri and G.
africana groups), and the G. americana complex (with G. sulcipennis and G. americana groups).

The general term for sculpture of Types 2, 3, and 4 is nodulate (Harris, 1979: 15). As detailed
below, each nodule is hypothesized as representing only part of an original sculpticell.

PHYLOGENETIC RELATIONSHIPS OF THE GALERITINI

A reconstructed phylogeny of the Galeritini (Fig. 1), based on features of adults (Ball, in
press), provides a framework for an evolutionary analysis of sculpture patterns. Each node is
designated by a capital letter (A-H), in alphabetical sequence, depending upon recency of
common ancestry, except for the terminal two nodes. Aspects of elytral sculpture were used as a
major feature to reconstruct the phylogeny of the supraspecific taxa of the genus Galerita , but

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SUBTRIBE GALERITINA: TRANSFORMATION SERIES
IN MACROSCULPTURE OF ELYTRA

G. (G.) africana Group

Type I
Eunostus
Ancystroglossus
Trichognathus
<3 ( Progalentina)

G (G.) perrieri Group

Fig. 2. Subtribe Galeritina: transformation series in macrosculpture of elytra. Types IIA-IVB represent approximately the
basal one third, and Type I, the basal one quarter, of the left elytron. Species represented by illustrations are the following:
Type I - Eunostus herrarensis Alluaud, Ancystroglossus ovalipennis Reichardt, Trichognathus marginipennis Latreille,
G. (Progaleritina) mexicana Chaudoir, and G. (Galerita) perrieri Fairmaire; Subtype IIA - G. (Galerita) attelaboides
Fabricius; Subtype IIB - G. ( Galerita ) procera Gerstaecker; Type III - G. (Galerita) sulcipennis Reichardt; Subtype IVA -
G. (Galerita) ruficollis Dejean; and Subtype IVB - G. (Galerita) balli Reichardt. Scale bars represent 1.0 mm.

Elytral Sculpture in the Tribe Galeritini

355

SUBTRIBE PLANETINA : Planetes
MACROSCULPTURE - LEFT ELYTRON

BASAL PORTION DISC

Fig. 3. Subtribe Planetina: macrosculpture of the left elytron of Planetes bimaculatus MacLeay. Scale bars represent 250
nm.

not to reconstruct the phylogeny of the other genera.

Overall, the system reflects important changes in structure of the mouthparts and ovipositor.
There is also a striking increase in body size associated with node D, probably reflecting a
change in habits from that of hunting concealed in the leaf litter to running on the surface of
the forest floor, or in more open areas.

PATTERNS OF ELYTRAL SCULPTURE OF THE GALERITINI

Although the subtribe Planetina exhibits more plesiotypic features than does the Galeritina,
outgroup comparison shows that planetine adults have highly derived sculpture. Thus, the
subtribe Galeritina, with its greater range of sculpture types, is the focal group for elucidation

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of evolution of these systems, and the following analyses begin with consideration of this
subtribe.

Macrosculpture

Subtribe Galeritina. — Figure 2 illustrates the hypothesized trends in elytral
macrosculpture. The figures also illustrate the reduced basal ridge that is characteristic of the
adults of Galeritina. Four general types of macrosculpture are recognized, based on structural
and phylogenetic considerations.

Type I includes elytra with broad intervals that are either flat or slightly convex. I did not
formally distinguish between the slight difference involved. Type I is characteristic of three
genera, and of one subgenus and one species group of Galerita (sensu lato ).

Type II macrosculpture is characteristic of the G. africana group ( Galeritiola Jeannel, of
previous authors). It differs from Type I by having wider and deeper interneurs, and
consequently narrower intervals, the latter being carinate. Two subtypes of macrosculpture are
recognized: IIA, with nine carinae; IIB, with five carinae.

Type III macrosculpture is characteristic of the monobasic Middle American montane G.
sulcipennis group. The elytral intervals are more elevated than in Type I, and are classified as
costate. The figure, unfortunately, does not do justice to the difference between the two types of
sculpture.

Type IV macrosculpture is characterized by carinae and readily seen carinulae (Subtype
IV A), or if carinulae are not readily apparent, careful examination reveals vestiges of them
(Subtype IVB). Subtype IVB looks very much like IIA, but the carinae of IVB are not as high,
and the interneurs of IIA lack any indication of carinulae.

Subtribe Planetina. — Figure 3 illustrates macrosculpture for a specimen of Planetes. The
pattern is Subtype IVA. Carinulae appear to be nearly as wide as the carinae, but in fact there
is a substantial difference as the figure of a portion of the elytral disc, taken at higher
magnification, indicates. At working magnifications (ca. 5X - 50X), however, the carinae and
carinulae appear about equal, so that the elytra seem to have a densely packed system of
carinae, and thus seem quite different from the Subtype IVA elytra of Galerita.

Microsculpture

Subtribe Galeritina. — Figure 4 illustrates two general types of microsculpture
characteristic of galeritines: imbricate, which is confined to the basal area, principally basad of
the basal ridge; and nodulate, which is more or less extensive on the disc. The sculpticells of the
imbricate type are flat and broad, while the nodulate sculpticells are narrower and convex. Four
types of microsculpture are recognized on the elytral disc in the Galeritina, and their proposed
evolutionary trends are illustrated in Figure 5. Type I, which is characteristic of Eunostus the
sister group of the other three galeritine genera, exhibits markedly transverse, flat sculpticells
across most of the surface. Laterally, however, some sculpticells have small medio-apical
nodules.

Types 2-4 are characterized by widespread nodulate microsculpture, without microlines.
Type 2 exhibits a non-patterned arrangement of nodules, with Subtype 2 A having fewer
nodules than Subtype 2B. The former is characteristic of Ancystroglossus, the latter of
Trichognathus and subgenus Progaleritina.

In Type 3 microsculpture, which is confined to the G. perrieri species complex, the nodules
are in transverse rows: in 3A, exhibited by adults of the G. perrieri group, the nodules are short

Elytral Sculpture in the Tribe Galeritini

357

MICROSCULPTURE AT BASE
OF LEFT ELYTRON

Trichognathus marginipennis

Fig. 4. Macrosculpture at base of left elytron of Trichognathus marginipennis Latreille. Scale bar represents 1 50 fim.

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SUBTRIBE GALERITINA : TRANSFORMATION SERIES
IN MICROSCULPTURE OF ELYTRA

Fig. 5. Subtribe Galeritina: transformation series in microsculpture of elytra. Type I illustrates a portion of interneur 7 in
the basal third of the left elytron. Subtypes 2A-4B illustrate portions of interneur 3 and/or 4, in the basal third of the left
elytron. Species represented are: Type I - Eunostus herrarensis Alluaud; 2A, Ancystroglossus ovalipennis Reichardt; 2B,
Trichognathus marginipennis Latreille, and G. (Progaleritina) mexicana Chaudoir; 3A, G. ( Galerita ) perrieri Fairmaire;
3B, G. (Galerita) attelaboides Fabricius; 3C, G. ( Galerita ) procera Gerstaecker; 4A, G. ( Galerita ) sulcipennis Reichardt;
4B, G. (Galerita) ruficollis Dejean. Scale bars represent 50 nm.

Elytral Sculpture in the Tribe Galeritini

359

SUBTRIBE PLANETINA : Planetes
MICROSCULPTURE - LEFT ELYTRON

BASE DISC

Fig. 6. Subtribe Planetina: microsculpture of the left elytron of Planetes bimaculatus MacLeay. The illustrations represent
parts of the left elytron: the basal tenth, toward the sutural margin; and a portion of interneur 3 and adjacent carinulae.
Scale bars represent 10 /mi.

and uniform across the elytral surface; for 3B and 3C, characteristic of the G. africana group,
the nodules are longer than those of 3 A, but inter se are relatively shorter (3B) or longer (3C),
flattened basally, and in fairly well marked transverse rows, between carinae. On the tops of the
carinae, the sculpticells are elongate and flat, and closely adpressed.

Type 4 microsculpture is exhibited by adults of the G. americana complex. Of the two
Subtypes, 4A {G. sulcipennis group) is most like that of the G. perrieri group. The difference is
seen in the elongate and flattened nodules on the top of the elytral costae. In Subtype 4B
(exhibited by adults of the G. americana group), the nodules are longer and the transverse rows
between adjacent carinae and carinulae are better defined. In those adults exhibiting Subtype
IVB macrosculpture (i.e., with carinae reduced), locations of atrophied carinulae are indicated

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by the markedly elongate sculpticells that are a characteristic feature of the tops of carinae and
carinulae.

Subtribe Planetina. — Figure 6 illustrates microsculpture characteristic of Planetes adults.
The elytral base has imbricate, flat, and essentially isodiametric sculpticells. The disc exhibits
long, keeled nodules arranged in transverse rows between adjacent carinae and carinulae. The
sculpticells of the latter are very narrow and linear.

EVOLUTION OF ELYTRAL SCULPTURE OF THE GALERITINI
The Pattern

Figures 7 and 8 illustrate and summarize the hypothesis of evolution of sculpture. Figure 8
is a continuation of Fig. 7. For the labelled nodes except G, the hypothesized ancestral
combination of sculptural features is illustrated, based on features of extant galeritines,
macrosculpture above, microsculpture below. The ancestral states for node G are the same as
for F. For each of the extant groups whose sculptural features differ from those of the ancestral
stock, illustrations are also provided.

Features of the common ancestor. --These are inferred from the most plesiotypic sculptural
features of extant adult galeritines. They are Type I macrosculpture, and
imbricate-isodiametric microsculpture, the latter as seen on the elytral base of Planetes adults.

Macrosculpture. — The reconstructed phylogeny suggests that from Ancestor A to F or G in
subtribe Galeritina, there were no significant changes in macrosculpture. From Ancestor G,
with Type I macrosculpture, Type II developed, and further differentiated into two subtypes, in
the G. africana group, with Subtype IIB losing four carinae. From Ancestor F, Type III
sculpture emerged in Ancestor H, and from the latter, Type IV, which in turn differentiated
into two subtypes, in the G. americana group.

To determine polarity of Type IV sculpture, I relied on correlation of characters, for this
part of the transformation series is not ordained by the reconstructed phylogeny presented in
Figure 1. Subtype IVB is associated with the derived features of brachyptery and life in
montane environments, in the northern part of the Neotropical Region. Subtype IVA, on the
other hand, is associated with the ancestral features of macroptery and life in lowland
environments, over extensive areas of the tropics. Reichardt (1967: 158) postulated, and I
agree, that the traces of carinulae are evidence of loss, associated with reduction of wings and
loss of flight, rather than that the traces represent the precursors of fully developed carinulae.

Although there is no sign in Type III of developing carinulae, or widened interneurs to
foreshadow development of Type IV sculpture, a costate condition (Type III) could be a
reasonable step between nearly flat (Type I) and carinate (Type IV) conditions.

In the lineage that gave rise to the Planetina, macrosculpture Type IVA also arose.
Although intermediate extant forms are unknown, it seems unlikely that the change from the
postulated ancestral condition occurred without intermediate changes like those proposed for
the Galeritina.

Microsculpture. — Although changes in macrosculpture came relatively late in the
Galeritina lineage, the pattern for microsculpture suggests an early striking change, followed
by less marked differentiation. I suggest that imbricate isodiametric sculpture of Ancestor A
changed in Ancestor B to transverse sculpture, with some sculpticells exhibiting nodules. This
was followed on the surface apicad of the basal ridge, by spread of the nodules over the disc,
and disappearance of the plesiotypic lines that marked the sculpticells. The number of nodules
increased, and took on an arrangement in rather irregular transverse rows (Ancestor F,

Elytral Sculpture in the Tribe Galeritini

361

TRIBE GALERITINI : RECONSTRUCTED PHYLOGENY
OF GENERA AND EVOLUTION OF SCULPTURE
OF ELYTRA

Planetes

Eunostus

Ancystroglossus

Trichognathus

Galerita

ANCESTOR

Fig. 7. Tribe Galeritini: reconstructed phylogeny of genera and evolution of sculpture of elytra. Sculpture of elytra of
Planetes, Eunostus , and Ancestors A-D are each represented by a pair of Figures, of which the lower is microsculpture, and
the upper macrosculpture. The figure for Ancestor C also represents Ancystroglossus , and those for Ancestor D also
represent Trichognathus and Galerita. The illustrations are of the left elytron, basal portion, as explained in captions for
Fig. 2 (macrosculpture) and Fig. 5 (microsculpture). Specimens represented are: Ancestor A - microsculpture, Planetes
bimaculatus MacLeay, and macrosculpture, Ancystroglossus ovalipennis Reichardt; Planetes bimaculatus\ Ancestor B -
microsculpture, Eunostus herrarensis Alluaud, and microsculpture, A. ovalipennis ; Eunostus herrarensis\ Ancestor C, A.
ovalipennis ; Ancestor D, Trichognathus marginipennis Latreille. Scale bars represent at low magnification, 500 at
high magnification, 50 fim.

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GENUS Galenta : RECONSTRUCTED PHYLOGENY
OF SUBGENERA, SPECIES COMPLEXES AND
GROUPS, AND EVOLUTION OF SCULPTURE
OF ELYTRA

Subgenus

Progaleritma perrieri

Group

africana

Group

sulcipennis

Group

americana

Group

ANCESTOR

Go I e r 1 1 a

americana Complex

( s_ s t r i c t o )

ANCESTOR

Fig. 6. Genus Galerita-. reconstructed phylogeny of subgenera, species complexes and groups, and evolution of sculpture of
the elytra. Sculpture of the elytra of Ancestor E, Ancestor F, the G. africana group, Ancestor H, and the G. americana
group are each represented by a pair of Figures, of which the lower is microsculpture and the upper macrosculpture. The
Figures for Ancestor E also represent subgenus Progaleritina-, for Ancestor F, also Ancestor G and the G. perrieri group;
for Ancestor H, also the G. sulcipennis group. The figures are of the left elytron, basal portion, as explained in the caption
for Fig. 2 (macrosculpture) and Fig. 5 (microsculpture). Specimens represented are of these species: Ancestor E - G.
(Progaleritina) mexicana Chaudoir; Ancestor F - G. ( Galerita ) perrieri Fairmaire; G. africana group - G. (Galerita)
attelaboides Fabricius; Ancestor H - G. ( Galerita ) sulcipennis Reichardt; and G. americana group - G. (Galerita)
ruficollis Dejean. Scale bars represent at low magnification, 500 nm; at high magnification, 50 pm, and at very high
magniFication, 5 p.m.

Elytral Sculpture in the Tribe Galeritini

363

Subtype 3A). As a more complex macrosculpture evolved, the transverse rows of nodules were
confined to the interneurs (Subtypes 3B, 3C, 4 A, and 4B).

Subtype 3A microsculpture seems easily derived from Subtype 2B by development of a more
orderly arrangement of nodules. Subtypes 3B and 3C are derived from 3A by a still more
ordered arrangement of nodules, and possibly by fusion of pairs of nodules, in adjacent rows, to
yield nodules that are fewer and longer. The transverse rows of nodules, confined to the
interneurs (Subtypes 3B, 3C, 4A, and 4B), probably decreased in number by fusion of members
of adjacent rows, and, consequently, the individual nodules became longer (Subtypes 3C and
4B). On the elytral base, transverse imbricate sculpture was retained. Transformation of 3 A to
4A and the latter to 4B is virtually self-evident, parallelling the transformation of 3A to 3B,
and to 3C.

It is important to note that the transverse sculpticells on the elytral disc of Eunostus adults
are not imbricate. This change is interpreted as a loss, and a reversion to a state more
plesiotypic than is exhibited by the sculpture of Ancestor A.

The planetine lineage adults evolved, on the elytral surface apicad of the basal ridge,
nodulate microsculpture with long nodules, similar to that of Subtype 4B. As for the
macrosculpture, intermediate steps are not known for evolution of the microsculpture, between
the hypothetical ancestral condition and that of the extant species of Planetes. On the basal
area of the elytra, the imbricate isodiametric sculpture was retained.

Microsculpture of the elytral base that is characteristic of Planetes seems the most
plesiotypic pattern among extant Galeritini. The discal sculpture, on the other hand, is highly
derived, with no known extant antecedants.

Convergence among taxa. — The same derived patterns of macrosculpture and
microsculpture are represented in planetines and galeritines, and within distantly related
members of the Galeritina. Adults of Planetes and of the G. americana group exhibit the
complex type of elytral macrosculpture, with development of a system of alternating carinae
and pairs of carinulae. Similarly, within the genus Galerita, a system of carinate intervals has
evolved independently in different groups of the subgenus Galerita. Also, Planetes , and the
Galerita americana and africana groups, have evolved independently a pattern of long,
transversely aligned nodular microsculpture, and elongate sculpticells on the tops of the
carinae.

Loss of carinae or carinulae has occurred independently in the G. africana group (carinae
lost), and in the G. americana group (carinulae lost). Although these losses involve different
structures, the end result in each lineage is similar.

Parallel development of macrosculpture and microsculpture. — As noted above, carinate
macrosculpture has had correlated with it development of long narrow nodules, transversely
arranged between intervals.

Significance of the Pattern

In order to highlight general implications of this study, brief comments are offered about
historical, developmental, and functional significance of the evolutionary pattern of sculpture of
the Galeritini. From an historical perspective, I suggest that the highly complex surface of the
arthropod cuticle exhibits patterns of variation that are amenable to phylogenetic analysis. This
study suggests that features of the cuticle are sufficiently stable that old patterns persist. For
example, if the estimate of age of Galerita is correct (Ball, in press, based on vicariant
distribution patterns of extant taxa), the subgenera of this genus pre-date the beginning of the

Quaest. Ent., 1985,21 (3)

364

Ball

Tertiary Period, and the other genera likely originated still earlier. Therefore, the
microsculpture patterns ought to be of a similar range of ages. Thus, seemingly minor and
inconsequential details of surface structure have potentially as much value as have other, more
obvious structural features for phylogenetic analysis and taxonomic use.

The examples of convergence are interesting, for they show that even rather complex and
detailed similarities may arise independently. Thus, it is important to evaluate critically such
similarities if one uses cuticular features in phylogenetic analysis.

Similarities between the patterns of evolution of macrosculpture and microsculpture
exhibited among the species whose adults have carinate intervals may result from a functional
relationship (see below) of genetically independent elements, or they may be the result of one
system constraining the other. If these derived forms of sculpture were genetically independent
of one another, then they would constitute separate character states for use in analysis of
phylogenetic relationship. Perhaps, however, development of carinae somehow channels or
influences development of the transverse pattern of and elongation of the nodules. If this is so,
and if one wanted to use the characters as sources of evidence about evolutionary relationship,
it would be necessary to determine the extent of the develomental relationship. If
microsculpture type were totally dependent upon type of macrosculpture, then one would have
but a single character, rather than two, with which to evaluate propinquity of relationship.

Previous authors (Hinton and Gibbs, 1969: 962; Lindroth, 1974: 261-263; Erwin, 1979: 547;
Allen and Ball: 544; Goulet, 1983: 375; and Ball and Shpeley, 1983: 800) have considered the
ecological role played by surface features of carabids, proposing that irregularities and dullness
contribute to cryptic patterns, while brilliance caused by marked reflectivity or iridescence
yields flash patterns which are confusing to potential predators. Alternatively (Erwin, 1979:
547), it has been proposed that since different patterns of sculpture are associated with
different types of habitats, the patterns might function to protect an insect’s body against
unfavorable environmental influences. For example, a grated pattern (i.e., diffraction grating)
might be especially effective in shedding mud and water, and thus of value to insects living in
wet, sticky environments. Using the analogy of a corrugated iron roof, the correlation of elytral
carinae with longitudinally directed nodules looks like a run-off system for shedding unwanted
material that comes in contact with the cuticle. Perhaps this system has therefore a similar
function to that of a grated system of microsculpture, the different solutions being the result of
selection for different types of environmental impediments.

More specifically, adults of Eunostus exhibit the transverse pattern of microsculpture. In
other carabid taxa, this pattern is correlated with life in tightly-packed leaf litter, but I do not
know if this is the type of habitat frequented by Eunostus. The nodular forms of microsculpture
are characteristic of all other galeritine groups, whose range of habitats collectively extends
from closed canopy rain forest to open woodland and riparian situations. So, the functional
significance of the different types of microsculpture is not likely to be found by seeking
correlates with different habitats. Correlation might be found at the level of microhabitats,
when these have been determined for galeritines.

In spite of my inability to demonstrate its adaptive significance, since this evolutionary
pattern has developed and has been maintained for an extended period of time, and since the
features are exposed to environmental pressures including potential predators that rely on
eyesight while hunting, it seems reasonable to infer that natural selection has influenced and is
maintaining this structural system. Futhermore, in view of the rather small steps in at least
portions of the transformation series, it seems reasonable to infer sustained directional

Elytral Sculpture in the Tribe Galeritini

365

selection, perhaps associated with either changes in habitat, or with improved design for
occupying the old habitats. (Ball, in press).

ANOTHER INTERPRETATION OF EVOLUTION OF ELYTRAL SCULPTURE IN

THE GALERITINI

Reichardt (1967: 158) considered evolution of macrosculpture of the elytra of subgenus
Galerita. Assuming that Subtype IVA sculpture was plesiotypic for this group, which he
ranked as a genus, and to which he related Planetes, he proposed that Types I, II, and III and
Subtype IVB were derived from the former Subtype: for II and IVB, by simple loss of
carinulae; and for Types I and II, both by loss of the carinulae and reversion from carinate to
costate or nearly flat intervals. In turn, this notion was based on two considerations: evident
reduction of the carinulae in adults of highland species in Middle America, and association of
this loss with brachyptery, an apotypic condition. However, he did not take account of the fact
that associated with macrosculpture Types I and III is a plesiotypic form of microsculpture, nor
that in the G. africana group (with Type II macrosculpture) there is no evidence that carinulae
had ever been present.

Having taken account of these facts, and as well having shown elsewhere (Ball, in press)
that the subgenus Galerita and Planetes are not closely related to one another, and
consequently there is no need on the basis of out-group comparison to postulate that Subtype
IVA macrosculpture is plesiotypic, I believe that Reichardt’s hypothesis of the evolution of
elytral macrosculpture in the Galeritini can be rejected.

CONCLUDING STATEMENT

In this paper, I have recognized and described the types of sculpture exhibited by
representative galeritines, using both structural and phylogenetic considerations to do so. I have
demonstrated a marked correlation between microsculpture pattern and the reconstructed
phylogeny that I had made previously. Underlying the reconstructed phylogeny based on
structural features, there ought to be a correlated series of ecological transformations. When
the latter are found and analyzed, I believe we will have the basis for understanding in both
functional and historical terms the patterns of evolution of elytral sculpture postulated here.

ACKNOWLEDGEMENTS

I offer thanks for the loan of especially important material for this study: to P. Basilewsky
(Musee Royal de l’Afrique Centrale, Tervuren, Belgique), for making available a specimen of
Galerita perrieri Fairmaire; and to D. H. Kavanaugh (California Academy of Science, San
Francisco, California), for the loan of specimens of Planetes and Eunostus.

Technical assistance was provided by various members of the staff of my Department. D.
Shpeley and G. D. Braybrook collaborated in undertaking the work with the SEM. J. S. Scott
did the layout and prepared the plates, which are vital components of this presentation. I. E.
Bergum assisted with preparation of the final copy of the manuscript.

My associates R. S. Anderson and J. R. Spence reviewed a preliminary draft of the
manuscript, and offered useful suggestions for improvement of presentation. Though I accepted
most of their proposals and made extensive revisions, I declined to accept all of them. I am,
nonetheless, grateful for their thoughtful, thorough reviews of both form and substance.

Quaest. Ent., 1985, 21 (3)

366

Ball

I must also acknowledge members of the audience at Hamburg, at the XVII International
Congress, whose comments following my oral presentation, caused me to modify some of the
statements that I made there.

The research on which this study was based was financed by Grant A- 1399, Natural
Sciences and Engineering Research Council of Canada. Funding that made possible the oral
presentation at Hamburg was provided partially by the NSERC grant, and partially by the
University of Alberta Endowment Fund for the Future.

I am very grateful for the generous and friendly cooperation that made this study a pleasure
to undertake.

REFERENCES CITED

Allen, R. T. and G. E. Ball. 1980. Synopsis of Mexican taxa of the Loxandrus series
(Coleoptera: Carabidae: Pterostichini). Transactions of the American Entomological
Society, 105: 481-576.

Ball, G. E. 1975. Pericaline Lebiini: notes on classification, a synopsis of the New World
genera, and a revision of the genus Phloeoxena Chaudoir (Coleoptera: Carabidae).
Quaestiones Entomologicae, 11: 143-242.

Ball, G. E. (in press). Reconstructed phylogeny and geographical history of the Tribe Galeritini
(Coleoptera: Carabidae). In, Taxonomy, phylogeny, and zoogeography of beetles and ants: a
volume dedicated to the memory of Philip Jackson Darlington, Jr., 1904-1983 (G. E. Ball,
Editor). W. Junk bv Publishers, Dordrecht, Boston, London.

Ball, G. E. and A. P. Nimmo. 1983. Synopsis of the species of subgenus Progaleritina
(Coleoptera: Carabidae: Galerita Fabricius). Transactions of the American Entomological
Society, 109: 295-356.

Ball, G. E. and D. Shpeley. 1983. The species of the eucheiloid Pericalina: classification and
evolutionary considerations (Coleoptera: Carabidae: Lebiini). The Canadian Entomologist,
115:743-806.

Basilewsky, P. 1963. Revision des Galeritininae d’Afrique et de Madagascar (Coleoptera:
Carabidae). Annales, Musee Royal de l’Afrique Centrale, No. 120, 93 pp.

Erwin, T. L. 1979. Thoughts on the evolutionary history of ground beetles: hypotheses
generated from comparative faunal analyses of lowland forest sites in temperate and tropical
regions (Coleoptera: Carabidae), pp. 539-592. In, Carabid beetles: their evolution, natural
history, and classification. Proceedings of the First International Symposium of
Carabidology. (T. L. Erwin, G. E. Ball, D. R. Whitehead, and A. Halpern, Editors). Dr. W.
Junk bv Publishers, The Hague, Boston, London. X+ 644 pp.

Goulet, H. 1983. The genera of Holarctic Elaphrini and species of Elaphrus Fabricius
(Coleoptera: Carabidae): classification, phylogeny, and zoogeography. Quaestiones
Entomologicae, 19: 219-482.

Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, No.
28. State of California Department of Food and Agriculture Division of Plant Industry
Laboratory Sciences. Sacramento, California, 31 pp.

Hinton, H. E. 1970. Some little known surface structures, pp. 41-58. In, Insect ultrastructure.
Symposium of the Royal Entomological Society of London, Number Five. (A. C. Neville,
Editor). Blackwell Scientific Publications, Oxford and Edinburgh.

Hinton, H. E. and D. F. Gibbs, 1969. An electron microscopic study of the diffraction gratings

Elytral Sculpture in the Tribe Galeritini

367

of some Carabid beetles. Journal of Insect Physiology, 15: 959-962.

Jeannel, R. 1941. Coleopteres carabiques, premier partie. Faune de France, 39: 1-571. Paul
LeChevalier, Paris.

Lindroth, C. H. 1969. The ground-beetles (Carabidae excl. Cicindelinae) of Canada and
Alaska. Opuscula Entomologica, Supplementum 34, pp. 945-1 192.

Lindroth, C. H. 1974. On the elytral microsculpture of carabid beetles (Col. Carabidae).
Entomologica Scandinavica, 5: 251-264.

Reichardt, H. 1965. The Asian species of Galeritula Strand. Breviora, No. 225: 1-16.
Reichardt, H. 1967. A monographic revision of the American Galeritini (Coleoptera:
Carabidae). Arquivos de Zoologia do Estado do Sao Paulo, 15: 1-76.

Quaest. Ent., 1985,21 (3)

369

BOOK REVIEW

D.C.M. Manson. 1984. Fauna of New Zealand; Number 4, Eriophyoidea except Eriophyinae
(Arachnida: Acari); Number 5, Eriophyinae (Arachnida: Acari: Eriophyoidea). Science
Information Publishing Centre, DSIR, Wellington, New Zealand. NZ $10.50 (Number 4, 142
pp.), NZ $9.00 (Number 5, 123 pp.).

These two volumes represent the first attempt at comprehensive systematic treatment of the
Eriophyoidea of New Zealand. The author includes 109 species, of which 62 are recorded for
the first time from New Zealand and 54 are new to science. The first volume (Number 4) deals
with 49 known species listed in the families Sierraphytoptidae and Diptilomiopidae, and the
subfamilies Cecidophyinae and Phyllocoptinae of the family Eriophyidae, while the second one
(Number 5) includes 60 species of the eriophyid subfamily Eriophyinae.

In Number 4 the author begins with a brief introduction, followed by a useful historical
review of the study of eriophyoid mites in New Zealand. He then discusses one of the most
significant and controversial recent problems in the nomenclature of Eriophyoidea, and wisely
opts to follow the ruling of the International Commission on Zoological Nomenclature
concerning the retention of the pre - 1971 usage of the generic names Aceria, Eriophyes, and
Phytopus.

The section on morphology is thorough and effectively introduces the reader to the terms
used in the systematic sections. Unfortunately, the author has chosen to employ some very
unusual concepts and inappropriate terms for describing certain acarine structures. For
example, he refers to “three main body divisions - the rostrum, the dorsal or cephalothoracic
shield, and the abdomen”. In this case, “rostrum” and “abdomen” are imprecise terms
apparently being used incorrectly in place of “gnathosoma” and “idiosoma”, respectively, for
the two generally-accepted, main regions of the acarine body. The dorsal shield is, in fact,
simply a sclerite on the prodorsal region of the idiosoma. Other inaccurately applied terms,
such as “claw” for solenidion and “featherclaw” for empodium, are used following the
traditional but incorrect practices of many specialists on Eriophyoidea.

The next part, on the life cycle of eriophyoid mites, is a concise account outlining the
so-called simple and complex types of life cycles in Eriophyoidea, and emphasizing the
importance of recognizing the deutogyne form in species with the latter. This is followed by a
comprehensive discussion of the different types of damage that various eriophyoid mites cause
to host plants. The author notes that members of several species of Eriophyinae apparently, are
regularly found associated with two or more distinct types of damage on hosts of the genus
Nothofagus. As he points out, this finding suggests that the exclusive use of symptomatic
damage to hosts in establishing the identity of eriophyoid mites, so prevalent in early works on
the group, and still permitted by the International Code of Zoological Nomenclature, should be
strongly discouraged.

The last 120 pages of Number 4, and all of Number 5, are devoted to systematic treatment
of the fauna. Clear, straightforward keys and diagnostic descriptions are presented for the
protogyne females of all taxa, providing an essential framework for future taxonomic work on
the New Zealand fauna. A comprehensive set of fully adequate figures is included for each
species, illustrating the diagnostic character states used in the keys and descriptions.
Inexplicably, the author has chosen to use the family name Sierraphytoptidae for mites having
3 or 4 setae on the prodorsal shield even though the name Phytoptidae, with 67 years priority, is
available.

Quaest. Ent., 1985, 21 (3)

370

Book Review

Dr. Manson has admirably brought together existing information on the systematics of the
Eriophyoidea of New Zealand, and these attractively produced volumes will be an important
addition to the libraries of all students of the group.

Ian M. Smith,
Assitant Director,
Biosystematics Research Institute,
Ottawa, Ontario

Abstracts are required, one in English and one in another language,
preferably French.

Communications regarding subscriptions and exchanges should be
addressed to the Subscription Manager, and regarding manuscripts to the
Editor.

Published quarterly by:

Department of Entomology
University of Alberta
Edmonton, Alberta, CANADA
T6G 2E3

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Return Undeliverable mail to the address above. Return Postage Guaranteed.

371.1

QUAESTIONES ENTOMOLOGICAE ISSN 0033-5037

A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.

Volume 21 Number 4 1985

FAUNAL INFLUENCES ON SOIL STRUCTURE

Proceedings of a Symposium held at the University of Alberta, Edmonton, Alberta, CANADA,

June 11-13, 1984

Oil, Toil and Soil: An Introduction to the Symposium 371.3

Analytic and Synthetic Contributions 371.5

Kevan-Soil Zoology, Then and Now-Mostly Then 371.7

Pawluk-Soil Micromorphology and Soil Fauna: Problems and Importance 473

Rusek-Soil Microstructures - Contributions on Specific Soil Organisms 497

Parkinson-Some Impacts of Fungal-Faunal Interactions in Soil 515

Edwards-Earthworms in Soil Formation, Structure and Fertility 517

Norton- Aspects of the Biology and Systematics of Soil Arachnids, particularly

Saprophagous and Mycophagous Mites 523

Hoffman-Biological and Systematic Problems Involving Soil Dwelling

Arthropods 543

Fjellberg-Recent Advances and Future Needs in the Study of Collembola Biology

and Systematics 559

Greenslade-Pterygote Insects and the Soil: Their Diversity, Their Effects on Soils

and the Problem of Species Identification 571

Dindal-Soil Animals and Soil Fabric Production 587

Mermut-Faunal Influence on Soil Microfabrics and Other Soil Properties 595

Foster-7/i situ Localization of Organic Matter in Soils 609

Altemiiller-The Importance of Soil Fauna in Regulating Soil Microstructure and

Soil Management in Forests 635

Hill-Soil Fauna and Agriculture: Past Findings and Future Priorities 637

McGill and Spence-Soil Fauna and Soil Structure: Feedback between Size and

Architecture 645

Addenda: Techniques, Equipment, Additional References, and Priorities for Future

Study 655

McKeague and Fox-Soil Micromorphology 657

Hill and Behan-Pelletier-Priorities for the Integrated Development of Soil

Micromorphology and Soil Zoology: Results of a Brainstorming Session 665

Norton-A Variation of the Merchant-Crossley Soil Microarthropod Extractor 669

371.2

Norton and Sanders-Superior Micro-needles for Manipulating and Dissecting

Soil Invertebrates 673

Behan-Pelletier, Hill, Fjellberg, Norton and Tomlin-Soil Invertebrates: Major

Reference Texts 675

Index 687

371.3

OIL, TOIL AND SOIL:

An Introduction to the Symposium

The machinery of human interaction is facilitated by several lubricants, of which coffee is
one of the most important in our society. Morning coffee break in the Department of Soil
Science at the University of Alberta collected the people who conceived this symposium. On a
cold day in November 1982, F. D. Cook, W. B. McGill, S. Pawluk, J. A. Robertson (all of the
Department of Soil Science), H. V. Danks (Biological Survey of Canada (Terrestrial
Arthropods)), and I warmed our fingers on pottery mugs and discussed common interests about
soil arthropods, around the table. A brief prepared by the Biological Survey of Canada (1982)
drew our attention to the unexplored possibilities of opening dialogue between soil zoologists
and pedologists in Canada. It was apparent that our understanding of soil arthropods in
Canada was deficient. However, it appeared that we might know more than it seemed possible,
if we could bring together people working on soil biology from quite different perspectives. We
surmised that such a colloquium could contribute to assessment of needs and identification of
priorities for soil biology in Canada. These became central objectives for organizing a soil
animal conference.

After the creative dizziness of conception comes the toil of pregnancy and the ever present
possibility of abortion. This is the pedestrian but onerous phase of organizing a conference. In
this task, F. D. Cook, S. Pawluk and I were joined by N. Juma (Department of Soil Science)
and J. A. Campbell (Alberta Environment, Research Management Division). V.
Behan-Pelletier (Biosystematics Research Institute) was an adjunct member of this committee
and contributed valuable advice and enthusiasm throughout the planning phase.

It is well known that soil animals contribute to soil function through effects on litter
breakdown and nutrient cycling. However, we felt that the effects of soil animals on soil
structure were less widely appreciated and because of our collective interests, we decided to
focus the symposium on these interactions. Our nebulous “soil animal conference” became
“Faunal Influences on Soil Structure”. G. E. Ball (Chairman, Department of Entomology) and
W. B. McGill (Chairman, Department of Soil Science) provided strong continuing support for
our efforts and the Faculty of Agriculture and Forestry contributed seed money from the
Endowment Fund for the Future.

As we developed the program, our concept of the symposium grew and we became most
interested in attracting scholars of international reputation who might catalyse and contribute
to the interaction between Canadian pedologists and soil biologists. At this point our squeaking
wheels outran our budget and so we sought another important lubricant of human interaction -
money. The response was generous and gratifying. We received financial support for the
scientific program from the Natural Sciences and Engineering Research Council of Canada,
Alberta Agriculture, Alberta Environment, The Alberta Research Council and the Canadian
Society of Soil Science. Additional financial support from within the University community
was provided by the Conference Fund Committee, Faculty of Agriculture and Forestry and the
Departments of Entomology, Forest Science and Soil Science. The City of Edmonton and the
Province of Alberta, respectively, agreed to host coffee breaks and the closing banquet with
hospitality grants. We on the Organizing Committee are most grateful for this support.

The birth of this conference in June 1984 was more party than pain. About 80 participants
representing ten countries arrived and three days of non-stop interaction followed. The
prevailing spirit was that of a class reunion despite the fact that participants came from the two

Quaest. Ent., 1985,21 (4)

371.4

rather isolated schools of soil zoology and pedology. It is a pleasure to acknowledge the
assistance of H. van Blodeau and the staff at Lister Hall who arranged a pleasant and relaxed
environment for the conference. S. Greenberg and V. Smyth of the Faculty of Extension
organized and manned the registration desk assisted by several student volunteers.

The following collection of papers that were formally delivered at “Faunal Influences on Soil
Structure” is but a pale reflection of what actually transpired. The Hon. F. D. Bradley
(Minister of the Environment, Province of Alberta) and J. Gordin Kaplan (Vice-President
(Research), University of Alberta) opened the conference by stressing the great potential
significance of soil research in the contexts of agriculture and land management. The formal
papers published or abstracted in this collection served as a starting point for the flurry of
intellectual exchange that characterized the meeting. Many participants contributed posters
summarizing their current work about animals in the soil. These and the workshop sessions
were in a sense the heart of the conference and maintained a strong pulse of enthusiastic
discussion.

It is our hope and belief that the record of this conference shall not end at the last page of
this volume. Instead, we predict that a growing dialogue between pedologists and soil zoologists
will lead to a new, more synthetic kind of soil science that includes soil biology at its core. There
is much toil ahead but it should pay off handsomely in terms of applications and by increasing
understanding of a fascinating part of the earths biota. A central message of this conference is
that these two objectives must travel hand in hand.

Finally, I wish to acknowledge the help and support of G. E. Ball who, in the capacity of
editor of Quaestiones Entomologicae encouraged us to put this issue together and bore with us
through the most difficult aspect of this project. We also thank S. M. Subbarao, publication
manager for the journal, for her patience. Publication of these proceedings was made possible
by support from the Endowment Fund for the Future of the Faculty of Agriculture and Foresty
at the University of Alberta.

John R. Spence, Chairman
The Organizing Committee
Department of Entomology
University of Alberta
Edmonton, Alberta T6G 2E3

371.5

ANALYTIC AND SYNTHETIC CONTRIBUTIONS

Quaest. Ent., 1985, 21 (4)

371.6

SOIL ZOOLOGY, THEN AND NOW - MOSTLY THEN

D. Keith McE. Kevan
Department of Entomology and

Lyman Entomological Museum and Research Laboratory

Macdonald College Campus, McGill University

21, 111 Lakeshore Road

Ste-Anne de Bellevue, Que. H9X ICO

CANADA

Quaestiones Entomologicae
21:371.7-472 1985

ABSTRACT

Knowledge of the animals that inhabit soil remained fragmentary and virtually restricted
to a few conspicuous species until the latter part of the 19th Century, despite the publication,
in 1549, of the first attempt at a thesis on the subject by Georg Bauer (Agricola). Even the
writings of far-seeing naturalists, like White in 1789, and Darwin in 1840, did not arouse
interest in the field. It was probably P.E. Muller in 1879, who first drew particular attention
to the importance of invertebrate animals generally in humus formation. Darwin s book on
earthworms, and the “formation of vegetable mould”, published in 1881, and Drummond’s
suggestions, in 1887, regarding an analgous role for termites were landmarks, but, with the
exception of a few workers, like Berlese and Diem at the turn of the century, little attention
was paid to other animals in the soil, save incidentally to other investigations. Russell’s
famous Soil Conditions and Plant Growth could say little about the soil fauna other than
earthworms. Prior to the Second World War, Bornebusch, in 1930, and Jacot, in 1936,
attempted to broaden the horizons of both zoologists and pedologists, but it was not until the
end of the war years beginning with Forsslund’s work in Sweden, published in 1945, that soil
fauna studies really got under way. From the pedological, rather than the zoological point of
view, a book by Kubi'ena, published in 1948, set the stage. Then, in addition to research
publications, several books on different aspects of soil fauna in general appeared from 1949 to
1951 by: Gilyarov, Franz, Kuhnelt, and Delamare de Boutteville. The first international
colloquium on soil fauna was held in 1955, since when there have been many, the latest before
the present one in 1982. There has nevertheless ( with a few notable exceptions) been a general
lack of interest in the fauna on the part of pedologists, and reluctance to intrude into the
realms of so-called “soil science” by soil zoologists, to mutual disadvantage. There is still an
almost complete absence of appreciation, especially among those who determine the directions
of soil research, that we are still without the means of proper identification of innumerable
members of the soil fauna, and that the understanding of basic soil ecology and the
pedological importance of the fauna is impossible without this.

RESUME

L’auteur passe en revue le developpement des connaissance sur les animaux qui habitent dans le sol. depuis les
debuts jusqu'h maintenant. Ces connaissances demeurerent fragmentaires el pratiquemenl restreintes d quelques esptces
frappantes jusque dans la deuxieme moilie du XIXiime sitcle, et ce malgre la parution. en 1549. d'un premier essai de

371.8

Kevan

th&se sur le sujet par Georg Bauer (Agricola). Meme les Merits de naturalistes clairvoyants, tels que White (1789) et
Danvin (1840), n’e mule rent que peu d’interet dans ce domaine. P.E. Muller (1879) fut probablement le premier it porter
une attention particuliere au role important des invertebres dans la formation de ihumus. L’ouvrage de Darwin (1881)
sur les vers de terre et «la formation des moisissures vegetales» et les suggestions de Drummond (1887) concernant un
role analogue chez les termites constitu&rent des evenements marquants, mais, h I’exception de quelques chercheurs tels
que Berlese et Diem it la fin du si&cle, la plupart porterent peu d’attention aux autres animaux vivant dans le sol, sauf
accessoirement durant le cours de d’autres travaux. Le fameux ouvrage de Russell paru en 1912 et intitule Soil
Conditions and Plant Growth contient peu d’ informations sur la faune des sols autre que les vers de terre. Avant la
Deuxieme Guerre Mondiale, Bornebusch (1930) et Jacot (1936) essayerent d’elargir les horizons des zoologistes et des
pedologues, mais ce ne fut qu'd la fin de la guerre que I’etude de la faune des sols prit vraiment son essor avec les travaux
de Forsslund en Suede en 1945. Du point de vue pedologique plutot que zoologique, l’ ouvrage de Kubi'ena (1948) etablit
le domaine. Par la suite, en plus d’articles scientifiques, plusieurs ouvrages traitant de differents aspects de la faune des
sols en general parurent en succession rapide: Ghilarov (1949), Franz (1950), Kuhnelt (1950) et Delamare de Boutteville
(1951). Le premier colloque international sur la faune des sols eut lieu en 1955 (Kevan, 1955) et fut suivi par plusieurs
autres, dont le dernier precedent celui-ci eut lieu en 1982 (Lebrun et al., 1983). Neammoins, on remarque en general un
manque d’interet dans la faune des sols chez les pedologues (mis h part quelques exceptions notables), de meme qu'une
hesitation de la part des zoologistes etudiant la faune des sols d s’ingerer dans le domaine des soi-disant «sciences des
sols»; cette attitude constitue un desavantage mutuel. II existe un manque quasi total d’ appreciation, particulierement
chez ceux qui decident de I’orientation de la recherche sur les sols, du fait que nous ne disposons toujours pas d’outils
adequats pour identifier les innombrables membres de la faune des sols, et que notre comprehension des elements de base
de I’ecologie des sols et de I’importance pedologique de la faune ne pourra s’ameliorer sans cela.

Table of Contents

Introduction 371.8

Ancient World 372

The Early and Middle Mediaeval Periods 382

The Later Mediaeval Period 387

The Renaissance 390

Mid- 17th to Mid- 18th Centuries 402

Up to the Middle of the 1 9th Century 415

1850 to 1900 421

1900 to 1945 425

The Post-War Period to the 1960’s 431

Recent Times 435

Conclusion 436

Notes 438

References 442

Index 466

INTRODUCTION

Prehistoric man was well aware of other creatures that shared his environment, and he
undoubtedly associated some of these, such as various “worms”, ants and termites with the
earth beneath his feet. Like his present-day counterparts among the Bushmen of southern
Africa and the Aboriginals of Australia, too, he probably obtained an appreciable part of his
food by digging for insect grubs. Nevertheless, the nearest thing, of which I am aware, to direct
evidence for this acquaintance with such humble creatures is what seems to be a presumed
amulet in the form of a possible Necrophorus burying-beetle of the Magdalenian culture of
southern Germany, some 25,000-30,000 years ago (Peters & Topfer, 1932; Schimitschek,
1977)(Fig. la). Another representation of a subterranean insect (though of a cave-, not a
soil-inhabiting one) is also from the Magdalenian culture, but from southern France and

Soil zoology

371.9

C cl- tkarsiu.5

Fig. 1. Artifacts from the Magdalenian culture of Europe, (a) The oldest known representation of an insect, probably a
Necrophorus burying-beetle; amulet made of Tertiary carbon from Hegau, Baden, Germany, 25,000 to 30,000 years old;
after Peters and Toepfer (1932). (b) Troglophilus camel-cricket scratched on bison bone, Caverne des Trois Freres,
Ariege, France, some 20,000 years old; after Begouen and Begouen (1928). Fig. 2. Ancient Egyptian stylized scarab seals
(right member of each pair, various dates), compared with actual insect sketched on ovals (left member of each pair).
After Petrie (1917).

Quaest. Ent., 1985, 21 (4)

372

Kevan

apparently of considerably later date, though probably some 20,000 years old. This is in the
form of a picture, scratched on a bison bone, clearly representing a species of the camel-cricket
genus Troglophilus, which does not now occur in the region (Begouen & Begouen, 1928;
Chopard, 1928; Schimitschek, 1977)(Fig. lb). I know of little if anything else which antedates
the ancient civilizations of Near, Middle and Far East that is relevant to our present theme.

THE ANCIENT WORLD

From very early times (though there is little direct evidence from earlier than the 3rd
Millennium B.C.E.), scarab beetles were revered, depicted and modelled in Egypt as symbols of
Kheper (Fig. 2), a manifestation of the all-powerful Sun-god, Ra or Re (see, for example,
Newberry, 1905; Petrie, 1917; Bodenheimer, 1928, 1949, 1960; Efflatoun, 1929; Schimitschek,
1968, 1977; Harpaz, 1973). It is thus unlikely that the priestly class was entirely unaware of
the biology of such important creatures, parts of whose lives are intimately associated with soil.
Nevertheless, so far as I can discover, and despite implications repeated by Harpaz (1973) to
the contrary, there seems to be no written record of anything that may have been known at the
time (with or without religious or philosophical association), other than what the adult beetles
looked like and that they rolled dung-balls (Bodenheimer, 1928, 1949, 1960). Whether
“worms” attacking ancient Egyptian crops were specifically cutworms ( e.g Agrotis ypsilon ),
as suggested by Efflatoun (1929), is a moot point.

We can infer from ancient sources, dating back to the 2nd Millennium B.C.E., that cicadas
have been known from time immemorial to be part-time denizens of the soil, emerging in
“ghostly” or “spiritual” form, as from the grave, symbolizing, notably in China, purity,
immortality and/or resurrection after death (Brentjes, 1954, 1964; Schimitschek, 1968, 1977;
Kevan, 1978; Riegel, 1981). They also had less lofty significance in “magic” and medicine and
(as nymphs) as food (Chou, 1980; Riegel, 1981). The ancient Hellenes later, ultimately from
the east (H. Kuhn, 1935; Brentjes, 1954, referring also to two other publications by Kuhn from
1943), acquired a reverence for cicadas. Especially among the people of Attica and Ionia, these
insects came to symbolize an almost religious bond between man and his native soil. This sacred
significance did not, however, preclude cicada nymphs from being dug up in large numbers by
the ancient Greeks and used as food, as they were in China (cf. Kevan, 1978: 28, 29, 42, 45,
49). The oft-cited Athenian hair-ornaments called tettiges (i.e., cicadas) were probably based,
if any actual insects were involved, upon soil-dwelling nymphs and not on winged adult cicadas
(certainly not on grasshoppers as misguided western tradition has it!). This topic has been
briefly discussed fairly recently by Kevan (1978: 435-436), but see also Hauser (1906-1908)
and Brentjes (1954, 1958). Probably of similar antiquity to the Old World tradition, though
without tangible evidence of this, are the Amerindian legends of both cicadas and ants being
among the first creatures to emerge through the soil from the centre of the earth to populate its
surface (see Kevan, 1983b). The soil fauna is thus something that has always been of interest to
civilized as well as to primitive man.

Although legends and artifacts form a significant part of our source material relating to
early knowledge and belief, we tend to place greater emphasis on the written word. In this
regard, other than the earliest ancient Egyptian hieroglyphs for scarab beetles, and at least as
old in written origin (though not in existing writing) as some of these, the earliest known
references relating to our theme are those of the ancient Sumerians of present-day southern
Iraq. Some 4,000 or more years ago, at the very latest, these enlightened people were certainly

Soil zoology

373

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familiar enough with various members of the soil fauna to have applied different generic and
specific (binominal) names to a number of them and to have written these down. Of such
names we know a few from the Harra Hubullu, a compendium prepared in the 9th Century
B.C.E. by the successors and neighbours to the Sumerians, the Akkadians, giving equivalents in
the two languages (Landsberger and Krumbiegel, 1934; Bodenheimer, 1949, 1960; Harpaz,
1973)(Fig. 3). The Sumerians of the early 18th Century B.C.E. (the time relevant to the later
Akkadian text), and probably much earlier, distinguished between at least seven kinds of ants
or Kisi (including Kisi ririga, or flying ants, and Kisi kurra, light-coloured and perhaps
termites, not ants) and two kinds of earthworms of the annelid genus Mar, the Mar gal (or Mar
dib) or Mar tab and Mar Sasur. They also had binomina for what seem to have been a mole
cricket ( Gryllotalpa ), which they called Ub pad ; for (field) crickets, known as Buru
zapaag(-tira)) or Buru balag(-gana)) ( Buru being the generic name for orthopteroid insects);
and for a small, self-burying orthopteroid named Buru saharra , or “dust locust” (which, very
tentatively, I have identified elsewhere as the pyrgomorphid Tenuitarsus angustus
(Blanchard)).1 What the Sumerians or Akkadians knew about these animals, however, we have
no idea, though we can surmise that the former probably knew much more than history or
archaeology will ever reveal. After all, in the fifth part of their most ancient of epics, the
“Flood” legend of Gilgamesh, they apparently associated adult dragonflies with the moulting of
their aquatic nymphs (Sandars, 1959) thousands of years (so far as we are aware) before
anyone else did so.2

Not quite so ancient as Sumerian sources are the early Sanskrit Vedic books of late in the
2nd Millenium B.C.E. These refer not infrequently to ants and/or termites, as indicated by
various verse quotations given by Kevan (1978), though, apart from an association with
subsurface moisture in two examples, the references are little related to soil.3 Kevan ( op . c/7.)
also gives later examples from early literature (including old Tamil, or Sangum, and later
Sanskrit) relating to ants and termites, but mention of their role in the soil is again minimal.

Quaest. Ent., 1985, 21 (4)

374

Kevan

Fig. 4. “Ezekiel’s wheels” (see text), (a), (b) Artists’ fanciful interpretations of Ezekiel’s cherubim; (a) after G. Eicke,
1964, in Schimitschek (1968); (b) after W. Merian, 1650; for reference, see Hogue (1983). (c) Scarabaeus cicatricosus in
flight, showing salient features compatible with Ezekiel’s account; after Hogue (1983). Fig. 5. Chinese mole cricket,
Gryllotalpa orientalis, from the ancient illustrated encyclopaedia of Pu Shang, the Erh-Ya\ from a late printed edition of
about A.D. 1600. After Bodenheimer (1928).

Soil zoology

375

The allusions to cicadas and ants in early Hellenic literature of the Homerian and
immediately subsequent periods (9th-8th Century B.C.E.) are likewise mostly unrelated to the
soil, and it is only in considerably later works that we come across surviving references to their
“earthly” associations (see Bodenheimer, 1928, or, more briefly, Morge, 1973).

Although much Israelitic tradition is very ancient, most of the Hebrew scriptures as we
know them, with some exceptions, were written down no earlier, and often much later, than the
6th Century B.C.E. These Scriptures, though referring quite frequently to insects, make
virtually no unequivocal references to the soil fauna (the allusions to ants, in Proverbs VI, 6,
and XXX, 24-25, being near exceptions).4 Such references as there are to “worms” ( tolaath ),
where epigeic, plant-feeding insect larvae are not involved, are almost invariably to
flesh-feeding dipterous maggots. Bodenheimer (1928, 1929, 1960) reviews briefly the insects
referred to in the Hebrew Scriptures. There is, in addition, one brief reference, in the Book of
Micah, VII, 17 (“King James”, 1611 version: “... like crawling things of the earth they shall
come trembling out of their close places ... ”) that might conceivably refer to earthworms
(indeed an alternative translation uses “worms ... move”). The principal interest of this
uninformative passage lies in its possible antiquity. The book in question probably dates from
about 720 B.C.E., though its later sections (including Chapter VII) could well be by a later
author.5

Much more intriguing, on account of its controversial nature, is the somewhat later book of
the prophet Ezekiel, originally dating from about 590 B.C.E. The Egyptian scarab cult, to
which reference has already been made, eventually became widespread in the Near and Middle
East, including Babylonia, where Ezekiel, like other Israelites of note, was captive. It is possible
that, being a priest, he studied these religiously important insects out of interest, if not
conviction. It has been concluded (Sajo, 1910; Schimitschek, 1968; Hogue, 1983) that he gave
them the name cherubim , couching his description (made, probably, in the glaring sun) in
allegorical, pseudoreligious terms (as would befit such venerable creatures) to the subsequent
(possibly intentional) mystification of all and sundry (Fig. 4a, b). If an entomological
interpretation of Ezekiel’s cherubim be accepted, however, there is little in his account that
goes much beyond the identification and description of the scarab beetles (Fig. 4c) and their
dung-balls, though there is a hint of something more (Hogue, 1983).

In the 5th Century B.C.E., about a century after Ezekiel was allegedly peering myopically
at scarabs by the Chebar canal in Babylonia, the Chinese illustrated encyclopaedia of Pu
Shang, known as the Erh-ya, made its first known appearance (to be followed by numerous
editions throughout the centuries). In it, soil fauna, including mole crickets (Fig. 5),
scarabaeoid beetles (and their dung-balls and larvae), ants of several kinds, cicadas and
centipedes were all included (Bodenheimer, 1928, 1929), though how many of these were in the
“first edition”, I do not know. So far as I can tell, little was included on the direct soil
association of any but the mole crickets and cicada nymphs. Other ancient Chinese literature,
e.g., in the form of early “herbals” or pen-ts’ao , is referred to by Chou (1957, 1980) and
Konishi & ltd (1973), but the soil fauna is scarcely considered.

Although there were a number of early Hellenic literary references to insects and other
terrestrial invertebrates (see, for example, Bodenheimer, 1928; Morge, 1973; Kevan, 1978),
virtually none of the surviving writings mentioned soil-inhabiting animals, with the exception of
brief allusions to gigantic, subterranean gold-digging “ants” in “India”, which eventually
became the mythical “ant-lions”, and a hateful, biting creature known as the amphisbaina
(amphisbaena). The latter, mentioned by Aiskhulos (Aeschylus, 5th Century B.C.E.) in his

Quaest. Ent., 1985,21 (4)

376

Kevan

Agamemnon (see Druce, 1910), was traditionally (from later sources) a two-headed, poisonous
burrowing serpent. Of both of these denizens of the earth, more will be said later, but it may be
noted here that the giant ants are mentioned in the Histories Apodexis of Herodotos
Halikarnesseos (Herodotus) of the mid 5th Century B.C.E. (Rawlinson, 1910). Quatrefages
(1854) suggested that large termite mounds, rather than ant-hills, provided a basis for the
legend. Herodotos also mentions, in passing, the underground activities of Greek ants, but it
was not until the time of Aristoteles Asklepiados (Aristotle) that we have anything approaching
scientific observation.

Of Aristoteles’ five “notebooks” on zoology of about 320 B.C.E., four, now known by their
Latin titles of Historia Animalium , De Generatione Animalium, De Partibus Animalium and
De Incessu Animalium , the original Greek texts being long since lost, mention a few
soil-dwelling creatures (see D’A.W. Thompson, 1910; Platt, 1910; Ogle, 1911; Farquharson,
1912). Though western scholars have, for centuries, been “brainwashed” into accepting
Aristoteles as the founder of biology, he was really a late-comer, if an extremely important one,
to the scene. His contributions to various aspects of the science were undoubtedly of immense
significance, but he may well have transmitted many ideas from already ancient Middle
Eastern (or even Oriental) sources of which we have no record. (One can scarcely imagine that
the Sumerians, for example, did not bring about the dissemination of valuable zoological
information). For all his great erudition and commendable powers of observation, Aristoteles
contributed surprisingly little knowledge of the soil fauna (see the short entomological review
by Bodenheimer 1928, 1929, and the even shorter one by Morge, 1973, based upon it). Indeed
he had some very peculiar ideas on mould, decay and humification. With great originality (!) he
observed ( Historia Animalium , I, 1), that “some creatures dwell under ground, as the lizard
and the snake. ” (he had just mentioned that some provide themselves with homes, including
mole and ant, so he did not immediately cite these again as examples); other versions of the text
read: “some make themselves holes; others not so” (D’A.W. Thompson, 1910). He gave a
reasonably good, succinct description of the life-history of cicadas, including the subterranean
nymphal stage, or tettigometra (which he pronounced to be good to eat), and he also gave some
notes on the biology of scarab beetles ( kantharoi ) and their dung-balls, and on the life of ants.
Other soil fauna which he briefly described were myriapods, earthworms (“Yer emseqa or
“earth’s entrails”) and moles. Both millipedes and centipedes were said to remain active after
being cut into pieces (though there seems to be some confusion here with their marine,
annelid- worm, counterparts). His two main references to earthworms were oddly confused with
the origin of eels. Moles, he stated, cannot burrow if they are transported from one location to
another. Aristoteles also mentioned certain “marginal” soil animals: woodlice, scorpions,
pseudoscorpions, mites on insects (Oudemans, 1926), and digger wasps, burrowing bees and
bumblebees. He did not mention the amphisbaena. His disciple Theophrastos, though making
valuable contributions to entomology in the course of his botanical studies, did not refer to
subterranean insects except for (cut)worms (?) that attack both roots and stems of wheat.
Later, we have the opinion of Kleanthes, about 270 B.C.E., that ants behave only instinctively -
though Plutarch, about 100 A.D., believed later that they reacted intelligently (see
Bodenheimer, 1928; Morge, 1973). Shortly before 200 B.C.E., the Roman playwright Titus
Macchius Plautus (in Mostellaria, III, 2) refers to “ tarmes ” (some kind of wood-feeding
insects, conceivably termites) boring from below ground (Kevan, 1978: 425).

The Macedonian physician Nikandros (Nicander) of Kolophon, in his Theriakos and
Alexipharmakos of about the mid 2nd Century B.C.E., wrote extensively (in verse) on animals.

Soil zoology

377

Fig. 6. The dreaded, two-headed, subterranean Amphisbaina (see p. 375): illustrations from Byzantine manuscripts
descended from Nikandros’ Theriaka of 2nd Century B.C.E., taken from Kadar (1978 : pi. 8, 55, 101). Top, from Cod.
Paris. Suppl. Gr. 247, 10th Century; middle, from Cod. New York Pierpoint Morgan M. 652), 10th Century; bottom, from
cod. Bonon. (Bologna) Bibl. Univ. Gr. 3632, early 15th Century.

without knowing very much about his subject. He had much to say about serpents and
scorpions, and, with reference to the former, there is, in The riakos, a formula for utilizing the
skin of the abhorrent amphisbaena against various maladies (cf. Druce, 1910; Kevan, 1978; 63,
444; the latter gives a fragment of the Greek original with a somewhat “unorthodox”,
inaccurate English translation from a Latin version). Much the same is given (in prose) by his
approximate contemporary, ApollodOros of Athens, in his Bibliothe ka, and it would seem likely
that this was Nikandros’ immediate source. The rather small, poisonous, half blind
underground serpent (Fig. 6), with a head at each end and the capacity to progress in opposing

Quaest. Ent., 1985,21 (4)

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dnando fuo lolirum po:cac in oic ribum

geftis Oenocat

3ncnfuio facttercsmonUratfpinimaUs

»frcs ^ Piligir (dco

0U39 Cquia iudeus non amaO.indcmm

vatmbmincfim fccura fumre

e'eft ef formica

Dumcaloun terra non requiefeit ca

b C

Fig. 10. Illustrations of (presumably) the isopod crustacean Armadillidium vulgare (Onoi hoi hypo tds hydrias) from
10th-Century Byzantine manuscripts descended from Dioskorides’ Pharmaka of 1st Century A.D., taken from Kadar
(1978: pi. 72, 89). Above from Cod. New York, Pierpoint Morgan M. 652; below from Cod. Vatican. Gr. 284. Fig. 11.
Part of the account of the Ant from a late metrical version of the Physiologus attributed to Bishop Theobald of Monte
Cassino, early 11th Century. From the printed version, Physiologus Theobaldi Episcopide naturis duo decim animalium,
Koln, 1492 (see Rendell, 1928).

Soil zoology

381

directions (simultaneously ?!), seems to have been known and feared since before the time of
Aiskhulos and Herodotos, though Nikandros did not mention its venom, or its mode of
progression. It was probably based originally on the harmless burrowing, worm-like reptile we
now call Trogonophis (the related modern genus Amphisbaena is tropical), rather than on the
blind-snake, Typhlops, which was also known at the time.6

Nikandros also made various references to insects, though nothing definite was said about
their relationships with the soil. Judging by Byzantine illustrations (Fig. 9), copied from
generation to generation (see Kadar, 1978), he seems to have known, in the Aristotelean
tradition, about centipedes ( skolopendra ) and possibly millipedes (foulos), though both may
more usually have been (? marine) annelids. (With reference to the latter, see discussion of a
Greek 2nd Century A.D. poem by Neumenios, in Kevan, 1978: 334). Kadar (1978) indicates
that another, later and much better-known physician, active in the middle of the 1st Century
A.D., Pedanios Dioskorides, also knew of some soil animals, such as earthworms and the others
referred to by earlier authors, including Aristoteles and Nikandros (Fig. 9, 10). These he listed
in his Pharmaka ( De Materia Medico). Woodlice of the genus Armadillidium (presumably)
were apparently known to him as onoi hoy hypo tbs hydrlas. In the 1st Century also, the poet
Marcus Annaeus Lucanus (Lucan, 39-65 A.D.) emphasized the venomousness of the
burrowing amphisbaena in his Pharsalia (see Druce, 1910).

The Aristotelean tradition, somewhat embellished, was carried on by the Roman Gaius
Plinius Secundus, or Pliny the Elder (77 A.D.; see Rackham, 1940). Though a great compiler,
he was unoriginal and often gullible. In his Naturalis Historia, he had little to say (and nothing
new) about the soil fauna, though he did mention cicadas (including their subterranean
tettigometra nymphs) and scarabaeoid beetles. He also seems to have been responsible (after
Strabon, or Strabo, in his Geographikos of 23 A.D.) for “popularizing” the early myth of the
fabulous, gigantic, dog-like, gold-digging “ants” of “India”, to which reference has already been
made (see above, and George, 1981). The “ant-lion” {myrme coleon, later mirmicoleon, etc.,
sometimes also called “ant-dog”) made its way into early versions of the symbolistic Christian
Physiologus (below and cf. Fig. 12), to which the lost 2nd-Century Greek Peri Zoon of the
Syrian monk Tationos may also have contributed.

“ Physiologus ,” or “The Naturalist, “ was presumably originally the pseudonym adopted by,
or for, an unknown compiler of the book that bears the name. The latter originated in the
eastern Mediterranean (Alexandria or Syria ?), probably towards the end of the 1st or in the
early 2nd Century A.D., but the Christian Physiologus perhaps came a century or so later.
Extant material goes back to about the 5th Century (James, 1928; Rendell, 1928;
Bodenheimer, 1928; McCulloch, 1962; George, 1981). It was copied in several versions and
eventually led to the Mediaeval “Bestiaries” ( Libri Bestiarum). The earliest Greek versions
included 63 chapters, of which 56 were devoted to different animals (C. Peters in Bodenheimer,
1928; Morge, 1973). Of these, only the ant, the “ant-lion” and the scarab concern us here. The
second of these, though fabulous as presented, and having mythical attributes, had, like other
strange animals in the Physiologus , a basis of factual existence.7 As explained by George
(1981), this was probably the badger-like ratel ( Mellivora capensis ), which burrows in sand
and soil, though the pangolin ( Manis ) had been considered previously to be a likely candidate
(Rawlinson, 1910, and others noted by George, 1981).

Before concluding our brief review of the so-called “classical” period of the Western World,
we should perhaps note that there were various other works that had a bearing on applied
entomology, from Cato, 235 B.C.E., to Palladius, ca. 380 A.D. (Bodenheimer, 1928; Morge,

Quaest. Ent., 1985, 21 (4)

382

Kevan

1973). Although virtually none have any direct relation to the present context, we should
perhaps mention the account of field crickets given by Publius Nigidius Figulus, mid 1st.
Century B.C.E., in his De Animalibus (cf. Wotton, 1552; Bodenheimer, 1928), as this was
distorted much later by Rhabanus Maurus (see p. 384?). Field crickets burrow backwards in
the soil (and chirp at night); they may be hunted by inserting an ant on a hair, blowing away
the dust the while; they can then be dragged forth together with the ant [which clings to it.
Greek references to catching crickets in much the same way, by means of a strand, go back
much further into antiquity.] Collumella, ca. 50 A.D., mentions ants, snails and miscellaneous
caterpillars. Aelianus, about 200 A.D., dispensed numerous moral tales involving animals,
including ants in particular, but also the “ant-lion” and amphisbaena, but these had not even a
pseudoscientific signifance. Gaius Iulius Solinus, in his Collectranea Rerum Memorabilium
(later Polyhistor ) of the second half of the 3rd Century, transferred the ant-lion from India to
Ethiopia. He also maintained that the amphisbaena had two heads.

In the Orient, this late classical period of the Occident was not particularly notable for
known observations on soil-inhabiting animals, though, in China, there was, as in the West, a
keen interest in “herbals”, or pen-ts’ao, which included medically important insects. The
earliest proper pharmacopoeia, the Shen-nung Pen-Tsao Ching was apparently compiled about
the 2nd Century A.D. (Chou, 1957, 1980; Konishi & Ito, 1973). Scarabaeoid beetles and mole
crickets were among the soil-inhabiting fauna mentioned.

THE EARLY AND MIDDLE MEDIAEVAL PERIODS

By the second half of the 5th Century A.D., with the rise of barbarism and the demise of
“classical” traditions, European culture was sinking to its lowest ebb. Give or take a century or
so, this was also true, to a greater or lesser degree, of most other civilizations including that of
China (and possibly also in the Americas). In about 500 A.D., however, another edition of the
Chinese Shen-nung Pen-Tsao Ching pharmacopeoia by T’ao Hung-Ching was prooduced
(Konishi & Ito, 1973), though it does not seem to have added much actual information to the
2nd Century version already noted.

For long after the decline of the Western Roman Empire there was no science in Europe,
though some semblance of culture and scholarship eked out a rather precarious existence in
Ireland (devoid of Greek) and in the Eastern (Byzantine) Empire. In zoology, apart from some
tanscriptions of old Hellenic works in the latter region (cf. Kadar, 1978), only the Physiologus
persisted, but even that was placed on the list of proscribed and heretical writings by the
Roman Church in 496 A.D. The ban was not lifted for just over a century. The earliest
Mediaeval Latin versions of the Physiologus , judging by the oldest surviving copies now
available (8th and 9th Centuries) varied little in substance, accounting for some 43 animals
(mainly mammals and birds, as noted by George, 1981) in 48 or 49 chapters (see also James,
1928; T.M. White, 1954; McCulloch, 1962). There was thus a slight reduction from the “late
classical” 56 animals already mentioned for the Greek text by Bodenheimer (1928) and Morge
(1973). The Latin versions, for example, did not refer to scarab beetles. Among soil
inhabitants, these insects symbolized heresy and their dung-balls evil thoughts; and ants were
symbols of provident virtue (encouragement of the “work ethic” - which some to-day might also
regard as heresy! - among the peasantry being important politically to both religious and
secular institutions). Ants were also noted for their wisdom, particularly, in the present context,
for their astuteness in biting grains in two to prevent their germination when stored in the soil.

Soil zoology

383

giuetuie roiot cn gnu ttntoun ti a

rtumful>l4bit$coatm. M3

Fig. 12. “Ant-lions” or “ant-dogs” (see text, p. 383) from an early 14th-Century Anglo-Latin “Bestiary,” in the British
Library, London (MS Royal, 2B, VII, fol. 96; cf. George, 1981).

(That ants do neither this, nor reject barley in preference to wheat, was immaterial). The
“ant-lion” (or “ant-dog”), derived from the gold-digging ants of the ancients (cf. George,
1981), was hybrid - yes, between lion and ant! - that symbolized man’s ambivalence: its
carnivorous front part, dominated by its vegetarian rear part, and vice versa , meant that it
could eat neither meat nor plant material and so, though paradoxically surviving, it perished
due to starvation soon after birth! Unlike its later namesake it did not live in pits in sand or soil.
At least one Mediaeval illustration later showed it as a dog-like mammal inhabiting mounds of
earth (Fig. 12).

By the 7th Century, the great plague of the mid 6th Century in the Mediterranean region
had come and gone, and the worst was over - “culturewise” at least -in both Occident and
Orient. It was early in that century that the glimmerings of biological science began to revive
when Isidoro de Sevilla (Isodorus Hispaniensis) produced his encyclopaedic Origines sive
Etymologiae. This not only borrowed from, but was later to contribute additional material to,
the Physiologus , resulting in the development of the second “family” of Libri Bestiarum or
“Bestiaries”. Amongst the animals considered by Isidoro were the amphisbaena, the mole, and
a handful of invertebrates, including earthworms and one or two soil-dwelling beetles
(Bodenheimer, 1928, 1929; Morge,1973). Isidoro’s “cicadas”, however, were Cercopidae
(originating in the saliva of cuckoos, not in the soil like true cicadas). In passing we might also
mention, in the 7th Century, Aldhelm (639-709 A.D.), England’s first great scholar and senior
contemporary of Northumbria’s Baeda or “Venerable Bede”. When prior of Malmesbury,
Aldhelm composed, in Latin verse around 695 A.D.,8 his famous 100 “Riddles” ( Aenigmata
Aldhelmi) as part of his Epistola ad Acircium (Letter to Aldfrith [King of Northumbria]; see

Quaest. Ent., 1985, 21 (4)

384

Kevan

Pitman, 1925). The reason for referring to Aldhelm here is not that he really mentioned the soil
fauna, but to draw attention to a general omission in histories of biology. Aldhelm, though not
prolific in the field, was one of the few first-hand recorders of nature during the millennium
since Aristoteles. His only riddle remotely associated with soil fauna uncharacteristically
concerned the “ Myrmicoleon ” or “ant-lion” in the mythical, symbolistic tradition of the
Physiologus.9 Baeda (Bede, 673-735 A.D.) in his Natura Rerum , of about 725, did not, so far
as I know, refer to the soil fauna at all.

Also, in passing, we might mention the anonymous Old English epic poem. The Deeds of
Beowulf probably the oldest surviving major poem in a western “modern” language. This deals,
in part, with events of the early 6th Century, but was apparently composed in the late 7th, or
more likely early 8th Century (the only known manuscript is late 10th Century). Beowulf, the
mighty hero, was eventually wounded by a gigantic, fire-breathing, subterranean Wyrm or
Worm (alternatively, Dragon - see Earle, 1892), which may be equated with The Mediaeval
“ Daemon subterraneum truculentus ” (see footnote to Table I).

In the first part of the 9th Century, the German bishop Rhabanus Maurus completed his De
Universo , which, though it drew heavily on Isidoro de Sevilla, was a much more erudite work
than his. In it (Bodenheimer, 1928; Morge, 1973) he mentioned “ vermes ” of various sorts
(including anything from fleas to clothes-moth larvae), some of which may have been true
(annelid) earthworms or possibly terrestrial beetle larvae. He also referred to “ scarabaeus ”
beetles ( Geotrupes ), to (field) crickets ( Gryllus , s. str., which burrow backwards into the soil
and which are hunted by ants wielding hairs - a distortion from Nigidius, see p. 382), and to
ants, with their various virtues. These last also included the fearsome, Indian giant gold-diggers
of the ancients, formerly confused with “ant-lions”, and transferred by him to “Aethiopia” (in
accordance with Solinus, antea, and Isidoro). For apparently the first time, too, a true insect
ant-lion ( Myrmeleon )10 was mentioned, under the latinized name of formicaleon (perhaps to
distinguish it from the mythical myrmicoleon). It is described as a veritable lion amongst ants,
burrowing in the dust and killing its victims as they carry along their loads. His “ cicadae ”,
however, like those of Isidoro, were Cercopidae and their nymphs not soil-dwelling.

By contrast, in the early 9th Century (and probably long before), the development of cicadas
(ts’an) from eggs laid in the soil was widely known in China, as exemplified by a poem by Po
Chu-I, quoted by Kevan (1983a: 42-43). Kevan (op. cit .) also quotes other Chinese and
Sanskrit poems of the period (late 8th to 9th Centuries) that refer to cicadas, mole crickets,
termites and/or ants, though few are pedologically oriented.

A notable western scholar of the middle 9th Century was the Irishman, John (the) Scot
(Johannes Scotus Erigena, ca. 810 - ca. 877; the Scots, sensu stricto, originally came from
Ireland!). His De Dmsione Naturale, written betwen 865 and 870, included much original
thinking - presumably contributing to its subsequent condemnation by the Roman Church -
but, as it drew mainly on “Pseudo-Dyonisius” and similar authors of antiquity, it again gives us
nothing to note on soil fauna. The 9th Century was also notable for the rise of Saracen11
scholarship. Early in this period, there were translations into Syriac and Arabic of old Hellenic
writings, including those of Aristoteles, now lost in the original. In the middle of the century,
however, at about the time that John Scot was most active, an independent zoological work, the
Kitabal-Hayawan ( Books on Animals ), was compiled by Al-Gahiz (or Aljahid). Regrettably it
too, included virtually nothing on soil animals, other than some generalities on beetles in Book
3, and on ants in Book 4 (Bodenheimer, 1928). The later, better-known author, Ibn-Sina
(“Avicenna”), of the late 10th to early 11th Century, was, it seems, merely a translator, whose

Soil zoology

385

most valuable contribution was to be among those who helped to preserve the writings of
Aristoteles. He did, however, discuss the amphisbaena, whose Arabic name was given as
auksimem. During the 10th Century, too, other old Hellenic texts were being transcribed under
the influence of the Byzantine rulers of Constantinople (Kadar, 1978), but nothing original
transpired. Thus it was that, by 1000 A.D., soil zoology, like most other scientific disciplines,
had progressed little further than these early works - where they had not, in fact, retrogressed.

The 11th and 12th Centuries, pedobiologically, were no more fertile, though imagination
and moralizing (e.g., in respect of ants) increased slightly in the shrunken Physiologus (see, for
example, James, 1928; Rendell, 1928; McCulloch, 1962) and the appearance in church
architecture of relatively uncomplicated forms of the amphisbaena (Druce, 1910). We may,
however, mention a few works marginally associated with soil fauna, though the late
1 lth-Century comments by Shlomo Jizechaki (or Rashi), on insects mentioned in the Talmud,
cited by Morge (1973), do not seem relevant. In China, there was a revived interest in
pharmacopoeias and the old pen-tsao’s were restructured along taxonomic lines in the form of
the Cheng-Lei Pen-tsao ( Reorganized Pharmacopoeia ) by T’ang Shen-Wein in 1108 (Konishi
& I to, 1973). This discussed, amongst vermin and other lowly creatures, scarabaeoid beetles
and mole crickets (Bodenheimer, 1928, 1929). Not long afterwards, in Germany, the
Benedictine abbess, Hildegard (“St. Hildegardis”)12 began compiling her Libris Physicis , which
may be said to date from about the middle of the century. Her work differed from earlier
“herbals” as it was based on personal experience and local usage, not upon established
“authority” and hearsay. Field crickets (which she called “cicadae”) had certain medicinal
properties; and she also mentioned ants.

Another mid- 12th Century author, of great erudition (according to himself) and extreme
verbosity, was Ioannes Tzetzes of Constantinople, whose enormous metrical (one cannot say
poetical!) work, Biblos Histdrike (commonly called Chilediades), written about 1165-1170,
included a fair amount of animal lore among his (un)natural history verses. However, he said
nothing not already written by earlier authors. Examples of his writings (on cicadas, though not
in soil) are given by Kevan (1983a). The mid 12th Century also produced the work of the
Saracen scholar Ibn-Rashid (or “Averroes”), another major translator of Aristoteles, but not, a
contributor. In this period, too, we should mention the credulous Anglo-Latin work of the
Englishman Alexander Neckam, De Naturis Rerum , of 1170 {cf Wright, 1863a; Raven,
1947), and of the even more credulous Norman-Welsh Silvester Gerald de Barri, or Giraldus
Cambrensis, Topographia Hibernicae , of 1182 {cf. Wright, 1863b; Raven, 1947), if only in a
negative sense. Though both works refer quite extensively to natural (and unnatural) history,
including mention of insects, spiders and other invertebrates, the former refers, among
soil-dwellers, only to the mythical amphisb(a)ena and the “seps” (probably based on a gecko,
but which could mean almost anything from a poisonous serpent-lizard to a woodlouse or a
myriapod), and the latter to the badger which is said to dig burrows in the earth.

The late 12th Century was the time when “Bestiaries” {Libri Bestiarum) not only started to
become more elaborate, but when the numbers and complexity of “species” (real as well as
“derived”) mentioned therein increased {cf James, 1928; T.M. White, 1954; McCulloch,
1962). In the early 12th Century, they had typically included relatively few (about 36)
chapters, like the Latin Physiologus. By some curious turn of events (most likely due to
inadvertent omission of a passage by some copyist, though I have not seen this theory
advanced), the “ mermecolion ” later generally became confused with the margarita , or pearl, as
was the case in the 12th-Century manuscript discussed by James (1928) and T.M. White

Quaest. Ent., 1985,21 (4)

386

Kevan

l? auifi uvidfcJr. tUri didbtffanob; iff wWi uvpul<

or 6ici r~r r —

ORitntt
dettkU)

u><Vpi3

priiujlraf

*w few

a&4 dt&mtoo aa ftojpuom eft <jd> vt*u

Fig. 13. “Millipede” (resembling Glomeris ; possibly an isopod crustacean Armadillidium) from 13th/14th-Century
manuscript in the British Library, London (MS Harley 3244), apparently copied from an earlier (12th Century) Latin
“Bestiary” (probably English). In the present manuscript, above this illustration is one of “ vermes ” (“life history” of
earthworm) and another of spiders (with 7 pairs of legs!); below are mouse-like (or more probably shrew-like) “scorpions”
(named in bottom line of text as shown); most of the above will be found (in white on black) in Davis (1958). Fig. 14.
Industrious (8-legged!) ants carrying “grain” (pupal coccoons). From a Mediaeval “Bestiary” of (?) 13th Century
(Pierpoint Morgan Library, New York, MS. 81, f. 31 1 - cf. Rowland, 1973).

Soil zoology

387

(1954). Thereafter it tended to disappear altogether. The manuscript mentioned above included
some chapters relevant to the present context: on mole, ant, “ amphivena ” (amphisbaena, but
winged and no longer soil-inhabiting - cf. Fig. 7, 8) and “vermis”. The last included
earthworms, but also (in the tradition of Isidoro and Rhabanus Maurus) a wide range of
arthropods, amongst which were scorpions, spiders, “millipedes” that rolled up into a ball (i.e.,
either Glomeris diplopods or Armadillidium isopods - cf. Fig. 13) and “termites” (by which
seemed to be understood, almost any kind of wood-feeding insect other than true termites!).

In China, during this period, versions of the Erh-ya encyclopaedia and the pen-tsao
pharmacopoeias with their occasional references to soil fauna continued to appear.

THE LATER MEDIAEVAL PERIOD

We may continue the story in China with a single reference of marginal pedobiological
interest. Ever since the later T’ang-dynasty period (8th Century), crickets had been admired
and kept for their songs, but, by later centuries, cricket fighting had become an important part
of Chinese culture. As large wagers were made on the outcomes of the encounters, much care
was lavished on the contestants. This demanded a basic knowledge of cricket biology
(particularly as regards their care and maintenance). As fighting crickets are all
ground-dwelling species, a number of which burrow in soil, a fair amount was known of such
species. An extensive manual on the subject was written by a member of the Sung-dynasty
court, Kia Se-Tao, at the beginning of the 13th Century. It was called Tsu-chi King , or The
Cricket Book (see Chou, 1957, 1980; Petit & Theodorides, 1962). Needless to say there were
successors in Ming-dynasty and later times.

With reference to crickets, it is also interesting to note that these were mentioned in the
longer, 71 -chapter, version of Bestiaire written in northern France before 1218 by one Pierre
(called “le Picard” or “de Beauvais”). The insects were called cri(s)non or gresillon and were
said to sing so much that they lose their appetites, forget everything else, let themselves be
hunted and die singing. (This is really a distorted cicada myth.) One 13th Century manuscript
of this work illustrates the cricket in front of a hole in the soil, though a 14th-Century one
shows crickets on a hearth (McCulloch, 1962). 13

In 13th-Century Christian Europe, though the “Bestiaries” (Fig. 14) remained the main
sources of zoological (mis)information, scholarship began slowly to emerge from the stagnant
morass into which it had sunk. To some extent this resulted from, and in others it paralleled,
the Saracen advances in knowledge and the rereading of classical authors. Three major
encyclopaedias compiled by members of the Christian Dominican order, and one by a
Franciscan, all written between 1230 and 1270, referred to a few members of the soil fauna.
The works are briefly reviewed by Bodenheimer (1928, 1929) and, through him, by Morge
(1973). They are those of Thomas de Cantimpre, or Catimpratornus ( Liber de Naturis Rerum ,
1233-1248), of his apparent mentor, Albert von Bollstadt, or Albertus Magnus ( De
Animalibus, in his Opus Naturarum), 1255-1270, of Vincent de Beauvais, or Vincentius
Bellovacensis ( Speculum Maius Tripartitum [naturale, historiale et doctrinale] , the relevant
parts, I, Books 17-23, also ca , mid 13th Century), and of Bartholomew (Glanville? the)
English, or Bartolomaeus Anglicus ( De Proprietatibus Rerum , of roughly the same date - see
Raven, 1947). These authors, between them, mentioned moles, earthworms, amphisbaenas,
ants (including their larvae and pupae), true ant-lions, crickets (often confused with cicadas),
various beetles (including ground-beetles) and their larvae, and so on, but, apart from Albertus’

Quaest. Ent., 1985, 21 (4)

388

Kevan

&yj*y ija%) k>V^^J^J
•‘^jj > boA iMy iy a^U- Jeij ^ l^A I %L|I;>)

15 &0b am* i

opim-' <i5

# Y>,/n*cuf vermis toryvrr cyiqnms (td .irumo

'N- - cIjruB dicintr cnim cum fcrp.tinhnf'htahrrt
u'rttmen ft p/erumy, in^vnu, c*of Inpcrxrr pu
^tic <>mino

Fig. 15. Scarabaeoid beetles ( khunfusa ) from the Arabic manuscript (Munich Codex) of H. Al-Qazwini’s
Nuzhat-ul-Qulub, originally written in 1341. After Bodenheimer (1928), who says that the illustration probably
originated during the lifetime of the author, and perhaps under his supervision. Fig. 16. The Bishop of Lausanne
excommunicating cockchafers in the 15th Century. Copied from a contemporary illustration, after Bodenheimer (1928).
Fig. 17. Mole-cricket, Opimacus , now Gryllotalpa\ water-colour from Book IV of the Codex Animalium of Petrus
Candidus Decembrus, ca. 1460. After Bodenheimer (1928).

Soil zoology

389

denial that the amphisbaena had two heads, they still really had nothing to say on these animals
that had not been said previously, mostly by the Physiologus and by Aristoteles.14 In the very
early 1300’s (1304-1309), Pietro de Crescenzi (Petrus Crescentii), in his Ruralium
Commodorum Libri XII, dealt with crop pests, though mostly on the basis of reports by
classical authors. Once more, soil-inhabiting forms do not appear to have been considered,
though he did recommend certain remedial measures for the control of ants (Bodenheimer,
1928; Morge, 1973).

Meanwhile, the Saracen scholars were gradually expanding knowledge in many fields
(though scarcely in relation to soil fauna). The cosmography of Zakartya bin-Mohammad
bin-Mahmud Al-Kummunt Al-Qazwint (cf Wiedemann, 1916; Bodenheimer, 1928), the
'Aja'kh al-Makhluqat (Wonders of Creation ), completed in 1263, refers briefly to earthworms,
ants, scarabaeoid beetles and crickets. This work was drawn upon and expanded by another
Al-Qazwtnt (Hamdullah Al-Mustaufa of that ilk) in his encyclopaeida, Nuzhat-ul-Qulub
(Hearts' Delight ), of 1341 (cf. Stephenson, 1928). Soil animals mentioned included termites
(aradat; they eat earth and are attacked by ants), woodlice (/z/w5ru-l-qabban), earthworms
( kharatin ; with medicinal and aphrodisiac properties), beetles (khunfusa; including small
scarabaeoids, Fig. 15), various “worms” (dud; including insect larvae of divers kinds, some
subterranean), crickets (sarsari in Persian; tatuk in Arabic; with medicinal properties) and ants
(naml; various kinds enumerated).

A little prior to this work, in 1320, we have what is probably the first involvement of the
Christian Church in the control of soil pests - though against the aerial adults - the
excommunication of May-beetles (Melolontha) at Avignon. Similar exercises in exorcism (Fig.
16) continued for centuries, since pest outbreaks always diminished thereafter - eventually!

One of the landmarks of Mediaeval biological literature was undoubtedly the great
zoological lexicon, the Hayat al-Hayawan (Life of Animals), completed in the late 14th
Century, by the Egyptian scholar Kamal Ad-Din Ad-Damirt (see Jayakar, 1906, 1908;
Bodenheimer, 1928, 1929; Morge, 1973). Soil-inhabiting animals were mostly of the same
kinds, with much the same information, as included by the Al-Qazwini’s: Termites (al-'arada,
as-surfah), various insect larvae (al-asan\ including some subterranean), field crickets
(al-gudgud, sharrar al-lail), scarab beetles (al-gua\; dung-feeding by larvae noted); “worms”
(ad-dud; including earthworms and a range of insect larvae, but also termites and [parasitic]
nematodes), a “worm” that rolls up in a ball (ash-sha‘hamat al ard; either an
Armadillidium-Mke isopod or a Glomeris-Yikz millipede), woodlice (/zzmar-kabban), earwigs
(al-‘ukuban) and dung beetles (qish'iban), as well as a whole range of ants (naml, generally;
al-gathlah, black; ad-dinnah; ad-dharr, small, red; as-simsimah; ash-shaisaban, male;
at-thathrag-, al-'ugrut, ? carpenter; ‘aygabuf; hayzabun; al-fazir; muq, winged; and
heigemana, very small). Although Ad-Damirt was comprehensive, he was not particularly
innovative or informative, especially in terms of soil-fauna relationships.

In these times, also, other writers of the Islamic world mention something of various
agricultural insect pests, but, to all intents and purposes, relating only to those above ground.
Several of the zoological and agricultural works, and copies later made from them, included
illustrations of the animals. These were, however, seldom, if ever, drawn from nature - locusts
were often bipedal and like birds, and (almost in the present context) crickets quadrupedal and
like newts!

When considering the 14th Century, one should perhaps not be surprised at the lack of
progress for, in the very middle of it, came the Black Death. This was by no means confined to

Quaest. Ent., 1985,21 (4)

390

Kevan

Europe, but it was most terrible there, especially from 1347 until 1350 This plague (followed
by severe typhus epidemics) had dreadful and lasting consequences for human activities . of all
ldnds including scholarship. At least a quarter, and probably a third of the entn e populatio
Europe died. Like Ad-Damtrt in Cairo, Cunrat von Megenberg, who translated Thomas de
Cantimpre’s De Naturis Rerum into German {Das Puch /= Buch] der Natur) about this time
(see Note 14), survived in Regensburg, but many scholars did not. Then, a century later, came
another major catastrophe for Christendom (though less so for scholarship); the Byzantine

Empire fell to the Osmanli Turks in 1453. f

Scholastic recovery from this second disaster was not so slow as from the Black Death,
the Renaissance, spurred on by the resulting economic revolution in its wake was alrea y
beginning. The Middle Ages had come and gone, and within less than half a century, both in
the Far East (where it had long been known) and in the West, the era of the almost untversal

use of printing had arrived. . Kllt

Belonging to the scientifically rather sterile transitional period of the early Renaissance,
Mediaeval in tradition, we may note in passing the beautifully Mustr ate l Codex
Petrus Candidus Decembrus, about 1460 (cf. Bodenheimer, 1928, 1929; Merge 1973). This
drew heavily on the 13th-Century European encyclopaedias (and on old Pl.mus Secundus.). ^
contributed nothing new, but, in its fourth volume, it included references to various worm
and insects, such as ants and “cicadas” (really crickets), and excelentwatercolou^ofamole
cricket (called Opimacus , Fig. 17), and of a true (if 4-legged) ant-1, on larva (set i Bodenhe mer
1928) Here, too, it would seem appropriate to place what would appear to be the first de
evidence since the ancient Sumerians of discrimination between species of earthworms.
Treatyse of Fysshynge wyth an Angle, attributed (possibly erroneously) to Dame Julyana
Barnes (born ? ca. 1388), Prioress of Sopwell Nunnery, Hertfordshire England, recommends
the “great angle Twytch” (probably Lumbricus terrestris or Allolobophora
catching eels, but “red” worms for all other fish. A manuscript (perhaps of as late as 1479 a

versions also exist, that under the name of Julyana Berners (1496) being the earliest.

THE RENAISSANCE

The European Renaissance was no sudden phenomenon, and it developed at different times
in different places, but we can think of it as occupying much of the 15th through to he ^m d
of the 17th Centuries. In the Orient, also, there were roughly coincident changes in Chinese
philosophy] but these were not so marked nor did they so radically affect attitudes toward
learning in general and science in particular. The European entomological and assoc ated
literature of this period is briefly reviewed by Beier (1973), though he makes wtually
reference to soil fauna. Some scattered information on the topic is, however, b

Bodenheimer’s “History of Entomology” (1928, 1929). less

Immediately after the widespread adoption of printing, already mentioned, her was les

immediate change in biological knowledge than might be anticipated, fnd °ld
prevailed in printed, rather than manuscript form. Bartholomaeus Anglicus mid-1 3th-Centu y
De Proprietatibus Rerum appeared in a first printed edition in 14 , an unra

Megenburg’s mid- 14th-Century Das Puch /= Buch / der Natur in 1475 became the
illustrated, printed natural history book (see above and Fig. 18). Not long afterwards appeared
(Fysshynge Wyth an Angle, see above) and the first edition (of many) of Or, us (or Harms)

Soil zoology

391

Quaest. Ent., 1985, 21 (4)

Fig. 18. Folding woodblock “plate” including ants and earthworms, from Cunrat von Megenberg’s (1475) Das Pitch der
Natur from Mss. of the middle of the previous century. Fig. 19. Ants emerging from the ground, illustrating the section
“Formica” in an early printed version of Ortus Sanitatis (see p. 390), ca. 1500. After Bodenheimer (1928).

392

Kevan

Fig. 20. Mushrooms, earthworms, snail, etc., 16th Century. Above, woodcut from Lyons (French) edition, 1572, of
Mattioli’s (1548) Commentario. Below, embroidery (34x34cm) based on the same, by (or under the direction of)
Elizabeth, Lady Shrewsbury (“Bess of Hardwick”), ca. 1580; one of a series now at Hardwick Hall, Derbyshire, England;
note prominent isopods above the earthworm. [From a photograph.]

Soil zoology

393

GEORGII A G R 1>

COLAE DE ANIMANTIBVS
fiibtcrrancis Liber.

orpvs fubterraneum, ut
res ip fa demonftrat,in ani-
matum diftribuitur,8£ina-
nimatum . quod autem ani
mi expers eft, rurlus diuidi
turinid quodfuafponte erumpit exter
ra,&Tnidquod exeadcmeffoditur.De
altero inanimi genere dixi in quatuor li-
bris de natura eorum quae effluut ex ter-
ra infcriptis,dealtero in decern denatu-
ra foftilium : nunc de fubterraneis ani-
mantibus dicam. Cum uero genus ani-
mantium omne conftet ex quatuor de-
mentis , 8C corpus humidum ac ficcum,
id eft aqua 8C terra, ad accipiendum apta
fint,necefte eft ea ipfa duo elementa ani
mantium matcriam die . Ex quo rurfus
illudquadam naturae neceffitate confe-
quitur , ut omne animal dC in aqua uel
terra gignatur, &C in eis commoretur at-
queuita fruatur*Nam beftiae uolucres,

a 4 etfi

Fig. 21 . The first page of text from Georg Bauer’s De Animantibus Subterraneis (Agricola, 1 549; Preface dated 1 548).
[Facsimile now in Macdonald College Library.)

Quaest. Ent., 1985,21 (4)

394

Kevan

Sanitatis , a sort of herbal-cum-pharmacopoeia based upon the various Mediaeval
encyclopaedias already mentioned. It has been attributed to Johann Wonnecke (or Dronnecke)
van C’aub (Johannes de Cube or John Cuba), though some consider him to have been merely
the translator of the work into German (cf. Wonnecke van C’aub, 1480, 1485). Jacobus
Meydenbach (1491) has also been credited with the authorship, though he was but the editor of
a somewhat later, better known, edition. Bodenheimer (1928, 1929) briefly reviews the latter
and later editions from an entomological viewpoint. (Insects were not mentioned in the smaller,
earlier edition). Soil-dwelling insects mentioned included ants (Fig. 19), true cicadas (as well as
Cercopidae), scarabaeoid beetles and field crickets. The Hortus Sanitatis spawned various
other herbals in the 16th Century, but, like the “Bestiaries” that were still popular, these
contributed nothing to the advancement of science, pedobiological or otherwise. In the
“Bestiaries” the religious and moral emphasis became, if anything, even greater and led to the
publication of works virtually devoid of interest in biology, such as the Reductoria Moralia of
Petrus Berchovius, 1521 (see Bodenheimer, 1928, 1929). Soil-inhabiting animals mentioned
therein included the same old range of ants, “ant-lions,” cicadas, crickets (deadly poisonous
insects!) and so forth.16 The Hortus Sanitatis also encouraged more practical books like the
Commentario of Pietro Andrea Mattioli or Matthiolus (1548) and the Naturalis Historiae of
Adam(us) Lonicer(us) (1551). The former, at least in some later editions, includes an
illustration showing earthworms (Fig. 20) and support for the view that the amphisbaena has
only one head, though reference to other soil fauna is virtually lacking. The latter work (see
also Bodenheimer, 1928) mentions ants and scarabaeoid beetles (notably the Rose Chafer,
Cetonia ), crickets, “earth flies” and “earth fleas”. It would be pleasing to think that the last
constituted the earliest specific reference to Collembola (among the most characteristic of all
the soil fauna), but, alas, flea-beetles are more probable, for control of garden pests was being
discussed.

Between the dates of publication of these two works (which maintain a Mediaeval quality)
came the first ever that we can really relate specifically to the soil fauna, though even this was
marginal and dealt very largely with vertebrates. This was De Animantibus Subterraneis (Fig.
21), published in Basel by Georg Bauer under the alias of Georgius Agricola (1549). 17
Although generally descriptive of the fauna, the book emphasizes animals that dig or tunnel in
the ground, some of which (such as rabbits and foxes) present-day soil zoologists scarcely
consider as constituting part of the true subterranean fauna. Little in respect of the latter is
actually discussed. The book was innovative in that it adopted an interdisciplinary, ecological
approach, though to-day most would probably consider it (even though it be in Latin!) to be
largely “waffle” (a feature by no means alien to many ecological writings!)

Agricola {op. cit.) divided his “subterranean” fauna into two main categories, “permanent”
and “occasional,” but this division did not apply to the entire life-cycles of the animals
considered. For example, ants, wasps (Vespula), hornets (Vespa) and crickets, as well as
scarabaeoid larvae, were all considered to belong to the category whose association with the
underground was “permanent”, whilst such insects as bees and cockroaches were but
“occasional”. Table I indicates those animals mentioned by him that (with the exception of
Blattae and “ Gryllus domesticus ”) we might consider to have rather more than a merely
temporary association with the soil or litter. Woodlice (“ Asellus ”) were, however, only
mentioned as hiding in cracks in walls and in houses (“ rimis parietum & domorum latebris
occulantus”) . The “ scolopendrae ” (centipedes and/or millipedes) were said to be found in
[fallen] tree-trunks or in wood placed upon the ground or in sticks loose in the earth

TABLE I

Alphabetical list of German Subterranean Animals that can Conceivably be Classed as Soil Fauna, as Categorized by Georg Bauer (Agricola

1549)

Soil zoology

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Soil zoology

397

(“ scolopendra in truncis arborum, aut in lignis supra terram locatis, aut inpalis terrae
infixis”). “Black” spiders (“ Aranei nigri”), which I take to mean wolf spiders (Lycosidae),
inhabit holes in the ground, as do field crickets (“ Gryllus agrestis ”). It was observed that the
latter (and cockchafers or “ Vermis in Maio”) dig in dry earth in order to construct their
burrows for the summer (House crickets for the winter also); field crickets die before winter;
cockchafers in early autumn. It is also of interest to note that even in this scientifically-based
work, credence is given to the existence of various subterranean demons. The (unknown)
amphisbaena is also mentioned, presumably still in its dread, mythical Mediaeval form (see
Note 6). Though Agricola’s small book, as the first treatise on soil zoology, may be said to
constitute something of a landmark in biological science, it had no impact on the study of soil
animals, or of ecology generally, either at the time or subsequently. It has, in fact, rarely been
cited.

De Differentiis Animalium, by the Oxford physician Edward Wotton (1552), should now be
briefly mentioned for its refreshing style. It was, however, but a concise account of classical,
zoological knowledge freed from the clutter of Mediaeval embellishment, and cannot be said to
have contributed anything new. The soil-inhabiting animals were virtually the same as those
mentioned by Aristoteles. Nigidius’ account of the burrowing and capture of crickets (see p.
382) as well as a reiteration of their alleged medicinal properties, however, is given.

About the same time, the Swedish Archbishop Olaf Ster (Olaus Magnus Gothus) published
his treatise on Scandinavia, including an account of its animals (Ster, 1555: Book XII). In
compelete contrast to Wotton, this was in the old, almost Mediaeval, tradition. Of
soil-inhabiting animals, only ants were considered, though a fair range of their types of “nests”
was covered. It was stated that a red (poisonous) species lived in mole-hills in meadows.
Bodenheimer (1928), who refers to the above, also notes that Hieronymus Cardanus (Geronimo
Cardano), in 1559, cited Albert von Bollstadt (see p. 387) on the question of true ant-lions, and
stated that West Indian ants inflict painful bites.

Bodenheimer (1928) also mentioned the fact that Johann Colerus, in his “Household Book”
of about 1500, referred to damage to plant roots by insects (for which remedies were
prescribed), and he also draws upon the behaviour of ants in relation to weather prediction.
Bodenheimer likewise draws attention to the local, but notably original observations of Dr.

T A L T Jt I S E T T o .

|Fig. 22. Mole-cricket, Talpa Insetto (now Gryllotalpa) from Ferrante Impcrato (1599).

\2uaest. Ent., 1985, 21 (4)

398

Kevan

6 $6 Vlyfsis Aldrouandi

Bicipitem veroefle Scolopendram verum non eft, fed bicipitcpoeta dixit, quod tails videa E
tunnam vt Ariftoteles reftatur & experimento quotidie deprehendi poteft,ScoIopendra ex
vtraq; parte graditur,ranqua vtrimq; caput habeat vnum,eriam ft induas partes dii'iccia fit,
tunc enim altera pars in caudam, altera mouetur in caput. Scd dum graditur, remorum in-
• nb'-'h'ifL ftar,latos mouet pedes, quos alarum inmodu geftar.ficuri poeta dixit. c ThccphraftasSco-
phnt.'c. u. lopendrasfcribitreperiricircaradices giadioIi,quia,inquir,corTgreganturfacillimeincaai.

- Quibufdam in regionibus rantacopia incrcuere,vt fugati ab eis populi proprios dcferuerinc
{Li.Z.c.iv. lares,quod TrerienfibuscontigiflefcribitTheophraftus teftefPlinio: idemde Rhytienft-
bus prodidit E ^EJianus . Qua? in hac tabula nurru i .depingitur Scolopendra eft marina, la-
tocorpore,fubcaftaneo>velurpedibusinnumeris,longiuIculis,aureicoloris. Num.a.co

lift1' 1 u* n t&f

lore

Fig. 23. A page from De Animalibus Insectis by Ulysse Aldrovandi (1602), showing various forms of Scolopendrae
(centipedes), including millipedes and marine polychaet worms (I). [Original in Lyman Collection, McGill Univeristy,
Macdonald College Campus.]

Soil zoology

399

: JLib.z. minirnorumanimalium T beatrum. |pp

Maxima terrefhis Scolopendra,ea quam vidcs craffitie & longitudrne efiq
color rorius corporis cx fufconigricantcfplendens. Singulis inciiuris pedun-
culus appendetluteus,/^ q^infnguJislateribus fcxaginta prorfum & rctror-
, fum a qua facilitate pi omovct. Num et caput vcrlus ingreditur, & in cau-
dam 3 ideoque a Nicandro &Rhodigino biceps dicitur. Partem inter ca-
put & aluum non fimplieem fed multiplicem habet : quo fit,ut pracifum hoc
genus vivere poffit. Irritatus hie Scolopendraramacritermordet, utLndo-
uicus Atmarus(qui nobis eum e Libya dono dcdir)quamvis chirothecis du-
pliciquc Jinteo munitus, vix eum manum ptrcntevn ferre potueritjaltecnim in
linteumosforcipatum adegerat^ diuque pendulus vix tandem excuti per-
mi hr.

Horum alium ex nova Hifpaniola allatum linea qua?dam flammea medium
perdorfum ornatj atqueameus later pilorumque color commendat: habet
cnim capillares pedes, atque armatimfe rollens celerrimectirrit. Hoc fumma
admiratione dignum eft,quum natura huic animalculo caput minimum dede-
rit, memoriam tam'en, vimque rationis amulam,nequecongio, nec urceo,
fed ampliffima quadam mcnfuratribui(Te:cumenim innumeri adfinf ptcles
quafi remiges,& acapite vcluticlavo alij permulrum diftant; novit'tamen
quifqueofficium fuum, & pro imperantis capitis mandato. in hanc vel illam
partem fc conferunt.

Alius item ad nosab Auguftini promontorio ex India perlatus, corpore
nennihil atque pedibus major, quifeptuaginta livefcentibus incifuris, & bis
totidem fpadiccis pedibus conftabat.

PluresScolopendrasrcperirinondubito, omnium fere colorum, prater
viridem: quamvisetiam Ardoynus de viridi mentionrm facit. Infitafin-
gulis proprietas(ex TheophraPi fenrentia)ad Gladioli herba’ radices fefc
conferre. Bubulasautemexuvias meretur Robertus ConPaniinus, eumque
fecutus Stcphanus, nec non Ardoynus ipfe : qui Scoiopendram. primum
ferpentern, deinde odtipedem, turn in cauda cornigerum,.. ultimo urdigra-
dum efte comminifcuntur. Taxandi Rhodoginus, Alberms & 'Avic/enna,
quod nullum Infc&um fupra viginti pedes habere temene affirmant, dlique
numcro Scoiopendram alligant. Quamvis etiam Nicandio-biceps dttrarur
hisverfibus:

Fig. 24. A page of “Scolopendrae” (myriapods) from Insectorum Theatrum , completed by Thomas Moffet in 1589
.(Moufet, 1634, posthumous). [Original in Lyman Collection, McGill University, Macdonald College Campus.)

Quaest. Ent., 1985, 21 (4)

400

Kevan

Johann Bauhin (1598) on “ Scarabaeus majatis ” (a geotrupid dor-beetle, illustrated), and what
were d-riy scarabaeoid larvae, made when graves were being dug near Mompe.gard
(Wurtemberg) About this time, too, (? 1593), the Neapolitan physician Ferrante Imperato
(1599) included, in his book of natural history curiosities, a reasonably good illustrated acc°“
of the insetw" (i.e., the mole cricket, Gryllotalpa, Fig. 22). The closing years of the 16th

Century also saw the invention, in 1590, of the microscope by the Dutchman Zachar,as
Jannsen which, if one may be permitted to turn a phrase “opened up a whole new can o
worms- for all of the biological sciences, and ultimately led to the discovery of the true nature

pharmacopoeia was restructured along biological rather
than pha m cological lines, to produce the Pen-tsao Kang-mu of Lia Shih-Chen, completed ,n
596 (Kon" hi and Ito, 1973). It referred to ants of three or four kinds, termites, burrowing
(polyphagid) and other cockroaches, earwigs, mole crickets, field crickets, ant-lions and vanous
ldnds of beetles such as Carabidae, Elateridae, Silphidae and Scarabaeoidea (including
coprophagous species) and their larvae. Instructions were given on how to collect many of these
insects In the Orient, however, general knowledge of the soil-associated (and ot er) au
made virtually no advance since “Mediaeval" times and, indeed, never did so until caught up m

the wake of 20th-Century western scientific advances.

^ The^adventof the 17th Century may be said to have heralded a new era of invertebra e
zoology and of entomoogy (sensu lata) in particular. This has a direct bearing on the
recognition of the more prominent members of the soil fauna. It did not, however, bring with

entomology ever written both real, belong to > the 16th

Century as they were too early to take advantage of the has P rioX of

Italian, Ulysse Aldrovandi (1602), De Animalibus Insect ,s (F g- 23 ). has pnorfy
publication but the first to be “ready for press” (virtually completed 3.III.1589) was he
Insectorum Theatrum (Fig. 24), compiled by Thomas Moffet, Moufet or Muffet the
London-born, much travelled physician of Scottish parentage who became P .ys^ctan at th
English court The work itself, though not published until long after its compilers dea

(Moufet, .634), was known to some author* Mot own
abortive imperfect continental edition in 1598 (Raven, lz )• . ,

observations with previously unpublished, posthumous manuscripts of EdwarJ "" (“ ^
the Swiss zoologist Conrad Gesner, and particularly of the , lustrator an ^ ollaborato^ - a other
English physician, Thomas Penny (no, Penn, as given by Bodenhe.mer, 1928, and B, r W
Th! English translation (under the name of Muffet) did not appear . ? S * * 8 J ^

quarter-century had passed, when it was combined in a «.ngl Z ^

Reverend Edward Topsell’s (1607, 1608) “Histories of Four-footed Beasts

S BmhAldrovandi and Moffet included invertebrates other than ^ terrestrial arthr°P°da'
case of the latter author, these were restricted mainly to various kinds of worms' ' hef ™,s
dealt in addition with slugs and, quite extensively, with echinoderms. As much o AWrovandJ
tome, in contrast to the smaller volume of Moffet, was taken up w, h all manner
non-scientific (though often fascinating) material, the latter ^ ™

scientifically the more satisfactory, though it, too, left much to betdfesiredt, ftrthwmms am

fairly wide range of soil- and litter-inhabiting animals, whic , apar ^om Although tht

slugs, are accounted for along with others by Bodenheimer (1928, 1929). Althoug

Soil zoology

401

publication of these works constituted a great stride forward for entomology, knowledge of the
biology of the animals concerned was, nevertheless, advanced very little. Moffet (Moufet,
1634), among soil-associated animals mentioned (and often figured) the following, in addition
to earthworms: ants (which are categorized in an almost Mediaeval fashion), mole crickets
(said to spend most of their lives in damp soil and to collect grains of wheat and oats, possibly
for the winter), earwigs, various kinds of beetles (such as scarabaeoids, including their dung
balls, ground-beetles, elaterids and staphylinids, as well as the subterranean larvae of some of
these, such as wireworms and whitegrubs), field crickets and cicadas (whose soil-inhabiting
nymphs were not emphasized). In addition to these insects there was mention of mites on
geotrupid beetles (see also Oudemans, 1926) and of millepedes, centipedes, isopods and
burrowing spiders, as well as scorpions. Aldrovandi (1602), besides discussing earthworms and
slugs, covered much the same range of soil-associated arthropods as Moffet. He noted that
chafers ( Melolontha ) dig “nests” in dry earth, assuming that, as with burrowing bees and
digger wasps, they oviposit there. He also referred to and figured earth nests of Geotrupidae
and mentioned root-feeding by whitegrubs and mole crickets. When writing on cicadas, he
mentioned the amphisbaena in connection with Nikandros (see p. 376), but he dealt with it
more fully, and illustrated it, in his posthumous, 1606, volume on reptiles and serpents,
maintaining that the animal did indeed have two heads, contrary to the declarations of other
authors (Druce, 1910).

In the same year that Aldrovandi (1602) published his De Animalibus Insectis, there
appeared in Strassburg the anonymous New Feld- und Ackerbau, a revised and “modernized”
version of Petrus de Crescenzi’s Ruralium Commodorum of the early 14th Century (see p.
387), advising on how to deal with various pests, including “earth lice”, whitegrubs and ants
(Bodenheimer, 1928). In the following year, Schwenckfeld (1603), in the 6th book of his
Theriotropheum, relating to the fauna of Silesia, dealt with insects in an alphabetical, but very
comprehensive manner (cf. Bodenheimer, 1928). Soil-associated animals mentioned included
the following: “ Ascarides terrenae" (various insect larvae including whitegrubs, cutworms, and
probably the maggots of bibionid Diptera, all of which damaged fields and cut off roots);
“ Cantharis formicaria latior ” ( Cetonia aurata , the Rose chafer, and its white-grub-like larva
and pupa living commensally in ants’ nests); “ Culices fematarii ” (various small,
manure-inhabiting dipterous flies); “ Curtilla ” (mole cricket, Gryllotalpa gryllotalpa, a pest of
roots which builds nests in the earth and lays yellowish eggs therein19); Formica (ants generally;
life history given; the idea that ants become winged when older is perpetuated); Fullo (here
meaning the Common earwig, Forficula auricularia, lives under tree-bark; the idea of
propensity to creep into human ears perpetuated, and remedies given); Gryllus agrestis (= G.
campestris, Field cricket; digs in dry earth and spends the summer underground in holes);
Scarabaeus pilularius (= Geotrupes stercorarius , dor-beetle; makes big balls of dung, using
; its feet, and lays its little larvae therein to protect them from winter cold); Scarabaeus bufonius
i (= Carabus auratus , a large ground-beetle; lives where toads are plentiful; people believe that
1 they copulate with these; they are likewise poisonous [!]); Spondylis (whitegrubs, Melolontha
I and similar larvae; garden pests which lie in the earth near plant roots which they completey
ij devour; used by anglers as fish-bait).

We have already referred to the Reverend Edward Topsell in connection with the English
| translation of Moffet (see p. 400), but his Historie of Serpents (Topsell, 1608) should perhaps
K receive brief mention here. This work was based mainly on the work of Conrad Gesner (see p.

| 400), but, despite its title, includes some information on invertebrates, virtually all, with the

—

I Quaest. Ent., 1985,21 (4)

402

Kevan

exception, so far as we are concerned here, of his notes on scorpions, taken from a
pre-publication copy of Moffet. He does, however, mention a “discourse of Wormes” by his
contemporary, Dr. Thomas Boreham. This does not now appear to be extant (Raven, 1947),
which is a pity, as it would seem to have been the first treatise on earthworms as such, other
than that attributed to Dame Juliana Barnes (see p. 390).

Before concluding this section, we should perhaps briefly mention Francis Bacon of Verulam
(St. Albans), one of the most noted philosophers of his age, if only to note that, in his
posthumous Sylva Sylvarum of 1627, he presented some observations and researches on insects
(most of which he pronounced to be generated in filth) and earthworms. Despite his erudition,
however, he had nothing to contribute to knowledge of the soil fauna (see Bodenheimer, 1928).
Bodenheimer {op. cit .) also refers to the 1645 Zootomia Democritaea of Marco Aurelio
Severino, which includes some observations on the anatomy of crickets and (?) earwigs.

MID-17TH TO MID-18TH CENTURIES

Although Francis Bacon (above) had introduced new philosophical concepts, it was not until
the 1640’s that we see the beginnings of the “Rise of the Naturalists” (the “Bionomic Era” of
Bodenheimer, 1928). Like other developments, this did not come about suddenly, but one
particular name may be mentioned here, that of Dr. Thomas Browne (later Sir Thomas Brown
- without an “e”!). This worthy English scholar began to raise biology to a scientific level by
questioning “authority” - almost “for the first time since Aristotle”, according to T.H. White
(1954), though John Scot (see p. 384) apparently found himself in disfavour on a similar
account several centuries previously. In his Pseudodoxia Epidemica, which went through
several subsequent editions, Browne (1646) refuted, or at least cast doubts upon, many widely
accepted beliefs (though, paradoxically, he was a firm believer in witchcraft and in the validity
of the Ptolemaic concept of the universe!). Apart from debunking mythical beasts, such as the
basilisk (and the amphisbaena) Browne made a number of sound observations. Among these he
noted that the (soil-associated, adult) earwig, Forficula auricularia, is winged, not apterous, as
generally supposed. He also noted the occurrence (though not specifically in connection with
soil) of the “red-coloured summer spider” or “tainct” (later “taint” or “tant”, presumably a
trombidiid earth-mite, identified by Oudemans (1926) as being Acarus holosericeus, described
and named much later by Linnaeus (1758) and now the type-species of the genus Trombidium.
This is probably the first report of a recognizable, free-living, soil-associated mite.20

Before proceeding further, we should perhaps mention here that knowledge of the tropical
fauna was increasing at this time. Particularly notable were the writings of Georg Marcgraf,
some of which were published posthumously (Marcgraf, 1648; see also Bodenheimer, 1929),
though others have only recently come to light. Apart from mentioning the termitophagous
activities of the South American ant-eater, various kinds of Brazilian insects are referred to.
These included digging scarabaeoid beetles (illustrated with numerous parasitic or phoretic
mites on the pronotum) and the jigger flea {Tunga penetrans ) which affects human feet by way
of the soil - as was known to the early Peruvians (cf. Morge, 1973) and had been known to
Europeans since the early 16th Century {cf. Kevan, 1977).

Such reports of this period really belong to an earlier age, and the same is true of the third
textbook of entomology to be published, that of John Jonston (1653), a much travelled Silesian
physician of Scottish extraction. The book (Fig. 25, 26) was really a combination and
condensation of Aldrovandi (1602), devoid of “non-scientific” matter, and Moffet (Moufet,

Soil zoology

403

Tor

miCGZ. A.ntcijJerL. j\I3rov

Tab

xvn

Fig. 25. Ants and their habitations. The upper part of PI. XVII (opposite p. 1 14) in the third text-book of entomology
(Jonston, 1653); from among the illustrations copied from Aldrovandi (1602). [Original in Lyman Collection, McGill
Univeristy, Macdonald College Campus.]

1634). It added nothing to what these two authors had included about soil invertebrates. To his
volume of four (not three as indicated in his title) “books” on insects, etc., Jonston appended
two more, culled from other authors, embracing serpents and dragons. These latter indicated
that belief in mythical Mediaeval dragons, basilisks, hydras and so on, were still current, if
declining. The “ Amphisboena ” (sic) was, however, no longer a two-headed monster (Fig. 26,
lower), but had taken its place among rational legless, burrowing reptiles, along with the rather
similar “ Scytale ” and “ Caecilia ”.21

It was shortly after the appearance of Jonston’s work that the first free-living nematodes
were discovered by Borel (1656). Although these were vinegar eelworms, Turbatrix aceti , and
not soil-inhabiting, their recognition had very important implications for soil zoology.
Fragments of the history of soil nematology will be found in Overgaard-Nielsen (1949) and, to
a minor extent, in Thorne (1961) and Chitwood and Chitwood (1974).

Quaest. Ent., 1985, 21 (4)

404

Kevan

kfttiw Mo uf

Tab xx\ I l

ScoUjptndne Mouf .

Sect . MfDciina.Ten'esit' .

S( ofojjenArcc Terr. Ahlr . ^

Tab. iv

A mj)i?t sheens G

reuittL,

Fig. 26. Above, “ Asellus ” (woodlouse, presumably Armadillidium) and “ Scolopendrae ” (myriapods, including marine
polychaet worms) from lower part of PI. XXIII of Jonston (1653); illustrations taken from Moufet (1634) and Aldrovandi
(1602) as indicated. Below, one-headed “ Amphisboena" from upper part of PI. IV of Jonston’s appended book on serpents.
[Original as Fig. 25.]

Soil zoology

405

The closing years of the sixth decade of the 17th Century also saw further discoveries in the
tropics. Bontius (1658) briefly mentioned a few marginally soil-associated, East Indian
arthropods, such as scorpions, cockroaches, ants, scarabaeoid and other beetles. Rochefort
(1658), besides cockroaches, discussed West Indian termites (mainly in wood) and the already
well-known jigger fleas ( Tunga penetrans). Piso (1658) noted the occurrence of several
root-feeding insect pests of sugar-cane and cassava in Brazil.

Sperling (posthumous, 1661) might now be mentioned, if only for his somewhat novel
approach to zoology in presenting the subject as a sort of catechism in the form of statements,
questions and answers. Many kinds of insects were referred to, but only the clever, industrious,
corn-gathering (!) ants concern us here {cf. Bodenheimer, 1928). Goedart (1662, 1667)
however, made one or two important observations, and he is generally regarded as being the
first naturalist for many centuries to rely mainly on his own observations, rather than on
written “authority” (though this distinction might more properly be claimed by Bauhin, 1598,
above). Part I of his work (1662) gives a good illustrated account of the biology and ecology of
the mole cricket, Gryllotalpa, and its subterranean nest and eggs. (Bodenheimer, 1928, who
discusses the complex authorship of the work, notes that Goedart claims to have invented the
name of the insect.) Part II (1667) contains a very good account of the crane-fly, Tipula
paludosa, and its leatherjacket larva, correctly suggesting a three-year life-cycle; he also
mentions a four-year cycle for the May-beetle ( Melolontha ) with its root-feeding larvae. Part
II is also important from the point of view of soil acarology and nematology as it draws
attention to, and illustrates for the first time, acariform mites and rhabditiform nematodes
(Fig. 27), which are shown in the decaying remains of an ink-cap ( Coprinus ) fungus.22

An increasing number of relevant observations were made by various authors during the
latter part of the 17th Century. Some of these may be briefly commented upon in the form of
the following list:

Hooke (1665): described and gave the first good illustration of a cryptostigmatic mite (Fig.
28) associated with mosses and fungi; Oudemans (1926) identified this member of a dominant
group of soil organisms as “ Acarus ” geniculatus\ Hooke (1665), in addition, reporting again on
the vinegar eelworm Turbatrix aceti (see p. 403), also discovered the nematode Panagrellus
redivivus that occurs in wallpaper paste, an important prelude to the discovery of species
directly associated with soil ( cf . Goedart, 1667, above).

Anonymous (1665): was the first report from North America of cicadas, the holes left by
their emergence from the soil, and their exuviae (Bodenheimer, 1929: 159, gives later
references also).

E. King (1667): gave a fairly detailed account of the biology of ants, including the pupal
nature of “ants’-eggs”.

Charleton (1668): made early observations on cryptostigmatid mites on bark {cf.
Oudemans, 1929); he also commented on various insects, including mole crickets and earwigs,
but his information was taken directly from Aldrovandi (1602) and Moffet (Moufet, 1634).

Redi (1668): did not make much direct contribution to knowledge of soil fauna, but
exploded the myth of spontaneous generation of insects, etc., from “filth” and other substrates,
including soil; he also referred to phoretic mites on ants, both winged and wingless, and on
beetles (see Oudemans, 1926; Bodenheimer, 1929).

Swammerdam (1669): amongst general observations, noted that certain invertebrates
developed without metamorphosis, namely, spiders and mites (probably not soil forms),
scorpions, isopods, myriapods, earthworms and slugs. It may also be noted here that, in the

Quaest. Ent., 1985, 21 (4)

406

Kevan

Fig. 27. Acariform mites and rhabitiform nematodes in the decaying remains of an ink-cap fungus, illustrated by Goedart (1667). Fig. 28. The “Wandering mite” illustrated by Flooke
(1665); the first good illustration of cryptostigmatid mite, see p. 403. Fig. 29. Adults, eggs and subterranean larvae of the horned scarab “ Nasicornis ,” from the upper part of PI. XXV11 of
Jan Swammerdam’s posthumous Bybel der Natuure (1737-38), completed by 1670. [Original in Lyman Collection, McGill University, Macdonald College Campus.]

Soil zoology

407

52-years-posthumous Bybel der Natuure (Swammerdam, 1737-38), detailed studies on ants
and their nests, and illustrations of the horned scarab beetle and its subterranean larva (Fig.
20) were published.

Wray (1670; i.e., the botanist John Ray before he changed the spelling of his name):
experimented with formic acid obtained from ants. It may also be noted here that, in 1672,
Francis Willughby, who was responsible for most of Wray’s later, posthumous, entomological
publication (Ray, 1710), died.

Kircher (1675, 1680): attempted to relate “science” with the animals of the Holy Bible and
to dismiss Redi’s (1668) work (above), maintaining that Noah’s Ark could not possibly have
accommodated representatives of all known living creatures, so that spontaneous generation
must be accepted for many; insects, etc., arose from dead material in the proper proportions; six
classes of such animals existed; from soil came earthworms and slugs, etc., and from excrement
and cadavers emerged scarab (and other) beetles (as well as wasps and bees); another group
included ants and crustaceans (which would include isopods). [An earlier work of Kircher (who
was a Jesuit priest), dating from 1665 and entitled Mundus Subterraneus ... (in 12 “books”),
and an even earlier one of 1657, with the same words in the title, sound like hopeful sources for
the historically inclined soil biologist, but they are basically theological!]

Holger Jacobensen [1676]: as indicated briefly by Petit and Theodorides (1962: 338) made
an important study of the anatomy of the mole cricket ( Gryllotalpa ), presumably in Acta
medica Hafniensis, but the work is unknown to me and unlisted in the principal entomological
bibliographic sources.

Lister (1678): referred to the red trombidiid earth mite, called “tant” {cf. Browne, 1646, see
p. 402), identified by Oudemans (1926) as “ Acarus ” (now Trombidium ) holosericeus, though
not actually in soil.

Wagner (1680): noted that the cockchafer ( Melolontha ) larva (whitegrub) was called
“ Inger ” or “ Enger ” (currently Engerling ) in German because it curled around roots, no kind of
which remained undamaged by them; significantly a three-year subterranean developmental
period was said to be required in Switzerland cf. a total life-span of four years in the
Netherlands, indicated by Goedart, p. 405). The ridiculous practice of excommunicating the
beetles, as at Lausanne, in earlier days was also noted ( cf . p. 389 and Fig. 16).

Claude Perrault [1680]: in Les Mecaniques des Animaux , described and discussed the
alimentary canal of the mole cricket ( Gryllotalpa ), according to Petit and Theodorides (1962:
332). He also published a small tract on Melolontha {cf. Bodenheimer, 1929: 307). Neither of
these works is known to me, nor are they listed in the principal entomological bibliographic
sources.

Knox (1681): was for many years a prisoner in Sri Lanka; his entomological observations
included pertinent comments on ants of various kinds (some of which excavated large holes in
the soil) and particularly on termites, their activities, depredations and mounds; his writings
seem to have been ignored in virtually all major termitological literature, but Bodenheimer
I (1929) quotes him from a German translation of 1689.

Mentzel and Ihle (1683): recorded phoretic mites, identified by Oudemans (1926, 1929) as
“ Acarus ” (now Parasitus) coleoptratorum , on geotrupid beetles.

Muralto (1683, 1684): discussed and illustrated the anatomy of the Wood cricket Nemobius
sylvestris, not a burrower), the Common earwig {Forficula auricularia ) and, more notably, the
mole cricket {Gryllotalpa).

Quaest. Ent., 1985,21 (4)

408

Kevan

Fig. 30. The first (?) recognizable illustration of Collembola ( Hypogastrura on snow) by Spielenberger (1684). After
Bodenheimer (1928). Fig. 31. Early illustrations of mites from Blankaart (1688). H, a species phoretic (?) on soil and
other insects, Parasitus coleoptratorum (Mesostigmata, Parasitidae), called “Luis van een vliegned torretje” or “luis van
de gekokerde vlieg”; I, Scarlet earth-mite, Trombidium holosericeum (Prostigmata, Trombidiidae), called
“Schaarlaken-roode Aard-spinneken” or “Scharlaken-Aard-spin.” Fig. 32. Phoretic (?) mite, Parasitus coleoptratorum,
from a geotrupid dung-beetle, illustrated by James Wilson (1702).

Spielenberger (1684): published the first (?) recognizable illustration of Collembola (a
species of Hypogastrura - Fig. 30) albeit from the surface of snow and not from soil (where
Collembola constitute one of the most numerous groups of animals). Insects have been reported
on snow since, at least, the times of Aristoteles and Plinius, but here their collembolan nature is
undoubted(c/. Bodenheimer, 1929).

Griendel (1687): gave detailed figures of wingless ants and with their pupal “eggs”.

Blankaart (1688): reported on various mites (Fig. 31), including those occurring on burying
beetles ( Necrophorus ) and a figure of the red trombidiid earth mite, identified by Oudemans

Soil zoology

409

(1926) as “ Acarus ” (now Trombidium ) holosericeus, though again above ground.

Mentzel (1688): illustrated the nymphal stage and exuviae of cicadas (cf. Bodenheimer,
1929).

The anonymous author, referred to by Bodenheimer (1928) as Hohaus ( ca . 1690), gave
details of the biology and damage caused by mole crickets ( Gryllotalpa ).

Kampfer [? ca. 1693] (1727-28): recorded observations on termites and ants in the Far East
(see Bodenheimer, 1929, who quotes a German version of 1749).

Leeuwenhoek (1695): was most famous for his development of the microscope, with all that
that implied for the future study of the soil fauna, but he did not contribute significantly to
such studies himself. We may, however, mention his notable, detailed, illustrated account of the
biology and ecology of the crane-fly, Tipula paludosa, and its root-feeding leatherjacket larva
(cf. Bodenheimer, 1928). In this, he “correctly recognized the limiting circumstances of
population dynamics” (Beier, 1973). Leeuwenhoek (1697) also made observations on ants, once
more commenting on the pupal nature of the so-called “ants’-eggs”. Each of these
contributions, however, had been largely anticipated some 30 years previously by Goedart
(1667) and E. King (1667) respectively (see p. 405).

Camerarius (1699/1700): again referred to Collembola on snow (cf. Spielenberger, 1684,
above).

Carrying forward the selected list of “soil fauna” publications into the 18th Century, we
may note the following:

Wilson (1702): gave a good illustration (Fig. 32) of a mite (Parasitus coleoptratorum) from
geotrupid dung-beetles.

Poupart (1704): described, for the first time, the life-history of the unusual rhagionid fly
Vermileo vermileo, the “ant-worm”, whose pit-dwelling larvae live in a similar manner to those
of myrmeleontid Neuroptera (true ant-lions).

Wilhelm Bosman [1704] of the Dutch East India Company resident in Guinea, according to
Bodenheimer (1929), quoting a 1708 German version, refers in his “Voyage to Guinea ... ” to
ants and to termites; the former he believed had a language; the latter were said to build
mounds twice as tall as a man [not an exaggeration], but he did not know if they had a “king”
as big as a [full-grown] fresh-water crayfish, as a Mr. Foquenberg would have it. [The queens
of some Macrotermes species are indeed almost as large as indicated.]

Sloane (1707): published the first of his two volumes on West Indian natural history, but
only the second of these is relevant here, and this did not appear for many years (Sloane, 1725),
see p. 411).

(W)ray [and Willughby] (1710) published (posthumously per Martin Lister) an early
classification system for “insects” (i.e., terrestrial invertebrates) that began a trend towards
orderly taxonomy. What might be termed soil fauna was included in the following categories:

I. “Ametamorphata” (without change)

A. 1, a: legless land animals living in earth - “ Lumbricus ” (all earthworms), slugs.

B, 1, a, x: 6-legged land animals (larger) - probably beetle larvae.

xx, yy: ditto (smaller not holding on to other animals) - including collembola,
“booklice” and some other dubious forms.

2, a: 8-legged, with tail - “ Scorpio ”

b: ditto, without tail - “ Araneus ”, “ Opilio ”, ticks, mites

Quaest. Ent., 1985,21 (4)

410

Kevan

3: 14-legged - “ Asellus ” (isopods and amphipods)

4: 24-legged - ? “bristletails”

5, a: Many-legged land animals - myriapods

II. “Metamorphumena” (making a change)

A. No resting pupal stage - “ Gryllus ”, “ Gryllotalpa ”, “ Cicada ”, “ Forficula ”.

B, 1, a: Moult to pupal stage visible, Coleoptera or Vaginipennia - “Scar abacus" (= all

beetles except staphylinids)

2: Moult to pupal stage concealed - “ Muscae ” (higher Diptera)

This work also included reference to mites infesting ground beetles and Lister’s appended De
Scarabaeis Britannicis including Scarabaeoidea, Carabidae, Elateridae and Staphylinidae) and
classification of British “insects”.

Reaumur (1713 ?): recorded Parasitus mites on geotrupid dung beetles (and on
bumblebees, etc.) - cf. Oudemans (1926).

Vallisnieri (1713): had a classification system in which his third major group of “insects”
comprised those that lived in the earth and in hard substances, but this was not adopted in his
later, major work of 1773 (see Bodenheimer, 1928).

Gunther (1718, 1719): in a sort of quarterly almanack, referred to cockchafers
(Melolontha, mainly swarming adults), mole cricket ( Gryllotalpa , illustrated as having a
curious proboscis), and cutworms ( Agrotis and other noctuid caterpillars damaging vegetable
roots, illustrated). A little later (Gunther, 1723), he again refers to cutworms attacking roots,
and to ants {cf. Bodenheimer, 1928, 1929).

Kolbe (1719): included reference to termites in South Africa.

Frisch (1720, 1722, 1727, 1736): in parts of a serially-published work on mainly economic
aspects of entomology in Germany, made, near the beginning, observations on (burrowing)
field crickets {Gryllus campestris), recommending their use in biological control of House
crickets {Acheta domesticus ); these, he suggested, would be driven out by their more aggressive
cousins. Although this was not a practical proposition, he properly stressed that control of pests
was not possible without adequate knowledge of their biology, the biological control of crickets
being an example of this. Among the few soil pests considered by him was (1727) the crane-fly
Tipula paludosa. The subterranean larvae of the Rose chafer, Cetonia aurata, were mentioned
in the 12th part of the work (1736). Frisch (1772) also records various uropodid and gamasid
mites on beetles (including Geotrupidae) in dung (see Oudemans, 1926).

Sloane (1725): in the delayed second volume of his work (see p. 409), discussed various
insects associated with soil in the West Indies (mainly Jamaica), notably ants, termites,
rootgrubs (scarabaeoid larvae) and the jigger flea (see also Kevan, 1977).

Linnaeus (1735): published the first, short, but regal-folio edition (Fig. 33) of what
eventually, in a different form, was to revolutionize many aspects of natural history, his
Systema Naturae. In it he distinguished the following animals that one may associate with soil:

I. QUADRUPEDIA: Ferae - Talpa (mole). Glires - Sorex (shrew).

III. AMPHIBIA: SERPENTIA {Corpus apodum ... ) - Anguis (“snakes”, including Caecilia)\

here Linnaeus notes some fabulous monsters (e.g., Dragon and Basilisk, but not

Soil zoology

411

C A R O LI

D O C T O R I S

SYSTEM A

S I V

R E G N A T R I

SYSTE MAT. ICE

CLASSES,

G E N i. R A ,

LI N N I , svec!,

MEDICINA,

NATURE,

A N A T U R JE

PROPOSITA

O R D I N E S ,

0 3 Ell OVA ! Quam amp l a Junt op a a Tua !

Quam ea omnia fapienter fectjli !

Quam plena, ejl terra poffejfione tua 1

PLLn. civ. »4.

L U C D U N I BATA VO RUM,

Apud TH I: O D O RU M HAAK. mdccxxxv.

El Typoo«a*hi*

JO ANNIS WILHELMI «»• GROOT.

I Fig. 33. Title page of Linnaeus' (1735) first (regal-folio) edition of the Systema Naturae.

Quaest. Ent., 1985, 21 (4)

412

Kevan

Amphisbaena).

V. INSECTA: Coleoptera - Forficula (“ Staphylinus ” or cockroach!, and Auricularia or
earwig), Scarabaeus (including Scarabaeus pillularis and Melolontha), Carabus (including
“ Cantharellus " auratus). Hemiptera - Gryllus (only “ Gryllotalpa ” is of present concern),
Formica (ants), Scorpio ( S . terrestris, as opposed to “S'. aquat .” or Nepa\ Linnaeus
obviously did not know a true scorpion and deduced their taxonomic position from old
descriptions). Aptera - Acarus (ticks, mites, etc., including Pediicfulus] Scarabaei [on
beetles] and Scorpio-araneus , pseudoscorpions), Araneus (spiders, etc., including
“ Tarantula ” and also Phalangium ), Oniscus (“ Asellus ” spp., isopods), Scolopendria
(including Scolop. terrestris or centipedes, “ Scolop . marina ” or polychaete worms, and
Julus or millipedes).

VI. VERMES: Reptilia - Lumbricus (including intestinum terrae, Aristoteles’ name in Latin
for earthworms, L. latus and the parasitic nematode, Ascaris ), Umax (slugs).

Subsequent editions before the 10th (Linnaeus, 1758) need not concern us here.23

Reaumur (1738): made references to, and illustrated, various dipterous larvae, including
some living in soil; he gave a rather full account of the life-history of the Narcissus bulb-fly,
Merodon equestris ; he also figured ant-lion larvae. Continuing his entomological Memoires
(Reaumur, 1740), he gave illustrated accounts of crane fly ( Tipula paludosa) and a bibionid (?
Bibio hortulans ) and their subterranean larvae; he also discussed and illustrated “ Cicada ornF
and its nymph. A little later, Reaumur (1742) illustrated the life history of “Formica-leo" the
ant-lion (Fig. 34); he also mentions the well known phoretic mite Parasitus coleoptratorum,
though not on beetles {cf. Oudemans, 1926). Reaumur never published the last four projected
volumes of his work, although his manuscripts are preserved in Paris and part of his sixth
volume, written about 1743-44, and dealing with ants, was published posthumously (Reaumur,
1926). 24

Geer (1740, 1743): for the first time, adequately described and discussed Collembola; he
also gave good illustrations (Fig. 35), though these were not the first for the group, as has been
stated by some (see Spielenberger, 1684, and p. 407). His specimens were found in winter on
tree-bark, but this does not detract from the importance of his contribution on these typical soil
hexapods. He observed their method of springing, their moulting, their exuviae, their eggs, and
the presence of the unique ventral collophore.

Linnaeus (1741): unlike Swammerdam, recognized winged male from winged female ants.
Baker (1743): reported mites on earwigs and geotrupid dung-beetles ( Anoetes polypori and
Parasitus coleoptratorum respectively; cf. Oudemans, 1926; Bodenheimer, 1929).

Needham (1743, 1745): identified the nematode Anguina tritici that causes “ear-cockle”25
in wheat (see also Thorne, 1961). This was the first plant-parasitic nematode to be discovered
and, though the “cockle” galls occur in the ears of pannicles of grasses, they fall to the ground
and the worms pass a significant part of their lives in the soil. The importance for soil zoology
of this discovery, therefore, was considerable.

Linnaeus (1745): reporting on his trip to Oeland and Gothland in 1741, mentioned carabid,
staphylinid and scarabaeoid beetles, ants and “ant-lions” (larvae of Myremeleon). His Fauna
Suecica (Linnaeus, 1746), following the classification of his Systema Naturae (above),
referred, amongst soil-associated animals, to mole crickets ( Gryllotalpa ) as garden pests, and
to various other root-feeding invertebrates, such as slugs. The depredations of root-worms,
presumably the larvae of the swift moth Hepialis humuli, attacking hops were noted (see also
Bodenheimer, 1929). He also noted phoretic mites (such as Parasitus coleoptratorum and

Soil zoology

413

J’l ,i'J- bslfyni • to -dc irt ist~- c£e*r Jnscct&r Ttrm- 6 .

/fuMssa^tt J Ou/f'

Fig. 34. The life history of the uFormica-leo" (Myremeleon formicarius ) from Memoirs pour Servir d iHistoire des
1 Insectes. VI. (Reaumur, 1842: pi. 32). [Original in Lyman Collection, McGill University, Macdonald College Campus.)

Quaest. Ent., 1985,21 (4)

414

Kevan

Fig. 35. Geer’s (1740) early illustrations of Collembola (see p. 412).

Uropodidae) on geotrupid dung beetles and other Coleoptera, and the red earth-miti
{Trombidium holosericeum ) and other mites within the soil (“ habitat in terra") - set
Oudemans (1926). This may, indeed, be the first record of mites actually in soil. Thi
prostigmatid mite, now known as Achiptera coleoptratus was recorded from under stones (“swi
lapidibus ”).

Gould (1747): in his account of English ants, gave information on the biology of severa
species; he also stated that ants are eaten by mole crickets (as well as by other enemies) am
noted that millipedes and earwigs, in particular, are among the commensals inhabiting ants
nests.

Baker (1747): gave an account of the damage to pastures in eastern England by whitegrubi
and cutworms.

Rosel (1749): included fine engravings and accounts of the mole cricket and field crickets.

Hughes (1750): referred to field crickets ( Gryllus assimilis ) under stones and lumps o
earth in Barbados; he also discussed ants and termites there.

Hill (1752): recorded red earth mites, presumably Trombidium holosericeum , as beinj
“very common under surface of earth”, as was apparently the “grey rough earth acarus,” {
cryptostigmatid mite determined by Oudemans (1926) as “ Acarus scaber" possibly a species o
Cepheus, and “the little black Acarus,” Pergamassus crassipes (cf. Oudemans, 1929).

Geer (1752): gave another account of the rhagionid dipteran Vermileo, the “ant- worm fly'
referred to much earlier by Poupart (1704), see p. 409.

Reaumur (1753): gave yet another illustrated account of the same insect.

Rosel (1755): described true ant-lions (Myremeleontidae) and illustrated his account witl
engravings very similar to those of Reaumur (1742, see p. 411), but of even higher quality
Rosel died in 1759, but some time prior to that he had prepared material for his fourth volum

Soil zoology

415

(Rosel, 1761) which showed phoretic mites on Necrophorus burying beetles {cf. Oudemans,
1926).

Kalm (1756a): gave a very comprehensive account of the “17-year” cicada (as
‘Gras-Hoppor”) in eastern North America. The same author (Kalm, 1756b, 1761) again
referred to the same insect and to field crickets, Gryllus “ niger ... ” (mostly G. pennsylvanicus,
sometimes G. veletis ), the latter overwintering in the soil, on one occasion at a depth of “ten
nches” (not piled up to that depth on the surface as mistranslations imply). Kalm is one of the
jarliest authors to state that he actually dug for invertebrates in the soil! Among the
overwintering insects that he found beneath the surface were various kinds of ants, carabid and
scarabaeoid beetles and their larvae (including June-beetles and whitegrubs, geotrupids and
lorned scarabs). Native, litter-dwelling (as well as imported, domiciliary) cockroaches are also
nentioned. Kalm’s so-called woodlice, however, were not isopods but ticks.

Osbeck (1757): commented upon several kinds of “ Scarabaeus ” beetles and the substrates in
vhich they occurred in Spain. Reference is also made to Spanish field crickets (see also
Bodenheimer, 1929).

Adanson (1757): observed various insects during his sojourn in Senegal, and among these
vere termites {cf. Bodenheimer, 1929). Apart from noting their destructiveness, describing
Tuitless efforts to combat their ravages, suggesting arsenic and fire for the purpose, he also
nade observations on the internal structure of termitaria. He was astute enough to conclude
hat the majority of the termites that were most destructive to his possessions were not of a kind
hat build conspicuous mounds; he noted their covered galleries. Reading Adanson, one gets the
mpression, perhaps for the first time, that here was an author who had some appreciation of
he intricate association between the soil and its (termite) fauna.

We have now reached that point in zoological history, when Linnaeus (1758) published the
l Oth edition of his Sy sterna Naturae. There was, indeed, no momentous biological discovery
issociated with this event, but the almost universal adoption of binominal nomenclature for all
inimals, which followed within a remarkably short time, ushered in a new era. So far as the soil
auna was concerned, the only immediate impact was to add a few more generic names to those
isted in earlier editions of the work (and eventually to provide a reference point for validating
he various names in the future). Additional, marginally soil-associated genera of Coleoptera
ncluded such as Mister , Silpha, Elater and Staphylinus (in its current sense, no longer a
:ockroach). We also find additions such as Termes (now specifically meaning termites) and the
mrrowing Field cricket, Gryllus campestris.

To conclude this section, we might just mention one, rather quaint but relevant agricultural
vork that really belongs to an earlier period, though published in the following decade. This is
he poem, written in the early 1760’s on St. Kitts, West Indies, entitled The Sugar Cane
Grainger, 1764). Kevan (1977) has extracted and commented on all the numerous
nvertebrate animals mentioned in it. These included a number of soil-related forms, such as
mrrowing land-crabs {Cardiosoma), scale insects on cane roots, ants, termites, crickets,
:ockroaches, jigger fleas and the human hookworm among the parasitic nematodes.

UP TO THE MIDDLE OF THE 19TH CENTURY

After the publication of the “Tenth Edition” of Linnaeus (1758), a new, if not universal,
•rderliness came about the field of zoology. While anatomical and biological studies increased,

L was, nevertheless, the discovery and description of previously unknown, or unrecognized.

}uaest. Ent., 1985, 21 (4)

416

Kevan

creatures that occupied most attention and which went ahead by leaps and bounds. The
“pre-Darwinian” systematic era had begun! Author after author added more and more species
to the known fauna of the world, and, though the proportion was not large, many of the animals
involved were associated with the soil and litter, if not as adults, then in their immature stages.

Clearly it would be profitless to attempt to enumerate in detailed succession the new
discoveries in the manner I have adopted hitherto in this review. We must confine our attention
to the more significant events for soil zoology and to the relevant publications. For the general
trends of the period, with many specific examples from entomology (though few mention the
soil fauna even indirectly), the reader is referred to Tuxen (1973), who begins a little earlier,
and to Lindroth (1973), who ends a little later than does this section. For an exhaustive account
of developments in acarology from 1759-1804, see Oudemans (1929). A full bibliography of
the literature on oligochaet annelids (including terrestrial forms) to 1894, is given by Beddard
(1895); Stephenson (1930) gives no historical review for these creatures; Reynolds and Cook
(1976) limit their brief remarks to taxonomy.

Before proceeding further, however, we should remember that there were, of course, others
than Linnaeus himself who crossed the “nomenclature boundary”, and amongst these, perhaps
the most important was his compatriot Karl De Geer, whose Memoires pour servir h Histoire
des Insects (so called in deference to the works of Reaumur, p. 411, and published in
Stockholm) began in 1752, before the “Tenth Edition”. The final volume, the seventh, however,
was not published until the year of its author’s death, 1778. It was in this volume that Geer’s
early observations on Collembola (see p. 413) were reprinted. The series did, of course, contain
many references to various other soil- and litter-associated insects, etc., but we shall not
enumerate these, other than, perhaps, to mention “ Acarus vegetans ”, a uropodid mite on
staphylinid beetles noted in 1768 (Oudemans, 1929).

In this period, we should also refer to the general entomological publications of such other
authors as J.C. Fabricius, P. Rossi, P.-A. Latreille, W. Kirby and W. Spence, and H.C.C.
Burmeister, to mention but a few. The first of these published his most important works from
1775 to 1798, but we shall mention only his Philosophia Entomologica ... (Fabricius, 1778).
This is because that particular work is regarded by some as being the first real textbook of
entomology, dealing as it did, with the subject scientifically and confining itself to non-marine
anthropods. Rossi is included here, not only for his important example of work on localized
faunas, his Mantissa Insectorum ... (Rossi, 1792, 1794), but more because he was the first ever
Professor of Entomology to be so designated (at the University of Pisa, 1801-1804). He did not,
in fact have much direct connection with soil fauna, but the recognition of entomology as a
discipline was to be of major importance to its study. Latreille’s numerous revisions of the
classification of arthropods, mainly between 1802 and 1829, are also of major general
significance, but, from a “soil” point of view, he also devoted much time to the study of ants
(Latreille, 1802).

Kirby and Spence (1815-1826) made an outstanding contribution to entomology by
“popularizing” the subject without degrading it. Nevertheless, they contributed little or nothing
beyond what was already known to the knowledge of the soil fauna as such. Admittedly, in the
first volume (1815 [& 1816]), they considered certain soil-inhabiting species amongst the pests
about which they wrote, but they added virtually nothing new. Similarly, in their second
volume (1817), though they devoted 75 pages to ants and termites, they limited discussion
almost entirely to their biology and behaviour, with little, if any, indication of the possible roles
of these insects as part of the soil fauna.

Soil zoology

417

Towards the close of the period treated in this section, Burmeister (1832) published the first
(general introductory) volume of his influential Handbuch der Entomologie. His final (5th)
volume did not appear until much later, in 1855.

We might now mention a few publications of more particular interest (direct or indirect) for
a study of the soil fauna, that appeared during the period considered here. We may begin by
noting Spallanzani’s (1769) account of free-living nematodes, to which he again referred many
years later, in 1787 {cf. Chitwood and Chitwood, 1974). Two references by O.F. Muller (1773,
1776) are also notable as they were the first to recognize the distinctness of that extremely
important group of soil animals, the enchytraeid oligochaet worms. “ Lumbricus ” (now
Lumbricillus ) lineatus and “L.” minutus (of dubious identity) were the species involved, both
from near the seashore {cf. Reynolds and Cook, 1976, who give a brief history of oligochaet
research generally). Also concerned with “worms,” once more with the “ear-cockle” nematode
of wheat, Anguina tritici, we may also mention Roffredi (1775) and Scopoli (1777), who,
respectively, began to unravel the life-history, and named the genus, though not the species.
[The latter was not done until Steinbuch (1799) worked mainly on a related species, A. agrostis
- see Thorne (1961).]

In the meantime, Schrank (1776, repeated 1781) was writing about mesostigmatid mites,
such as Pergamassus crassipes, and Collembola, like Onychiura ambulans , in soil under
flower-pots, and mites like Hologamasus lichenis under lichens (see Oudemans, 1929); and
O.F. Muller (1786 - cf Chitwood and Chitwood, 1974) made the first observations on truly
free-living fresh-water nematodes (many of which may occur in the water-film around soil
particles).

Another group of predominantly soil-inhabiting animals that were written about quite
extensively by European travellers to the tropics were termites. Notable among such authors
were Konig (1779) in respect of southern India and Sri Lanka (Fig. 36), and Smeathman
(1781) regarding tropical West Africa (Fig. 37). Fletcher (1922) translated Konig’s paper and
commented upon that of Smeathman; Thakur (1984) briefly notes that Konig was probably the
first author to investigate termites scientifically in Peninsular India and Sri Lanka (though
there had been much earlier reports from the latter by Knox, p. 407), and he clarifies the
nomenclature. Fungus-gardens, ectoparasites, the use of termites as human food, etc., are all
mentioned. Sparrmann (1784) [1783] also wrote about termites in Africa, but in respect of
South Africa. Among other things he observed their “piercing the soil.”

More significant, perhaps, and published a few years later (though mostly written earlier)
came the first edition of The Natural History and Antiquities of Selborne (G. White, 1789). In
this, White (see also Note 26, p. 440) not only made keen observations on mole crickets (Fig.
38), field crickets, harmful scarabaeoid and tipulid larvae, other insects and injurious slugs, but
stated that “worms seem to be great promoters of vegetation which would proceed but lamely
without them, by boring, perforating, and loosening the soil, and rendering it pervious to rains
and the fibres of plants, by drawing straws and stalks of leaves and twigs into it; and most of all,
by throwing up such infinite numbers of lumps of earth called worm-casts, which, being their
excrement, is a fine manure for grain and grass ... the earth without worms would soon become
cold, hard-bound, and void of fermentation; and consequently sterile ... ”26 Here, then, we
Finally see the beginnings of a clear understanding of the interaction between the soil and its
inhabitants!

Nevertheless, though the book was an immediate best-seller (and was even published within
three years in a German translation in 1792, the year before White died), this particular piece

Quaest. Ent., 1985, 21 (4)

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Kevan

Fig. 36. Termites from south India and Sri Lanka illustrated by Konig (1779). Nos. 10 and 1 1 are of Hospitalitermes
monoceros (Konig), from Sri Lanka; nos. 12-14 are of Anacanthotermes viarum (Konig), from South India; the others are
of uncertain identity.

Soil zoology

419

Fig. 37. The illustration of the large, mound-building West African termite, Macrotermes bellicosus, from Smeathman
(1781). Smeathman tells us that, in certain “English” parts of West Africa, termites had the dubious distinction of being
dubbed “Bugga Bug”!

Quaest. Ent., 1985,21 (4)

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Kevan

Fig. 38. Mole-cricket ( Gryllotalpa ). Illustration from an early edition of Gilbert White’s (1789) Natural History and
Antiquities of Selborne.

of wisdom was not followed up for almost half a century, except for the plagiarism by Bingley
(1803 and subsequent editions, see Note 26), until Charles Darwin read a paper on the subject
to the Geological Society of London on November 1st, 1837, soon after his return from his
famous voyage with H.M.S. Beagle (Darwin, 1840). — And that did not arouse much
immediate interest either!

Meanwhile, general systematic and biological works like those on ants by Latreille (1802)
and Huber (1810), and on mites by Schrank (1803-04) and Hermann (1804), went on; Morren
(1829) was experimenting with water relations of earthworms; Henle (1837) described the
terrestrial oligochaet genus Enchytraeus (type species E. albidus ) from decaying seaweed,
sewage beds and compost heaps; Bourlet (1839, 1841, 1842) and Nicolet (1841, 1847) had
begun to lay the foundations for the study of important group of soil organisms, the
Collembola; and Koch (1835-38, 1844, 1847) did the same for myriapods (and other non-insect
arthropods). One may note, too, that Dujardin (1842) wrote about Nematomorpha (gordiid
worms) and larger mermithid nematode parasites of insects (his new genus Mermis), which can
be associated with soil; and then later (Dujardin, 1845), while dealing chiefly with
endoparasitic helminths affecting vertebrates, he referred to free-living, soil-inhabiting
rhabditiform nematodes, as well as to the plant parasite, Anguina tritici. Hoffmeister
(1842-45), too, was beginning to distinguish between various species of lumbricid earthworms.

By this time, however, Ehrenberg (1837) had published his tract on the “living soil”,
drawing attention to the possible role of protozoa and other micro-organisms therein, and
Darwin (1840) had pointed the way to the scientific study of earthworms. Thus, soil biology, as
such, may now, perhaps, be said to have begun at last, however modestly. One should not,
however, be misled by the title of a paper by Schiodte (1849), “Specimen faunae subterraneae”,
for this, though an important landmark of its own, dealt with cave-dwelling, not soil
invertebrates.

Soil zoology

421

1850 TO 1900

By the middle of the 19th Century, real knowledge of the soil fauna as such was, in general,
only a little advanced from what it had been in the days of Aristoteles. Larger or more
conspicuous animals that inhabited the soil were reasonably familiar - such as moles, legless
lizards (even true amphisbaenids!) and amphibia, earthworms (though most of these were
simply lumped together as “ Lumbricus ”, certain slugs, isopods and myriapods, and a modest
array of larger insects, such as scarabaeoid and various other beetles and their larvae,
cutworms, cicada nymphs, ant-lion larvae, mole crickets and burrowing field crickets, crane-fly,
bibionid and other fly larvae, and, of course, various kinds of ants and termites. Incidental
knowledge had, however, begun to accumulate regarding smaller creatures, such as
Collembola, mites (especially those that lived on insects), enchytraeid worms and nematodes,
though mostly as little more than curiosities. Beyond the earthworms, and possibly ants and
termites, there was little concept of a soil-fauna community. Other animals were considered
largely in isolation, Ehrenberg’s (1854) atlas of soil inhabiting protozoa, etc., being an
exception.

A noteworthy early contribution to nematology, including free-living forms, was that of
Diesing (1850-51), later revised (Diesing, 1861). Also in relation to nematodes, a note by
Berkeley (1855) was of considerable interest as it focussed attention on an unidentified “vibro”
attacking the roots of cucumbers, probably the first discovery of a plant-parasitic eelworm
other than those causing ear-cockles of cereals and wild grasses ( Anguina ), which had again
received attention shortly before by Hardy (1850). [J. Kuhn’s (1857) Anguillula (now
Ditybachus ) dipsaci on teasel is often considered to be the “second” plant-parasitic nematode.]

Termites were also receiving further attention from a systematic point of view with the first
monograph on the group by Hagen (1855-60), while more information on soil-inhabiting
nematodes continued to accumulate. For example, Schulze (in Carus, 1857) described the
soil-inhabiting Diplogaster micans)\ Gervais and Beneden (1859) gave us more on the
ear-cockle eelworm ( Anguina tritici)', H.J. Carter (1859) mentioned tropical free-living
nematodes while writing on parasites of humans and, economically very importantly, Schacht
(1859) noted the occurrence of the cyst-forming, root eelworm ( Heterodera schachtii, though
not then named27) on sugar-beet. Claus (1862) and Eberth (1863) also contributed to our
knowledge of free-living nematodes, by which time, however there were only about 80 species
known, most of them marine (Overgaard-Nielsen, 1949). Lest it be thought that no progress
was being made at this time in the area of integrated soil biology, one should mention here the
writing of Post (1861-62), who again drew attention to the important role of living organisms
in the soil, but the time was not yet ripe for detailed investigations of this kind. More traditional
work of importance during the immediate period was the initiation of the continuing work of
Schiodte (1861-83) on the larvae of Coleoptera, very many of which live in soil or litter,
decaying vegetation, etc., and which were largely unknown at the time. Koch (1863) also
(posthumously) laid the foundations for a better understanding of the myriapods. (His son
continued in this field later).

A real beginning was also made on a concerted study of free-living nematodes (including
many soil forms) by Bastian (1865). Schneider’s (1866) monograph on nematodes in general
also appeared about the same time, but it was Butschli (1873) who provided the basis for the
present-day classification of free-living nematodes, of which he included 61 soil and fresh-water
species, 30 new (Overgaard-Nielsen, 1949). An interesting discovery was also made about this

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422

Kevan

time, when Lohde (1874) first observed nematode-trapping fungi in the soil. Then came the
first of J. G. de Man’s publications dealing specifically with soil nematodes (Man, 1876), of
which he described about 50, most new (Overgaard-Nielsen, 1949). This author continued to
publish on free-living species until 1921 (Thorne, 1961). Interest in terrestrial annelids,
especially earthworms, was increasing about this time, as indicated by the works of Eisen
(1871-1873), who was to continue with their systematics for many years, and of Perrier (1872,
1874), who also did some experimenting with them.

A milestone belonging to this period, for students of the soil fauna, was the publication of
the Monograph of the Collembola and Thysanura (which included Microcoryphia and
Diplura) by Lubbock (1873).28 Such biological and ecological information as was available was
included, though it was mainly a systematic work, as was customary (but necessary) at the
time. By the end of the decade, Plateau (1876) had studied digestion in myriapods; Hensen
(1877) had published the first important paper on the role of earthworms in soil fertility since
Darwin (1840); Vejdovsky (1877, 1879), in two works, with which I am unfamiliar, began to
put the enchytraeid annelids in order; and P.E. Muller (1879), who invented the terms “humus
form”, “mull” and “mor”, stressed that these latter were biological, not merely
physico-chemical, systems, in which the fauna in general (not merely earthworms), together
with other organisms, was intimately involved (see also P.E. Muller, 1884, 1889).

Darwin (1881), with his customary procrastination, now published The Formation of
Vegetable Mould through the Action of Worms (Fig. 39), which over-shadowed other valuable
but slightly later contributions on the subject by Hensen (1882) and Baur (1883). Vejdovsky
(1885) also made further studies of earthworms and other oligochaets (but not much on
Enchytraeidae as he had already dealt with these, as noted above). It is probable that Darwin’s
book, rather than stimulating further research on the interaction of fauna and soil, tended, by
its authoritativeness, to give the impression that there was little more to be said on the matter -
except where earthworms were rare or absent. Drummond (1887, 1888) developed the
hypothesis that termites were the tropical analogues of earthworms, but soil fauna studies as
such did not burgeon forth as might have been expected.

We should now turn our attention again to other groups of animals that are extremely
abundant in the soil, namely the myriapods and the mites. In respect of the former, Latzel
(1880, 1884) published a very important monograph for central Europe. Knowledge of mites
was gradually accumulating as a result of the efforts of various authors, but one in particular,
Antonio Berlese, should be mentioned. His Acari, etc., in Italia reperti, published over many
years (Berlese, 1882-1903) included large numbers of soil-inhabiting species. Before his major
contributions were made29, however, Michael (1884, 1888) had published an extremely
important monograph on the British “oribatid” (Cryptostigmatid) mites, which laid the
foundation for the study of these typical and abundant soil- and litter-inhabiting creatures.

Towards the close of the 19th Century we should refer to further work on economically
important root-feeding nematodes, for instance, the description of the root-knot eelworm of
coffee Meloidogyne exigua by Goeldi (1887) in Brazil, and a fine monograph on the
Sugar-beet eelworm, Heterodera schachtii, by Strubell (1888). One of the earliest workers to
realize the important role of the fauna in comminution of litter and in humus formation was
Keller (1887). Another was Kostychev (1889), who recognized that passage of organic matter
through the bodies of invertebrates (earthworms, millipedes, sciarid fly maggots), even if little
chemical change occurred, was important, the excrement being more readily broken down by
fungi. Related to this, though scarcely realized at the time, were the studies on the biology of

Soil zoology

423

Tower-like casting from near Nice, constructed of earth, voided
pronahly by a species of Periclueta : of natural size, copied from
a photograph.

A tower-like casting, probably ejected by a species of Perichmta,
from the Botanic Garden, Calcutta : of natural size, engraved
from a photog>aph.

A casting from the Nilgiri Mountains in South India; of
natural size, engraved from a photograph.

Fig. 39. Famous illustrations of earthworm castings published by Charles Darwin (1881); from photographs by Dr. King,
when keeper of the Botanic Gardens, Calcutta.

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424

Kevan

UNTERSUCHUNGEN

UBER DIE

Bodenfauna in den Alpen

INAUGURAL- DISSERTATION

ZUR

ERLANGUNG DER PHILOSOPHISCHEN DOKTORWURDE.

VORGELEGT DER

HOHEN PHILOSOPHISCHEN FAKULTAT DER UNIYERSITAT ZORICH

(MATHEMATISCH-NATURWISSENSCHAFTLICHE SEKTION)

VON

KONRAD DIEM

aus Hkkisau, API'ENZELL a.-kh.

Begutachtet von den Herren Prof. Dr. A. LANG

Prof. Dk. K. KELLER

ST. GALLEN

ZOLLIKOFER’SCHE BUCHDRUCKEREI

1903

... . -

Fig. 40. Title page of Konrad Diem’s (1903) thesis on the Soil Fauna of the Alps. The work was reprinted the same year in
Jahrbuch der Naturwissenschaft lichen Gesellschaft St. Gallen 1 901-1 902: 234 pp.

Soil zoology

425

WATER

JACKE

COLLECTING _
VESSEL

Fig. 41. Berlese funnels for extracting soil arthropods. Left, original pattern of Berlese (1905); right, early gas-operated
modification. After Kevan (1962a).

millipedes by Rath (1890, 1891). Further investigations on the effects of earthworms on soil
fertility were also carried out by Wollny (1890) and Djemil (1896), while various other
earthworms studies were undertaken by Bretscher from 1895 until the end of the century, when
he submitted for publication a paper on their biology, which appeared the following year
(Bretscher, 1901).

The 20th Century opened auspiciously from the point of view of soil zoology, for, under the
direction of Professor Conrad Keller (see above), Konrad Diem undertook a comparative
ecological study, from 1900 to 1902, of the animals inhabiting Swiss alpine soil and litter. In his
doctoral dissertation (Fig. 40) for the University of Zurich (Diem, 1903), he briefly defined the
term “ Bodenfauna ” (soil fauna) for the first time, and it would seem that his thesis was the
very first attempt at an integrated faunistic investigation of soil and litter habitats anywhere.
Though his methodology was quite unsophisticated, he considered not only the more
conspicuous animals (earthworms, myriapods, gastropod molluscs), but also Collembola,
nematodes and enchytraeids as best he could. Additional, less thoroughly treated groups were
beetle larvae, fly larvae and “others”. Mites seem to have been ignored. Indeed he admits to
having difficulties with the smaller forms of life.

Another important step forward at about this time was the invention of the “Berlese funnel”
for extracting mites and other small arthropods from soil and litter (Berlese, 1905). Berlese’s
name is still widely, but erroneously used for virtually all apparatus of a similar nature, but the
original, water-jacketed funnel was heated, eventually by a gas-ring from below (Fig. 41),
whereas all modern devices are unjacketed and heated (by an electric bulb or other device)
from above. These are modified from the “Tullgren funnel” (Tullgren, 1917).

During the early years of the century, general interest in earthworms continued. Parker and
Metcalf (1906) and Hurwitz (1910) were concerned with the reactions of these to salts and to
acids respectively. Russell (1910) and Bauge (1912) again stressed the question of earthworms
and soil fertility. Wieler (1914) also concerned himself with earthworms and soil reaction.

The publication by Russell (1912) of the First edition of Soil Conditions and Plant Growth
“provided an enormous stimulus to the comprehensive study of the soil and its living organisms,
but at that time not much could be included on the role of animals other than earthworms ... ”

1900 TO 1945

Quaest. Ent., 1985, 21 (4)

426

Kevan

(Kevan, 1962a), and there was little enough of that. A sort of “two solitudes”30 attitude on the
part of self-styled soil scientists on the one hand, and of zoologists, on the other, seems to be
traceable to this period. The former, for a long time, seldom paid attention to animals smaller
than earthworms (which they ignored if they could), whilst the few zoologists who deigned to
get their hands dirty were considerably retarded in their recognition of the pedological
significance of the soil fauna. Most early studies by the latter had a direct or indirect bias
towards crop pests, mostly insects or nematodes.

One of the earliest comprehensive studies of insects (and other arthropods) in the soil was
that of Cameron (191 3)31 in which the principal finding was that gravitational soil water was
destructive, and capillary soil water favourable, to them. The paper was not concerned with
effects of the fauna on the soil. On the other hand, Cobb (1915), in a pioneer paper attempting
to popularize “nematology” (the word was introduced into the language here), suggested,
though no basic information was available, that nematodes presumably bore an important
relationship to the fertility and biology of the soil. (He was in error, however, in his opinion that
the great majority of soil species possessed an oral “spear” and were injurious to plant roots).
He estimated that, in an acre of North American alluvial soil, there may be 3,000 million
nematodes in the top 3 inches (7.5cm). Cobb (1917, 1918) gives further estimates of nematode
populations in sand and soil.

While the First World War restricted work on the soil fauna (as it did other endeavours) in
most places, even in neutral countries, we may recall the appearance of Tullgren’s (1917)
funnel for extracting small arthropods (Fig. 42), and note the invention of another type of
funnel for soil-inhabiting nematodes (Fig. 43), that of Baermann (1917). The latter was
originally for retrieving the larvae of parasitic hook-worms from tropical soils, but it was
subsequently used for nematodes generally, including those occurring in the faeces of
vertebrates.

Probably the first Canadian contributions to integrated soil entomology were also published
at this time, those of Cameron (1917a, b), though the larger of these related to work done
previously in England. They stressed the importance of soil moisture and aeration, as well as of
other factors, on soil insect ecology.

Over the next decade or so, no major stride forward was made in the study of the soil fauna,
but we may mention a few publications of interest. Morris (1920) began his investigations on
soil insects, reporting on their occurrence in permanent pastures; Jegen (1920) discussed the
significance of enchytraeid worms in humus formation; Buckle (1921) investigated the fauna of
arable land; Arrhenius (1921), Moore (1922) and Phillips (1923) were concerned with the
effects of soil reaction (pH) on earthworms, whilst Salisbury (1923) looked at the question
from the opposite viewpoint, the influence of earthworms on soil reaction and stratification;
Micoletzky (1922) gave the fullest account to date of free-living soil nematodes; and
McColloch and Hayes (1922) discussed the reciprocal relations between soils and insects. The
year 1922 was also notable for the first serious attempt to use “wet” extraction methods for soil
arthropods (Fig. 44), as introduced by Morris (1922a) for the purpose of his studies on these
animals in arable land (Morris, 1922b, 1927). M. Thompson (1924) also undertook an
extensive study on soil arthropods. Soil nematode investigation, especially on applied aspects
also began to forge ahead. The distinctiveness and great importance of the Potato root eelworm
(later called the Golden nematode in America) came to the fore, the name Heterodera
rostochiensis being bestowed upon it by Wollenweber (1923); see papers by Wolleenweber
(1924) and Morgan (1925). Thorne (1927) investigated mononchid eelworms in arable soils in

Soil zoology

427

SAMPLE

if — GAUZE

-TIN CAN

SUPPORT

OUTER SUPPORTINC
WAXED CUP

(tip cut off)

INNER WAXED
J CUP

Fig. 42. Tullgren funnels for extracting soil arthropods. Left, original design, after Tullgren (1917); inset, right, modified
version of Haarlov (1947), after Kevan (1962a). Fig. 43. Baermann funnels for extracting soil nematodes. Left, basic
pattern of Baermann (1917); right, Anderson and Yanagihara field pattern. After Kevan (1962a).

Quaest. Ent., 1985,21 (4)

428

Kevan

Fig. 44. Morris “wet” extraction apparatus for soil arthropods; after Morris (1922a). a, ledge; b, shelf; c, First funnel; d,
first sieve with holes 3-5 mm in diameter; e, second funnel; f, second sieve, with holes 1-5 mm in diameter; g, third funnel;
h, third sieve, with 50 meshes to the inch; i, outlet; k, inlet; 1, rose. Fig. 45 Ladell flotation apparatus for extracting soil
arthropods; after Ladell (1936). a, cylinder in which soil is mixed with liquid; b, conical head Fitted to the top of the
cylinder, with a watertight connexion; c, combined stirrer and air bubbler supporting two sieves; d, stirring mechanism; e,
air pump; e,, manometer; f, small electric motor for stirring; g, soil sedimentation tank; h, glass reservoir containing the
solution; i, Buchner funnel; j, Filter flasks; 1, discharge outlet of cylinder; 2, handles; 3, rubber ring; 4, central tube of
hollow stirrer; 5, hexagonal box of stirrer; 6, air tubes; 7, air outlet; 16, threaded collar; 17, hexagonal nut; 18, brass tube
connexion; 19, flanged tube; 20, hexagonal back nut; 21, brass boss; 22, iron strap; 24, crank; 25, connecting arm; 27,
chute; 28, lip; 29, overflow pipe; 30, discharge outlet of tank; 31, handles; 32, tap funnel. Fig. 46. Flotation apparatus for
extracting soil arthropods. Above, Salt and Hollick (1944) apparatus (A. sieving; B. actual flotation; C, separation of
arthropods from vegetation). Below, large-scale tanks of Cockbill et al. (1945). Both after Kevan (1962a).

Soil zoology

429

paest. Ent.. 1985, 21 (4)

430

Kevan

the United States, and Cobb (1927) summarized what was known about “nemas” (nematodes
other than parasites of vertebrates) in general. Even in China these animals were receiving
attention in soil (H.D. Brown, 1929). An important paper was published by Handschin (1926)
on the Collembola of subterranean communities. Oudemans (1926, 1929) published the first
two parts of his historical bibliography of mites. A little-known, annotated bibliography of
cryptostigmatid or “moss” mites by Jacot (1929) should also be mentioned, and, so as not to
forget earthworms, we should note an important contribution by Stockli (1929) on the
significance of these in soil formation.

In concluding our brief review of the 1920’s, we should also refer to the comprehensive study
of insects and other invertebrates in the soils of pasture and arable land by Edwards (1929),
and to a characteristically American contribution to soil entomology - a machine (albeit still
hand-operated) to facilitate the rapid recovery of insect larvae (specifically elaterid wireworms)
from the earth (Lane and Shirck, 1928). Power-driven apparatus of increasing complexity was
introduced later, but this will not be considered further here (for references, see Lang, Akesson
and Carlson in Kevan, 1955: 351-355).

The beginning of the next decade was notable for the appearance of one of the classics of soil
zoology, The Fauna of Forest Soils, by C.H. Bornebusch (1930), which had a very great
impact on the study of the role of animals in forest soil ecology. In the same year, too, Mail
(1930) stimulated interest in the effect of low (winter) temperatures on the survival of soil
insects. Driedax (1931) published further investigations on the significance of earthworms for
plant growth, and, in the same year, Nazaroff (1931) advanced the interesting theory, later
generally accepted (Machado, 1983b), that large masses of spongy brown ironstone in lateritic
areas of the African (and other) tropics were a result of “the ferruginization of termitaria”.
Ulrich (1933) made a notable quantitative comparison of the macrofauna of forest litter
between good and poor stands of trees. Kollmansperger (1934) included extensive ecological
information in his thesis on German earthworms; and Rommell (1935) gave a positive example
of the role of myriapods in mull formation.

It was about this time that Jacot (1935, 1936), first in a short, popular article and then by a
scientific paper, began his eloquent, but not very successful, attempts to interest North
American biologists and/or pedologists in the fauna of soil and litter (especially of forest and
woodland) for its own sake and from a pedological viewpoint.

Frenzel’s (1936) monograph on invertebrates of all groups in meadow soils was the next
large work in the field after that of Bornebusch. It appeared in the same year as Ladell (1936)
proposed a new, rather complicated type of flotation apparatus for recovering insects and other
arthropods (Fig. 45). In the following year, Ford (1937) published an important ecological
paper on population fluctuation in Collembola and mites. A significant Canadian contribution
on populations of soil insects was that of K.M. King (1939). Jacot’s (1939) discovery that
phthiracarid mites hollow out conifer needles without apparent external change in their form
was also significant. Jacot (1940) also produced a very commendable, comprehensive review of
animals in soil and litter, but again he seemed unable to influence American biologists or
pedologists to any appreciable extent.

As the Second World War progressed, the need for food production focussed attention on
the soil (except for large areas where this was being fought over). Even papers like that of
Joachim and Kandiah (1940), comparing soils derived from termite mounds with those of
adjacent land, took on added significance, as did Adamson’s (1943) review of termites and soil
fertility. Even the mole ( Talpa europaea) and its relationship with earthworms and other soil

Soil zoology

431

invertebrates received its share of attention (MacDougall, 1942). From a pedological viewpoint,
the microscopic investigations of humus by Kubiena (1943) showed the abundance and
importance of the excrement of soil invertebrates for further decomposition of soil organic
matter, Gisin (1943), in neutral Switzerland, published the results of an extensive study of the
ecology of Collembola, and Starling (1944), in the eastern United States, studied the ecology of
pauropods, a little-known group that proved to be much more abundant in the soil than had
generally been believed. In England, however, intensive wartime studies of crop pests had been
in progress, particularly on wireworms (elaterid beetle larvae) which were of special
importance in crop fields that had previously remainded unploughed for long periods. From
these studies came, not only ecological information aimed at pest control, but the well-known
flotation soil-extraction technique of Salt and Hollick (1944) and the large-scale flotation tanks
of Cockbill et al. (1945) - see Fig. 46. And, of course, there was no lack of interest in
earthworms and soil fertility. This even stimulated the reissue of Darwin’s (1881) book on the
subject, with a foreword by Sir Albert Howard, in which the latter expressed strongly his
opinions against the use of “artificial” fertilizers.

Meantime, in neutral Sweden, extensive studies on the fauna of forest soil and litter had
been undertaken for some time. These resulted, near the end of the war, in another large
“land-mark” publication in the annals of soil zoology (Forsslund, 1945).

THE POST-WAR PERIOD TO THE 1960’S

Immediately after the war there was a gradual, then a rapid, expansion of work in soil
zoology, although the subject had, as yet, by no means developed into a discipline like
fresh-water biology. Pearse (1946) in the United States, Fenton (1947) in the United Kingdom,
and Gilyarov (1947) in the Soviet Union, produced important contributions on forest soil
faunas. Salt et al. (1948) continued with studies on pasture soils.

There was also considerable renewed interest in earthworms and their reciprocal
relationships with soil conditions (Evans and Guild, 1947, 1948; Evans, 1948; Guild, 1948;
Dawson, 1948; Dutt, 1948). Scandinavia, where soil fauna studies had always been pursued
with vigour, continued to produce impressive results. Haarlov (1947) modified the Tullgren
funnel so as to increase its efficiency (Fig. 42, inset); his was the basis of numerous subsequent
modifications, improvements and adaptations. Weis-Fogh (1948) related distribution of
Collembola and mites within the soil profile to pore-space, using an elegant, new technique.
Overgaard-Nielsen (1948) introduced new methods for nematode and rotifer extraction, and
Forsslund (1948) recorded unprecedented numbers of arthropods from forest soils.

The same year also saw the publication of Kubiena’s (1948) influential Entwicklungslehre
des Bodens. Although this was not the first time that this author had drawn the attention of
other pedologists to the importance of small arthropods in humus formation, nor was it to be the
last, it may be said that it was now, more than at any time previously, that “soil scientists” were
made to sit up and take notice of soil and litter fauna, other than earthworms, in the process of
soil formation.

The next few years saw the burgeoning of major works on the soil fauna. In addition to those
in more restricted fields, like that on the soil nematodes by Overgaard-Nielson (1949), they
included books covering a wide Field: Gilyarov’s (1949) Osobennosti Pochvy ... v Evolyutsii
Nasekomykh , dealing with the soil as a milieu for insect evolution; Franz’s (1950)
Bodenzoologie ..., largely relating to the importance of soil fauna for cultivation; Kiihnelt’s

Quaest. Ent., 1985, 21 (4)

432

Kevan

UNIVERSITY OF NOTTINGHAM
SCHOOL OF AGRICULTURE

Second Easter School in Agricultural Science

SYMPOSIUM and
COLLOQUIUM

SOIL ZOOLOGY

The University of Nottingham School of Agriculture,
Sutton Bonington, near Loughborough, Leicestershire,
England.

APRIL 1st to 7th, 1955 (inclusive).

Fig. 47. The historic, pale buff cover of the programme for the First international meeting of soil zoologists in 1955

Soil zoology

433

(1950) Bodenbiologie ..., a general text on the soil fauna, later translated into Spanish and
English, and Delamare Deboutteville’s (1951) Microfaune du Sol ... with an emphasis on
tropical as well as temperate conditions and taking into account soil formed above ground level
on trees. At this time too, there appeared another large research report on the (mainly
arthropod) fauna of temperate (beech) forest soil, which was to become a classic (Drift, 1951).
Then Hartmann (1951, 1952) stressed the importance of soil fauna in his books on forest soil
ecology. Soon afterwards came another important book, mostly relating to soil and litter fauna,
but one which is not widely known to soil zoologists, and even less to pedologists, partly because
of its title, The Biology of the Cryptic Fauna of Forests (Lawrence, 1953), and partly because
it relates mainly to southern Africa. Also regionally restricted was Fauna Pochv Latvi 7skoI
SSR (on the soil fauna of Latvia) by Eglitis (1954). Although it would seem invidious to try to
select any papers from the scientific journals, for particular mention, the valuable review by
Birch and Clark (1953) should perhaps be excepted.

Pedological works, such as those of Kubiena (1953), Handley (1954) and Wilde (1954),
now referred more and more to the importance of the soil fauna in humus formation, but it was
the year 1955 that saw Soil Zoology finally emerge as a discipline on its own. In the previous
year, almost exactly 30 years before this present meeting, I undertook, at the instigation of
Professor E.G. Hallsworth, to organize the first32 international colloquium in the field, to take
place from the 1st to the 7th April, 1955, at the University of Nottingham School of
Agriculture, Sutton Bonington, near Loughborough, England (Fig. 47). The meeting -
registration fee, then(\), 10 shillings ( ca . $2), accommodation - (all meals included!) £ 1.5.0
( ca . $5) per diem! - was an unqualified success, bringing together scientists from many
countries (though mainly European) and its proceedings (Kevan, 1955) published in record
time, became “out-of-print” almost at once and are now very hard to obtain on the second-hand
market. The Sixth International Congress of Soil Science, at its meeting the following year,
fostered a greater interest than hitherto in the soil fauna, and there were several papers given in
this field (International Society of Soil Science, 1956). This led to an ongoing series of
international Soil Zoology colloquia, beginning in association with the 15th International
Congress of Zoology in 1958 (International Congress of Soil Zoology, 1959; Murphy 1962) -
see also Note 32.

From the time of the 1955 meeting onwards, too, soil animals also became emphasized in
several major works on animal ecology, such as those of Tischler (1955), Macfadyen (1957,
1962) and Balogh (1958), in at least one introductory zoology textbook (Moment, 1958), and
in some general books on soil, such as those of Russell (1957), though the United States
Yearbook of Agriculture, Soil (Stefferud, 1957), had very little space devoted to the subject,
which was symptomatic of an unexpectedly retarded general interest in North America.
Though both Canada and the United States had had in the past, and continued to have, their
proponents of soil zoology (as distinct from those who worked with soil pests) they lagged far
behind Europe, both western and eastern, in the field. Courses and research in Soil Zoology, as
such, were, however, introduced by me into the McGill University (Macdonald College)
programme in 1958-59, where they uniquely continue. Canadian perspectives of the times were
also published (Kevan, 1959-61, 1962b).

In the late 1 950’s and in the 1960’s, there was a very large increase in the number of
publications on soil fauna in scientific periodicals. Of these, I will mention only a review by
Kiihnelt (1963), and a paper dealing with an important development in arthropod extraction
technique by Kempson et al. (1963) - see Fig. 48. Significantly, however, many books and

Quaest. Ent., 1985, 21 (4)

434

Kevan

Fig. 48. Extraction apparatus for small soil and litter arthropods; after Kempson et al. (1963). Slightly modified versions
of this type of equipment have resulted in the recovery of some of the highest recorded numbers of soil arthropods per unit

volume.

monographs (other than taxonomic works, which we will not cover here) devoted, wholly or in
large part, to the soil fauna appeared. Some of these were quite general, others restricted in
their topics; some were large, some small; some placed an emphasis on humus formation, soil
fertility etc., while others did not. They included: Pack’s (1956) book on the biology of
apterygote Hexapoda, important soil animals; the Spanish and first English editions of
Bodenbiologie (Kuhnelt, 1957, 1961); a text by Nosek (1957); a “popular” book by Farb
(1959); a research monograph by Haarlov (1960); Godfrey and Crocroft (1960) on the mole;
Principles of Nematology by Thorne (1961); a book with an agricultural bias by Kipenvarlitz
(1961); a general introductory text by Kevan (1962a), with an amended edition in 1968; a
South American faunal series edited by Delamare Deboutteville and Rapoport (1962-68); texts
by Schaller (1962, 1968) and Bachelier (1963), The Physiology of Earthworms by Laverack
(1963); small “popular” works by Dunger (1964) and Palissa (1964); a localized account for
the central Volga region by Aleinikova (1964); a voluminous tome on soil biology generally by
G. Muller (1964); and, by contrast, a little booklet for amateurs by Moreau (1965); Gilyarov’s
(1964, 1965) monograph for the identification of soil insect larvae and handbook for diagnosing

Soil zoology

435

soil types according to their fauna; a small hi^hschool teaching manual by Pramer (1965), and
a slightly more advanced introduction to soil life generally by Jackson and Raw (1966); a large
monograph by Loksa (1966); a thesis with special reference to tropical Africa by Maldague
(1967); a collection of specialist papers on various aspects of soil biology edited by Burges and
Raw (1967); a text by Brauns (1968); and other works by Dunger (1968) and by Lamotte and
Bourliere (1969), the last being of considerably wider significance than to purely soil ecology.

Meanwhile, further colloquia, symposia, etc., were being held in many places,
internationally, nationally and locally. Some of the proceedings of these meetings were
published as follows: Arnol’di et al. (1958), Murphy (1962), Dunger (1962), Klapp and
Wurmbach (1962), Doeksen and Drift (1963), Rapoport (1966), Byzova et al. (1966),
Minkevicius et al. (1966), Graff and Satchell (1967) and Aleinikova (1969). It is also
interesting to note the widening interest in the soil fauna that was developing among
non-zoologically oriented scientists. An address by P.W. Murphy to the Seventh International
Congress of Soil Science at Madison, Wisconsin, 1960, on the role of animals in soil formation,
was prepared for inclusion in one of the general sessions, though the text was never published
(Kevan and Murphy, 1960; Kevan, 1961); Section II of a 1965 symposium on soil-borne plant
pathogens began with an invitational address on soil fauna (Kevan, 1965); and the Ninth
International Congress of Soil Science in Adelaide, 1968, opened one of its principal sessions
with another such paper on soil fauna and humus formation (Kevan, 1968).

In this period, too, serial publications devoted very largely to research on soil fauna
appeared. The first of these was merely an annual information bulletin (newsletter) resulting
from a decision made at the Sixth International Congress of Soil Science (International Society
of Soil Science, 1956). It was called Microfaune du Sol and began under the directorship of J.
d’Aguillar in April, 1957. After the seventh number (in 1963) it was replaced by the larger and
more comprehensive Biologie du Sol , beginning April, 1964, which eventually became
Pedofauna in 1982 (nos. 35/36) under the direction of G. Wauthy. In the mean time, in 1961,
Ekkehard von Torne, in Austria, had started the scientific journal Pedobiologia, and C.
Delamaire Deboutteville, in France, launched Revue d’Ecologie et de la Biologie du Sol in
1963. Both journals remained predominantly zoological for many years.

RECENT TIMES

During the past few years, there has been no slackening off of work on the soil fauna, and it
would be quite impractical to try to review this here, even sketchily. Reference to abstracting
journals and to the proceedings of congresses, etc., however, will show that the field continues
to expand. One or two research publications for last year, taken at random, indicate that
activity varies widely from basic preliminary surveys of the myriapod fauna of northern North
America (Kevan, 1983, c, d) to the practical significance of the presence of heavy metals in the
food of larger soil invertebrates (A. Carter, 1983)33, both by Canadian authors, or as part of
general biological studies of soils derived from loose sediments (Loub and Haybach, 1983).

Books and monographs on soil-zoological subjects since 1970 include that of Gilyarov
, (1970), to some extent a morphological development from his earlier book (Gilyarov, 1949); a
j little “popular” account for home consumption by Haarlov (1970); a specialized compilation
i edited by Delamare Deboutteville (1970); two soil-fauna ecology books by Wallwork (1970,

1 1976); a small “popular” book, more relevant to litter than soil proper, by Savory (1971); an
i International Biological Programme handbook on qualitative ecology edited by Phillipson

i Quaest.Ent., 1985,21(4)

436

Kevan

(1971); a collection of contributed papers edited by Pesson (1971); Termites and Soils by Lee
and Wood (1971); Biology of Earthworms by Edwards and Lofty (1972); Economic
Nematology edited by Webster (1972); an advanced high school-level ecological text edited by
Andrews (1972) on soil organisms generally; a contribution to tundra ecology edited by
Tikhomirov (1973); and introductory text for soil ecology by Richards (1974); The Biology of
Free-Living Nematodes by Nicholas (1975); the enlarged new English edition of Soil Biology
(Bodenbiologie) by Kiihnelt et al. (1976); a text on soil invertebrates of the Soviet Far East by
Kurcheva (1977); a small, general soil ecology handbook by A.L. Brown (1978); some account
of the invertebrate fauna of brown and black soils in the Georgian Republic edited by Rekk
(1979); Nematodes in Soil Ecosystems edited by Freckman (1982) - in which some of the
contributions at last begin to consider the role of these animals in soil formation, as suggested
by Cobb (1915); the Marshall et al. (1982) assessment of the (unsatisfactory) position of
soil-faunal studies in Canada, which resulted in the present meetings; and Wallworks’ (1983)
little manual on earthworm biology. “Hot off the press”, I may mention a review on micro
arthropods and soil processes by Seastedt (1984) and a book on the distribution and ecology of
Collembola (Gilyarov and Chernova, 1984) which has just reached me from the U.S.S.R. Also
to appear shortly is a profusely (scanning-electron-microscope) illustrated work on European
soil arthropods by Eisenbeis and Wichard (1985).

For this same period we should also mention various colloquium and symposium
proceedings, including the following: Aguillar et al. (1971), Dindal (1973), Gilyarov (1973),
Vanek (1975), Gorny (1975), Lohm and Persson (1977), C.A. Edwards and Veeresh (1978),
Dindall (1980), Applehoff (1981), Veeresh (1981), Warden (1981), Satchell (1983), Lebrun et
al. (1983), Gregoire-Wibo et al. (1983) and (“stop press!”) Gilyarov (1984). It would also be
appropriate to note an increased soil zoology content in works not primarily concerned with
this; for example: in the 10th edition of Soil Conditions and Plant Growth (Russell and Russell,
1973); in The Role of Arthropods in Forest Ecosystems (Webb, 1977); in The Encyclopedia of
Soil Science (Kevan and Hill, 1979); in Recent Advances in Entomology in India (Prabhoo,
1981); in Soils, an Australian Viewpoint (Greenslade and Greenslade, 1983); in Laterization
Processes (Machado, 1983a, b); and in Acarology VI (Griffiths and Bowman, 1984).

CONCLUSION

In the foregoing I have included scarcely a reference to taxonomic literature, catalogues or
nomenclators, which may surprise those who know me primarily as a taxonomist. This is partly
because the subject is a vast one without definable boundaries, and partly because I hope that
currently usable taxonomic related literature (from which one can often deduce the historical
background) will be covered elsewhere at these meetings.

I would, however, stress that of all aspects of soil zoology, it is taxonomy that still needs the
greatest immediate expenditure of effort which means adequate financial support and
increasing numbers of continuing positions for qualified research workers in taxonomy. A
dozen years ago I spoke of taxonomy as being the Cinderella of the sciences (Kevan, 1973) and
I will not repeat what I said then. Since that time, however, the situation, at least in North
America, has, if anything, deteriorated, and the attitude or understanding (or both) of
non-taxonomists to the need for taxonomy has scarcely improved (see Note 33).

This does not mean that there has been no progress in the ability we now have to identify
accurately the various members of the soil fauna. Indeed, there have been, particularly in

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437

Europe (especially in the eastern countries), considerable advances, but even so, identification
is often difficult there, too, especially in the case of some groups. In most of the world,
including North America, any conscientious soil ecologist inevitably becomes bogged down by
taxonomic problems. If he does not identify his animals properly, his research becomes greatly
reduced in value, sometimes to the point of being worthless. On the other hand, without the
means of identification (except in a few groups like Northern Hemisphere Collembola), what is
he to do? The answer is, of course, “first things first; become a taxonomist!”. Some have indeed
followed this course, but the task-masters of most do not encourage it!

Soil zoology has come a long way from the days (mostly less than a century ago) when only
a few, relatively conspicuous forms (I will not say “species” as identification was seldom so
precise) were recognized, or even since the time of Gilbert White or Charles Darwin, when only
an inkling of the importance of animals in the humification process was evident. We have
reached all manner of degrees of sophistication in chemical, physical, statistical and electronic
techniques, but we still know very little about the soil fauna itself. And our ignorance will
persist until we can recognize one species of soil animal, in all its stages, from another, and
learn about the biology of each one that is important, and in what way it is so. This, in itself,
may not be immediately evident. Sometimes we may learn some answers, but mostly we do not!
It does no good, for example, to lump all the mites or Collembola together, as is often done, for
every species is different and each plays a different role. House sparrows are not usually
lumped together with hawks, or even with other finches, nor is “humic” acid the same as formic
and/or acetic! Yet this is the way in which “data” on the soil fauna have often been presented
by otherwise competent “scientists”!

Even in recent times, with a few notable exceptions, there has been, to mutual disadvantage,
a general scantiness of appreciation of the fauna on the part of “soil scientists” (how many of
them have ever taken an appropriate course relating to soil zoology, though a few may have a
smattering of microbiology?) and a reluctance to intrude into the realms of so-called “soil
science” by zoologists (though a few agricultural and forest entomologists may have been
exposed to an elementary course in “soil science” - without faunal content, of course!). There is
still an almost complete lack of concern or comprehension, especially, it would seem, among
those who determine (financially or otherwise) the directions of research, that we are still
without the means of proper identification of innumerable members of the soil fauna.
Furthermore there seems to be no realization that the understanding of basic soil ecology and
thus of the pedological importance of the fauna is impossible without arduous, long-term
taxonomic research and application, and that even “simple” identifications are very
time-consuming. The competence could be created, but, even where some exists, it is seldom
encouraged. Nay, it is discouraged! To admit to being involved in taxonomy seems to call for an
apology - if not an admission of failure to accomplish anything worth while! The pundits would
have others try to run before they have learnt to crawl, let alone to walk!

How far, then, have we really come since the Sumerians, or Aristoteles, or Albertus
Magnus, or Darwin, or even Konrad Diem? A little way, perhaps, though, in some respects, not
very far. We certainly know more species, but what do they do? As a result of these meetings,
but not overnight, perhaps we shall take one more faltering step forward! Who knows what we
might take two?

Our motto: “ Es wimmelt im Boden von Unbekanntemr (Gisin, 1947).

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Kevan

NOTES

1 The Akkadian for “ant” was given as zirbabw, the pale species (? termite) as zirbabu sadi; and the flying ant as
mutapriiu. (Note, the Akkadians, unlike the Sumerians, did not seem to associate taxonomically the wingless with the
winged ants; nor did they use a binominal system of nomenclature or seem to make much distinction between species; and
their successors, in turn, were no more enlightened.) The Akkadian for (annelid) worm was /sqippu or iTqapu; for “mole
cricket” (?) it was hallalua. , hallalia or hallullQ a; for “field cricket” it was sasiru qiste or sarsari; and for “dust locust” it
was erib turbuti. One might also add a legless reptile to the soil-dwelling fauna known to the Sumerians. This was called
Mui (generic for “serpent”) iginugal or Mus iginutug; the Akkadians called it puuhmahu or upputum. Scorpions (g/r;
Akkadian zuqaqipu) were, of course, also well known, as might be expected, and several different kinds were
distinguished.

2 Sandars’ (1959) translation is entomologically a little confused, using “nymph” for “imago” and “larva” for “exuviae”,
but it is quite comprehensible. The Akkadian word for dragonfly from which her version is translated is fc«/71u(m) and, for
the nymph, kirippu (see also R.C. Thompson, 1928, who refers to both as kinds of birds!). For various Sumerian and
Akkadian names for dragonflies, see Landsberger & Krumbiegel (1934). Gilgamesh, regrettably, does not refer to the soil
fauna, unless one includes scorpions as such; a passage of some length relates to the “Scorpion-men”. There is, however, a
mention of honey and one of flies. Otherwise, invertebrates include only the prize, finally lost, of the “flower” of
immortality at the bottom of the sea. This seems to have been based on a sea-urchin, rather than a sea-anemone, for the
alleged aphrodisiac properties of the former have been claimed throughout the ages.

3 The Rgveda (see Griffith, 1887) of ca. 1300 B.C.E. also makes reference to various kinds of “worms”, for example in
Book I, Hymn 191, and in Book II, Hymn 50, but these are to “poisonous” and parasitic species. The latter hymn is
particularly interesting because it almost undoubtedly refers to two very important parasites of man, Dracunculus
medinensis and Wuchereria bancrofti, neither of which, however, are soil-borne! The Atharvaveda (see Griffith, 1894), of
a rather later date, ? ca. 1000 B.C.E., has more numerous references to “worms”. A few involve plant injury, but most are
concerned with flesh-consuming dipterous maggots in carrion or wounds, or with helminthic endoparasites (e.g., Book II,
Hymns 31 and 32; III, 28; V, 23; VIII, 6; IX, 4; XI, 9). There is, however, in Book XII, Hymn 1, a single reference that
just might refer to earthworms:

“The worm, O Prithivi, each thing that in the Rain revives and stirs ...”

4 The New English Bible translation of 1970, in Deuteronomy [II Moses] XXVIII, 42, however, uses “mole cricket” for
the Hebrew zelazal , but this may be a mistranslation of an onomatopoeic word for a “whirring” (flying) locust of one kind
or another (see Kevan, 1978: 197, 385, 467). Prof. I. Harpaz (in litt. 1985), however, suggests a true cricket, such as
Gryllus bimaculatus.

5 Although of later date, it might also be appropriate here to mention the Jewish Talmud, which, like the Hebrew
Scriptures, refers quite frequently to insects (Bodenheimer, 1928, 1929, 1960). Again there is virtually no unequivocal
allusion to the soil fauna other than to ants, though Bodenheimer (1928) and Harpaz (1973) draw attention to the
suggested biological control of these by transporting soil between (widely separated) nests in order to bring about mutual
extermination by the inhabitants of each.

6 Herodotos (see Rawlinson, 1910) did not refer directly to the amphisbaena, but, briefly only, to gaint Libyan serpents
which were possibly part of the same tradition (though they may have been pythons from the south; the draco was the
largest of all serpents and was a constrictor). The Arabian Flying serpents, about which he has much more to say, and
which had featherless wings, ere clearly not soil-inhabiting amphisbaenas though, by Mediaeval times, the two seem to
have become amalgamated (Fig. 7, 8 lower). Not only did the amphisbaena have two heads, but, by then, it seized one with
the other and rolled along like a hoop! At the same time, it could also be Arabian and winged like the flying serpents of
Herodotos, and bipedal besides (c/. Druce, 1910; T.H. White, 1954; McCulloch, 1962; Rowland, 1973). It had illuminated
eyes (in contrast to Nikandros’ description) and, in later Mediaeval times, some believed that its glance, like that of the
bipedal Basilisk (discussed, for example, by the authors just mentioned), killed the beholder. An interesting feature of
Herodotos’ flying serpents was that, at the precise time of fertilization, the female seized the male by the neck and would
not let go until this had been bitten through. Surely the “flying serpents” were based on (elongate) praying mantids,
though I do not recall having seen this theory advanced elsewhere. Furthermore, Herodotos’ concept, that the male gets his
own back because the offspring devour the “womb” of the female, could be reconciled with the hatching of young mantids
from the ootheca, regarded as a detached female abdomen!. Thus the “classical” amphisbaena was originally based on a
small, burrowing, legless reptile, but the late Mediaeval version may owe its “existence” in large measure to aerial mantids!
The amphisbaena reverted to its true small form (only seeming to have a head at each end) in the 16th Century, though,
even by the end of that century, Aldrovandi accepted the two-headed concept - see Druce (1910), who cites various
classical and later authors in respect of it. There is a short precis of parts of Druce’s paper in T.M. White (1954). Further

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439

ancient beliefs that may be tied in with the mantis theory are discussed by Kevan (1985).

7 Although Latin versions of the Physiologus included less entries than the originals (the early 11th-Century metrical
version of Bishop Theobaldus - see Rendell, 1928 - had but twelve, though the ant remained - Fig. 1 1), fabulous beasts
multiplied in the later 12th- and 13th- Century “Bestiaries,” as did their mythical qualities and their importance in
symbolism, both religious and moral.

8 Later (705 A.D.) he became bishop of Sherborne. Aldhelm eventually went on to greater (posthumous) achievements, as
he later became a saint of the Roman Church, a position attained by remarkably few (former) inhabitants of Great
Britain!

9 Aldhelm also included riddles on the so-called “ Bombix ” or “silkworm” (perhaps not actually Bombyx itself, but some
other cocoon-spinning caterpillar); Apis the honey-bee; Locusta , the locust; “ Scnifes ,” the gadfly ( Tabanus ); “ Tippula ”,
the water-strider ( Gerris ), called “water-spider” by Pitman (1925); and “ Crabro ”, the hornet ( Vespa crabro). In addition,
his riddle on “writing-tablets” mentions “honey-laden bees”; and he has another, rather unlikely invertebrate on his list, the
“ Sanguisuga ” or medicinal leech ( Hirudo ). His riddle No. 18, “Myrmicoleon”, with Pitman’s (1925) English version,
reads as follows:

Dudum compositis ego nomen gesto figuris:

Ut leo, sic formica vocor sermone Pelasgo
Tropica nominibus signanspraesagia duplis.

Cum rostris avium nequeam resistere rostro.

Scrutetur sapiens, gemino cur nomine fungar!

I long have borne a name of hybrid form:

Both ant and lion I am called in Greek -
A double metaphor, foreboding doom:

My beak can not ward off the beaks of birds.

Let wise men search out why my names are twain.

10 In 10th-Century Byzantine illuminated manuscripts, presumably copies from much older documents, there are
representations of 8-legged, chelicerate (?) arthropods called “ myrmekion ” (see Kadar, 1978). These have been
interpreted as ants (though they look more like tail-less arachnids -? pseudoscorpions much enlarged); perhaps they are
supposed to be ant-lion larvae.

1 1 Sometimes called “Arabic” or “Islamic,” but both these terms impose unwarranted restrictions of race, language
and/or religion.

12 Born ca 1098, she became abbess of Disibodenberg (now Disenberg), in the diocese of Speyer, in 1 136; she founded a
new convent at Bingen in the Rupertsberg region, 1147, and died there 1179. She was famed for her visions, and
prophecies and regarded as a saint, but she was never formally canonized by the Roman Church.

13 The “Bestiaries” became especially popular in England (and thereby the English part of France). The trend begun in
the latter part of the 12th Century continued. Not only did fantasy increase, but so did the numbers of species mentioned,
up to more than 100 (James, 1928; T.M. White, 1954, McCulloch, 1962). Among “soil” animals (other than this cricket)
we begin to get further references (cf. Davis, 1958) to millipedes or woodlice (Fig. 13), as well as to earthworms, beetles,
etc., only some of which had appeared earlier, not in the Latin versions of the Physiologus, but in Byzantine Greek copies
of Nlkandros and Diaskorides (Kadar, 1978). These additions, by further transcription, became carried forward into later
centuries.

14 Neither referred to the amphisbaena. The works began to appear in various translations and, more than a century later,
in printed editions. That of Bartholomew, being the most concise, was the most popular; it was printed in over 40 separate
editions from 1470 onwards. The Liber de Naturis Rerum was often reproduced anonymously and usually attributed to
Albertus Magnus. An early Flemish translation, written (between 1265 and 1269) in metrical rhyming couplets, was that
by Jakob van Maerlant; it was called Der Naturen Bloeme (Bodenheimer, 1928). More notable, perhaps, was the German
version by Cunrat (or Conrad) von Megenberg, called Das Piich [= BuchJ der Natur , translated in the middle of the 14th
Century. This eventually, became the first printed book devoted exclusively to natural history, complete with woodblock
plates (Megenberg, 1475, Bodenheimer, 1928; Morge, 1973). One of these plates included illustrations of ants and
earthworms underground.

15 The Treatyse of Fysshynge was added to the second printed edition of her (1)Boke of Saint Albans , published by
Wynkyn de Worde, who changed the spelling of the lady’s name. The first known printed edition of the Boke was that of

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440

Kevan

I486, printed at St. Albans by an unknown “Schoolmaster”; it did not include this Treatyse. The discrepancy between
textual [1485($/c)] and bibliographic [1496] dates given by J.E. Satchell ( in Dindal, 1980: 848) may thus be explained.
[So far as I am aware, it is pure coincidence that two Satchells are involved here; the earthworm expert does not mention
his namesake, whose publisher was another Satchell.]

16 Such religious and moral works on animals later included those like Dierum Caniculorum by Simon Majolus, 1600
([earth] worms, ants, “ant-lions,” scarabaeoid beetles and cicadas mentioned), Animalium Historia Sacra by Wolfgang
Franz, 1612 (a similar range of soil inhabitants, though omitting “ant-lions” and including crickets), and later, more
famous Hierozoikon by Samuel Bochart(us), 1663 (also including references to a similar range of fauna) - see
Bodenheimer (1928, 1929).

17 Georg Bauer (1494-1555) was from Saxony and was appointed physician to the German mining town of Joachimsthal
in 1527. Thereafter he wrote many books on mining, metallurgy and chemistry. His interest in the subterranean fauna
clearly stemmed from his interest in mines and diggings, not vice versa.

18 Here might also be an appropriate place to mention what appears to be a recently perpetrated fallacy regarding the
16th Century. In discussing the gall-forming nematode, Anguina tritici , which passes part of its life-history in the soil, and
which causes what is known as ear-cockle of wheat, etc., Thorne (1961), as did others before him, suggested that the
parasite was referred to by William Shakespeare, in Love’s Labours Lost (Act I, Scene 4), when he wrote (about 1594,
first performed ca. 1595, printed 1598), “Sowed cockle, reap’d no corn.” It was not suggested that the causitive organism
was known (it was not discovered until 1743), but that infected seeds would not germinate. The Oxford English
Dictionary , however, gives no earlier than 1836 as the first use of the word “cockle” in this context. The “Corn cockle” is in
fact a caryophyllaceous weed, Lychnis galigo, though the name has also been misapplied to rye-grass ( Lolium ).

19 Also, although hated by farmers, careful attention was paid to mole crickets and their behaviour, for barley was seldom
planted in spring before their chirping was heard. It was also noted that hoopoes ( Upupa epops) eat mole crickets. This
may possibly stem from the old Greek play Ornithes (The Birds) by Aristophanes, in which hoopoes are said to dominate
over locusts or grasshoppers (see Kevan, 1978: 267-268), for these birds could scarcely be considered to be abundant in
Central Europe. It is more likely, however, that the Lapwing plover or Peewit ( Vanellus cristatus) was meant ( cf. Yapp,
1984). It is further noted that the head of a mole cricket worn around the neck cures fever - again probably derived from
an ancient source, for the wearing of a dead orthopteriod round the neck for this purpose is mentioned in a medical “jingle”
by Joseph Ursinus, 1541, as quoted by Bodenheimer (1928: 218), as well as by myself elsewhere.

20 Browne (1646) also partially exploded the fable of the (soil associated) ant and the “grasshopper” by pointing out,
Firstly, that a cicada, not a grasshopper, was involved (the complexities of this are discussed by Kevan, 1978), and
secondly, that the former insect lives for so short a time in summer that it need not “have recourse unto the providence of
the Pismire [ant] in Winter.” He was confused, however, as to what was a cicada, for these are virtually unknown in his
native England, and he regarded the inhabitants of “cuckoo-spit” as such. As both are Homoptera, this was not
unreasonable, though he said that from “cuckoo-spit ... some kind of Locust [sic] doth proceed. “ Had he paid a little more
attention to Aristoteles’ account of the life-history of cicadas, he should have known that they emerged from the soil.
Later, Erasmus Darwin took Browne to task, but that is another story!

21 The only two-headed serpent indicated was the “ Serpens Biceps ”, in which both heads were at the same end of the
animal, and which was apparently treated as a mere freak, as it had been previously regarded.

22 See also Oudemans (1926), who gives a Dutch title and a date of 1664; the mites he identifies as “ Hypopus ” [now
Anoetus ] feroniarunv, the nematodes (“ slangetjes ”) he says were Diplogaster [now Pristionchus] longicauda ; both
identifications were presumptuous, though credible.

23 The second edition, of 1740, substituted Fullo for Melolontha and added to the species in “ Scarabaeus ”; the genus
Cicada was added to the Hemiptera, from which Scorpio was removed to the Aptera; to the last was added the
collembolan Podura (from Geer, 1740), and the terrestrial isopods in “ Oniscus ” became “ Millipes ”. The 3rd edition, also
of 1740, differed little in content from the 1st. The 4th edition, of 1744, resembled the 2nd, but additional groups were
added, including Diptera and Hymenoptera, to the latter of which the ants were transferred. Subsequent editions (5th of
1747, 6th and 7th of 1748, 8th of 1753, from which botanical nomenclature dates, and 9th of 1756) gradually increased in
scope, but added nothing significant for our purpose.

24 The rest of this volume was to have dealt with Coleoptera. Volume 8 was to have been on orthopteroids, etc., and
volumes 9 and 10 on arachnids, myriapods and annelids (cf. Wheeler in Reaumur, 1926).

25 At the time, the condition was known as “malm”; the name “ear-cockle,” according to the Oxford English Dictionary,
did not appear (in print) until 1836.

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441

26 Gilbert White’s brother John included in the 1802 (posthumous) edition of “Selborne," various previously unpublished
Ms. notes by Gilbert (together with remarks by William Markwick). These included further comments on earthworms as
well as on ants, bugs, etc. Bingley (1803) plagiarized, almost verbatim, both original and supplement, though in dealing
with mole crickets he cited White as his authority.

27 This was not done for more than a decade, when Schmidt (1871) published his studies on the pest.

28 Sir John William Lubbock, Baronet (later Lord Avebury), who lived at Down in Kent, near Charles Darwin, was a
leading figure of the day, not only as a zoologist, Vice-President of the Royal Society and of the British Association, and
Vice-Chancellor (administrative head) of the University of London, but also as a prominent banker and Member of
Parliament. An even greater service than by his zoological writings that he rendered to mankind was to introduce the bill
which established August Bank Holiday (known also to a select few as St. Lubbock’s day!). His legacy lives on, even in
parts of Canada, though his name scarcely does so! As he was so busy, it is often suggested that most of his writing was
done for him by one or more “ghost” writers. This monograph, however, seems to have been entirely his own work (and he
has signed himself “From the Author” in a copy I possess). The excellent plates are acknowledged as being the work of a
Mr. Hollick, a deaf mute, and thus unique for the times.

29 Berlese’s most active period in this field was from 1897-1900. From 1904-1921 he continued to publish on all groups of
mites in Redia, 2-18.

30 I realize that this expression will not be familiar to all. Two Solitudes is the title of a well-known novel by McGill
University author (John) Hugh MacLennan. The book, published 1945, deals with the isolation of the “French” and
“English” cultures of Quebec, and of Montreal in particular.

31 Alfred E. Cameron, a Scot, came to the Canadian Department of Agriculture from England about 1916 (he was not
acceptable for the armed forces on account of his club foot). Later he became Professor of Zoology at the University of
Saskatchewan. Later still he was Reader in Agricultural Zoology at the University of Edinburgh, and it was there, about
25 years after his paper was written, that, as an undergraduate, I first read it and another in the same vein (Cameron
1917a) - my First taste of soil zoology. I did not immediately engage in this field, but I take this opportunity to express my
appreciation of what I owe to my late mentor.

32 It comes as a surprise to some to find that International Colloquia on Soil Zoology are now numbered as if they began
with the one held three years later at Rothamsted in 1958 (International Congress of Zoology, 1959; Murphy, 1962). Thus
the last one to be held in Louvain la-Neuve in 1982 (Lebrun et al., 1983) was numbered “VIII”, not “IX”! This is because
the Biology commission of the International Society of Soil Science seemed to consider that they had a prerogative
stemming from a decision made at their 1956 congress (Int. Soc. Soil Sci., 1956) at which, for the first time, the Society
had paid more than scant attention to the matter. The numbering therefore applies only to colloquia sponsored by the
Society. The intervening International Colloquia were as follows: II, Oosterbereek, Netherlands, 1962 (Doeksen and
Drift, 1963); III, Braunschweig, West Germany, 1966 (Graff and Satchell, 1967); IV, Dijon, France, 1970 (Aguillar et
al., 1971); V, Praha, Czechoslovakia, 1973 (Vanek, 1975); VI, Uppsala, Sweden, 1976 (Lohm and Persson, 1977); and
VII, Syracuse, New York, U.S.A., 1979 (Dindal, 1980). Colloquim “IX” will be in Moskva, U.S.S.R., 1985. It may also
be noted that the 1955 colloquium itself was likewise misnumbered, for it was called “The University of Nottingham
Second Easter School in Agricultural Science,” whereas it was really the first of its series. There had, indeed, been a not
very widely publicised series of pedology seminars conducted in 1953 by guest-lecturer W.L. Kubiena, but this was quite a
small affair without published “proceedings”. It may now be disclosed that it was dubbed the first “Easter School” only in
retrospect, the better to promote the “second”!

33 Including some of the same myriapods (millipedes), which, like most of the other animals referred to are unidentified.
This is typical of much work by “soil scientists” who emphasize “precision and accuracy of chemical analysis”, but who do
not even comment on the lack of this in the animal species investigated by them!

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Kevan

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Quaest. Ent., 1985, 21 (4)

466

Kevan

INDEX

SOIL FAUNA, ANCIENT,
MEDIAEVAL AND RENAISSANCE
LITERATURE ABOUT
V4/a’kh al-Makhluqat, 389
Aenigmata Aldhelmi, 383
Agamemnon , 375
Alexipharmakos, 376
Animalium Historia Sacra , 440
Aristoteles, writings, 376
Atharvaveda , 438
Beowulf, The Deeds of 384
Bestiaries, 383, 394
Biblos Historike , 385
Bibliotheca, 377
Boke of Saint Albans , 439
Byzantine literature, 381
Cheng-Lei Pen-tsao , 385
Chinese literature, 382, 384-385, 387, 400
Codex Animalium , 390
Collectranea Rerum Memorabilium see
Polyhistor, 382

Das Buch der Natur see De Naturis
Rerum, 439

Das Puch der Natur see De Naturis
Rerum, 390, 439
De Animalibus, 382, 387
De Animalibus Insectis, 400-401
De Animantibus Subterraneis, 394
De Differentiis Animalium, 397
De Divisione Naturale, 384
De Generatione Animalium, 376
De Incessu Animalium, 376
De Materia Medica, 381
De Naturis Rerum, 385, 390
De Partibus Animalium, 376
De Proprietatibus Rerum, 390
De Scarabaeis Britannicis, 410
De Universo, 384
Der Naturen Bloeme, 439
Dierum Caniculorum, 440
Egyptian literature, 389
Epistola ad Acircium, 383
Erh-ya, 375
Gedgraphikos, 381

Harra Hubullu, 373
Hayat al-Hayawan, 389
Hebrew literature, 375
Hellenic literature, 375
Herodotos, Writings of, 376
Hierozoikon, 440
Histories, Apodexis, 376
Historia Animalium, 376
Historie of Serpents, 401
Hortus Sanitatis, 390, 394
Insectorum Theatrum, 400
Islamic and Arabic literature, 384, 389
Kit ab al-Hayawan, 384
Les Mecaniques des Animaux, 407
Liber de Naturis Rerum, 387, 439
Libri Bestiarum, 381, 383, 385
Libris Physicis, 385
Mundus Subterraneus, 407
Naturalis Historia, 381
Naturalis Historiae, 394
New Feld- und Ackerbau, 401
Nuzhat-ul-Qulub, 389
Opus Naturarum, 387
Origines sive Etymologiae, 383
Ortus sanitatis see Hortus sanitatis, 390
Pen-tsao Kang-mu see Cheng-Lei
Pen-tsao, 400
Peri ZoQn, 381
Pharmaka, 381

Physiologus, 380-385, 389, 439
Polyhistor, 382
Rgyeda, 438

Reductoria Moralia, 394
Ruralium Commodorum, 401
Ruralium Commodorum Libri XII, 389
Sanskrit literature, 373
Saracen literature see Islamic and Arabic
literature, 384-385, 387, 389
skolopendra, 381

Speculum Maius Tripartitum, 387
Sylva Sylvarum, 402
The New English Bible, 438
Theriakos, 376-377
Theriotropheum, 401

Soil zoology

467

Topographia Hibernicae, 385
Treatyse of Fysshynge wyth an Angle ,
390, 439

Tsu-chi King , 387

NAMES OF ORGANISMS, REAL

AND MYTHICAL

‘aygabuf, 389

Acarus, 407, 409, 412

Acarus geniculatus, 405

Acarus holosericeus , 402

Acarus scaber, 414

Acarus vegetans , 416

Acheta domesticus , 395, 410

Achiptera coleoptratus, 414

ad-dinnah, 389

ad-dud, 389

Agrotis, 410

A grot is ypsilon, 372

al-'arada , 389

al-'ugrui, 389

al-‘ukuban , 389

a/-asari’, 389

al-fazir, 389

al-gathlah, 389

al-gu’al, 389

al-gudgud, 389

Allolobophora longa , 390

Ameise, 395

amphibia, 421

amphipods, 410

amphisbaena, 375-377, 381-383, 385,

387, 389, 394-395, 397, 401-402, 410,
421,438-439
amphivena , 387
Anguillula dipsaci, 421
Anguina, 421
Anguina agrostis, 417
Anguina tritici , 412, 417, 420-421, 440
Anguis fragilis, 395
Anoetes polypori, 412
ant-dogs, 381, 383

ant-lions, 375, 381-384, 387, 394, 397,
400, 409,412,414, 440
larvae, 412, 421, 439

larvar, 390

ants, 371.8, 372-373, 375-376,
381-385, 387, 389-390, 394-395,
397, 400-401, 405, 407, 409-410,
412, 414-416, 420-421, 438-440
ants, carpenter, 389
ants, winged, 405, 438
ants, wingless, 405, 408, 438
Apis , 439
Aptera, 412, 440
aquatic nymphs, 373
arachnids, 439
aradat , 389
Aranei nigri, 395, 397
Araneus, 409, 412
Armadillidium , 381, 387
as-simsimah, 389
as-surfah, 389
Ascarides terrenae , 40 1
Ascaris, 412

394-395,410,412
ash-shaisabcm , 389
at-thathrag, 389
auksimem , 385
Auricularia, 412
badgers, 385
basilisks, 403, 410
bees, 394, 407, 439
bees, burrowing, 376, 401
beetles, 384,387, 389,410, 439
beetles, burying, 371.8
beetles, excommunication of, 407
beetles, scarabaeoid, 375, 381-382, 385,
389, 394, 401-402, 405, 412, 415, 421,
440

beetles, scarabaeus ( = Geotrupes ), 384

beetles, soil-dwelling, 383

bergmenkel/kobel/guttel, 396

bergteufel , 396

Bibio hortulans , 412

black spiders ( = Lycosidae), 397

Blattae , 394-395

Blattodea,

cockroaches, 395
blindschleiche , 395
Blindworm, 395

(Ws/. Ent., 1985,21 (4)

468

Kevan

Bombyx , 439
booklice, 409
bristletails, 410
brotworme, 395
Bugga Bug, 419
bumblebees, 376, 410
Buru balag(-gana), 373
Buru saharra, 373
Buru zapaag(-tira), 373
Caecilia, 403, 410
camel-cricket, 372
Cantharellus auratus, 412
Cantharis formicaria latior, 401
Carabidae, 400,410,412,415
Carabus, 412
Carabus auratus , 401
Cardiosoma, 415
carpenter (ants), 389
caterpillars, 382, 410
centipedes, 375-376, 381, 394-395, 401,
412

Cercopidae, 383-384, 394
Cetonia, 394
Cetonia aurata, 401, 410
yfir emseqa ( = earthworms), 376
chafers, 401

cherubim (? = Scarabaeus ), 375
Chilopoda, 395
Cicada , 440
Cicada orni , 4 1 2

Cicadae ( = Cercopidae), 384-385
cicadas, 372, 375-376, 381, 383-385, 387,
390, 394, 401, 405, 409-410, 440
cicadas and Amerindian culture, 372
cicadas and Chinese culture, 372
cicadas and Greek culture, 372
clothes-moth larvae, 384
cockchafers, 397, 410
larva, 395

cockroaches, 394-395, 400, 405, 415
Coleoptera, 395, 410, 412, 414-415, 421,
440

Collembola, 394, 408-409, 412, 416-417,
420-421,425,430-431
Coprinus , 405
Crabro , 439

crane flies, 405, 409-410, 412, 421
cri(s)non, 387

crickets, 382, 387, 389-390, 394, 397, 402,
410,415,440
crickets, fighting, 387
crickets, house, 395, 397, 410
crustaceans, 407
cuckoos, 383
Culices fematarii, 401
Curtilla, 401

cutworms, 372, 376, 410, 421
Daemonum, 396
digger wasps, 376
Diplogaster , 440
Diplogaster micans, 42 1
Diplopoda, 387, 395
Diptera, 410, 440
Ditybachus dipsaci, 421
dor beetles, 400-401
draco, 438

Dracunculus medinensis, 438
dragonflies, 373, 438
dragons, 384, 403, 410
dud, 389
dung beetles, 389
dust locusts, 373, 438
ear-cockle eelworm, 421
earth fleas, 394
earth flies, 394
earth lice, 401
earth mites, 407
earth’s entrails, 376, 379
earthworms, 373, 375-376, 383-384, 387,
389, 394-395, 400-402, 405, 407, 409,
412, 420-422, 425, 430-431, 438-440
earwigs, 389, 400-402, 407, 414
echinoderms, 400
ectoparasites, 417
eels, 376
eelworms, 421
Egyptian scarab cult, 375
eims, 395
E later, 415

Elateridae, 400-401, 410

wireworms ( = larvae), 401, 430-431
emmet, 395

Soil zoology

469

Enchytraeidae, 422, 425
Enchytraeus , 420
Enchytraeus albidus, 420
endoparasites, 438
engerle, 395

Engerling ( = Enger =Inger ), 395, 407

erib turbuti, 438

Feldgrille, 395

feldheim, 395

feltmaus, 395

Ferae, 410

field crickets, 373, 382, 384-385, 389,
394-395, 397, 400-401, 410, 414-415,
417,421,438
flea beetles, 394
fleas, 384

flying ants, 373, 438
flying serpents, 438
Forficula, 410,412
Forficula auricularia, 401-402, 407
Formica , 395, 401, 412
Formica-leo, 412
formicaleon, 384
Formicidae,
see ants, 395
foxes, 394
Fullo, 401,440
gadflies, 439
gastropod molluscs, 425
Geotrupes , 384, 395
Geotrupes stercorarius , 401
Geotrupidae, 400-401, 407, 409-410, 412,
414-415
Gerris, 439
giT, 438
Glires, 410
Glomeris , 387
gordiid worms, 420
i Gradientum, 395
grasshoppers, 372, 440
great angle Twytch, 390
i gresillon , 387
I i ground-beetles,

see Carabidae, 387, 401
j grubs, 395

j Gryllotalpa , 373, 400, 405, 407, 409-410,

412

Gryllotalpa gryllotalpa , 401
Gryllus, 410,412

Gryllus ( =Acheta ) domesticus , 395

Gryllus agrestis, 395, 397, 401

Gryllus assimilis, 414

Gryllus campestris, 395, 401, 410, 415

Gryllus domesticus , 394

Gryllus niger, 415

Gryllus pennsylvanicus, 415

Gryllus veletis, 415

Gryllus, s. str , 384

hallali a, 438

hallalua, 438

hallullaa, 438

/nmaru-l-qabban, 389

hausheim, 395

hayzabun, 389

Heimchen, 395

Hemiptera, 412, 440

Hepialis humuli, 412

Heterodera rostochiensis, 426

Heterodera schachtii, 421-422

/i/mar-kabban, 389

Hirudo , 439

Hister, 4 1 5

Hologamasus lichenis , 417
Homoptera, 440
honey-bees, 439
hookworms, human, 415
hoopoes, 440
horned scarabs, 415
hornets, 394, 439
Hundertfiissler, 395
Hymenoptera, 440
Hypopus, 440
/oulos, 379, 381
iginugal , 438
Insecta (larvae), 395
intestinum terrae , 412
invertebrates, 383, 385
/Iqapu, 438
tfqippu, 438
Isopoda, 395

isopods, 387, 401, 405, 407, 410, 412, 421
jigger fleas, 402, 405, 410, 415

i Quaest. Ent., 1985, 21 (4)

Kevan

470

Julus, 412

June-beetles, 415

kantharoi, 376

kharatin , 389

khunfusa , 389

kirippu, 438

tfwi, 373

A7si kurra, 373

Kisi ririga, 373

kult\u(m), 438

land-crabs, burrowing, 415

Lapwing plover, 440

leech, medicinal, 439

Umax , 412

lizard, 376

lizards, legless, 421

Locusta, 439

locusts, 389, 438-440

Lumbricidae, 395

Lumbricillus lineatus, 417

Lumbricillus minutus, 417

Lumbricus , 395, 409, 412, 417, 421

Lumbricus latus, 412

Lumbricus terrestris, 390

Lycosidae, 395, 397

Macrotermes , 409

Magdalenian culture, insects in, 371.8
maggots, 375, 438
Maik'aferlarve , 395
Manis, 381
mantids, praying, 438
Mar , 373
Mardib, 373
Mar ga/, 373
Mar Sasur, 373
Mar tab, 373
mar gar it a, 385
Maulwurf, 395
May beetles, 405
larva, 395

May beetles, excommunication of, 389
meiworm, 395
Mellivora capensis , 381
Melolontha , 389, 395, 401, 405, 407, 410,
412, 440
larvae, 407

mermecolion, 385
Merodon equestris, 412
Metamorphumena, 410
millipedes, 376, 381, 387, 394-395, 412,
414, 422, 439
Millipes, 440
mirmicoleon, 381

mites, 401, 405, 409, 412, 421-422, 425,
431,440

mites on beetles, 408, 410, 415

mites on earwigs, 412

mites on insects, 376

mole crickets, 373, 375, 382, 384-385,

390, 400-401, 405, 407, 409-410, 412,
414,417,421,438,440-441
moles, 376,383, 387, 395,421
molwurff, 395
mononchid eelworms, 426
mwq, 389
Mus, 438
Mus iginutug, 438
Muscae , 410
mutaprisu, 438
Myremeleontidae, 414
myriapods, 385, 405, 410, 421-422, 425,
430

myrmecoleon, 381, 384
myrmekion , 439
Myrmeleon, 384
Myrmeleontidae, 409
Myrmicoleon, 384
naml, 389

Narcissus bulb-fly, 412
Natantium, 396
Necrophorus , 371.8, 408
nematodes, 389, 403, 405, 425
Nematomorpha, 420
Nemobius sylvestris, 407
Neuroptera, 409
Oniscoidea, 395
Oniscus, 412, 440
onoi hoy hypo tas hydrias, 38 1
Onychiura ambulans , 417
Opilio , 409
Opimacus , 390
pangolin, 381

Soil zoology

471

Parasitus coleoptratorum , 407-409, 412

Parasitus mites, 410

pauropods, 431

pearls, 385

Pediic[ulus /, 412

Peewit, 440

Pergamassus crassipes, 414, 417

Phalangium, 412

phoretic mites, 407

phthiracarid mites, 430

Podura, 440

polychaete worms, 412

Polyphagidae, 400

Potato root eelworm, 426

Pristionchus longicauda, 440

prostigmatid mite, 414

pseudoscorpions, 376, 412, 439

puuhmahu , 438

Pyrgomorphidae, 373

pythons, 438

qish'iban , 389

rabbits, 394

rainworm, 395

ratel, 381

Regenwurm, 395

Reptilia, 403, 412

root eelworm, 421

rootgrubs, 410

Rose chafers, 394, 40 1 , 4 1 0

sarsari , 389, 438

sasiru qiste, 438

Sanguisuga, 439

Scarabaeidae ( = sacrab beetles), 382
Scarabaeidae ( = scarab beetles), 372,
375-376, 389, 407

Scarabaeidae and Egyptian culture, 372
scarabaeoid larvae, 394
Scarabaeoidea, 400, 410
Scarabaeoidea (larva), 395
Scarabaeus, 395, 410, 412, 415
Scarabaeus bufonius, 401
Scarabaeus majalis, 400
Scarabaeus pilularius , 401
scarabs, 375
Schtiflein , 395
Schabe , 395

schefflein, 395
sciarid fly maggots, 422
Scnifes, 439
Scolopendra, 395
Scolopendrae , 394
Scolopendria, 412
Scolopendria marina , 412
Scolopendria terrestris, 412
Scorpio , 412, 440
Scorpio aquat., 412
Scorpio terrestris , 4 1 2
Scorpio-araneus , 4 1 2
Scorpion-men, 438

scorpions, 376-377, 387, 401-402, 405,
438

Scytale, 403
Serpens Biceps , 440
serpent-lizards, 385
Serpentium, 395
serpents, 377, 403, 438
serpents, flying, 438
Seufkdfer , 395
sharrar al-lail, 389
shrew(mouse), 395
silkworms, 439
Silpha , 415
Silphidae, 400
Slow-worm, 395

slugs, 400-401, 405, 407, 409, 412, 421

snails, 382

snakes, 376, 410

Sore*, 395,410

sowbugs, 395

spiders, 385, 387, 395,405,412
spiders, burrowing, 401
Spitzmaus, 395
Spondylis , 40 1
spring wibel, 395
Staphylinidae, 401, 410, 416
Staphylinus, 412,415
T 395
Tu0Ac 395

Sumerian entomology, 373
/atuk, 389
Tabanus , 439
7a//?a, 395,410

(Wj/. 1985,21 (4)

Kevan

472

Talpa europaea, 430

talpa insetto(= mole cricket), 400

tant, 407

Tarantula , 412

tarmes, 376

Tausendfussler, 395

tettiges, 372

Tenuitarsus angustus (Blanchard), 373
Termes, 415

termites, 371.8, 373, 376, 384, 387,

389, 400, 405, 407, 409-410,
414-417,421-422
termites as human food, 417
tettigometra, 376, 381
ticks, 409, 412
Tippula, 439

Tipula paludosa, 405, 409-410, 412

toads, 401

tola’ath, 375

Troglophilus, 372

Trogonophis, 381

Trombidium, 402

Trombidium holosericeum, 407, 409, 414
ts ’an, 384

Tunga penetrans , 402, 405
Turbatrix aceti, 403
Typhlops , 381
Ub pad , 373
upputum, 438
Upupa epops, 440
Uropodidae, 412
Vaginipennia, 410
Vanellus cristatus , 440
vermes, 384
Vermileo , 414
Vermileo vermileo , 409
Vermis in Maio , 397
Vermis in Maio netus, 395
Vermium , 395
Vespa, 394
Vespa crabro, 439
Vespula, 394
vinegar eelworms, 403
Volantium, 395
Vormela, 395
wasps, 394, 407

wasps, digger, 401
water-spiders, 439
water-striders, 439
weevils, 395
whitegrub, 395
whitegrubs, 395, 401, 415
Wibel (= Wiebel), 395
wolf spiders, 397
wood crickets, 407

woodlice, 376, 381, 385, 389, 394-395,
439

Wormlein, 395
worms, 375, 389, 438
Wurmchen, 395
Wurmlein, 395
Wuchereria bancrofti, 438
zelazal, 438
zirbabu, 438
zirbabu Sadi, 438
zuqaqipu, 438

SOIL MICROMORPHOLOGY AND SOIL FAUNA: PROBLEMS AND IMPORTANCE

S. Pawluk

Department of Soil Science
University of Alberta
Edmonton, Alberta T6G 2E3
CANADA

Quaestiones Entomologicae
21:473-496 1985

ABSTRACT

Surface soil layers were viewed microscopically along a pedogenic gradient from the
northern Arctic to the southern Parklands within the Interior Plains region and westward to
the Alpine and Interior Grasslands of British Columbia. In all instances it appears that soil
animals play a major role in structural development although the relationship between humus
form, synecology and microfabrics remains vague. Among all animals present microarthropod
influences are the most ubiquitous in their influence upon soil microstructures and humus
formation. Larger animals are more prominent in the Parkland region and appear to play a
major role in regulating humus form; while moder humus form is most evident in the cold
northern regions of the Arctic, proto-mull appears to be more characteristic of the warmer
Parkland environment. Humus form of the Interior Grasslands is generally characterized by
moder and little is evident for the action of larger soil animals upon development of soil
microstructure; the reason for this is not clearly understood.

RESUME

Les horizons superieurs des sols ont ete examines au microscope le long d’un gradient pedogenique s'etendant du
nord de I’Arctique jusqu’au Parklands des Plaines au sud et a I’ouest jusqu’aux Prairies alpines el des plateaux
interieurs de la Colombie-Britanhique. Dans tous les cas, il semble que la faune du sol joue un role primordial dans le
developpement de I’aspect structural, me me si les rapports entre la forme de I’humus, la microtexture et la synecologie
sont encore vagues. De toute la faune du sol, les microart hropodes contribuent le plus a la microstructure des sols et d la
formation de Ihumus. Les elements plus gros de la faune sont plus communs dans la region des Parklands et semblent y
jouer un role primordial dans la determination de la forme d’humus. La forme moder d'humus est plus repandue dans
les regions froides de I’Arctique alors que le proto-mull semble caracteriser davantage I’environnement plus chaud des
Parklands. La forme d'humus des Prairies de I’interieur est generalement cracterisee par du moder, et, pour des raisons
que I’on s’explique encore mal, on y observe peu d’evidence indiqant Paction des elements plus gros de la faune du sol sur
le developpement de la microstructure.

INTRODUCTION

The Problem

The principal problem facing those who work in micromorphology and formation of soil
microstructure in relation to faunal activity, is the general lack of clarity as to the importance
of soil animals in initiating and maintaining soil fabric rearrangement. Of secondary
importance is the need for more precise cataloguing of a specific feature or features that each
organism or group of organisms is capable of contributing to the reorganization of soil
materials.

474

Pawluk

Early Research

The importance of soil animals in soil structural development has long been recognized.
Contributions from earthworms have been singled out for special attention by early researchers
such as Charles Darwin and P.E. Muller. Kubiena (1953) described forest mull as comprising
earthworm casts and their residues and to this he attributed the ‘crumb’ structure that is so
characteristic of these layers. However, as Jacks (1963) accurately pointed out, while it is
generally accepted that earthworms create crumb structures of Russian Chernozems, these
animals are not generally all that common in North American Prairie soils and crumb mull
structures must be produced without earthworms.

Many previous researchers involved in this area of study were convinced other animals were
also important contributors to soil reconstruction at the microscopic level (Kubikova and
Rusek, 1976; Zachariae, 1963, 1964; Babel, 1973). They not only emphasized the ecological
importance and soil genetic contributions of faunal associations but in some instances were able
to assign unique fabric arrangements to manifestations of very specific biological activity. Even
in his initial work Dr. Kubiena (1953) insisted upon a firm genetic role for soil organisms in
reorganization of soil fabric.

Classification of Soil Microstructure and Faunal Activity

From 1938 to 1970 Dr. Kubiena published several textbooks and many scientific articles in
which he clearly set forth his concepts on this subject. His fabric type most closely associated
with faunal action was spongy microfabric. Spongy microfabrics were defined as consisting of
aggregates bound to each other in a manner that forms a system of interconected voids and
cavities. The internal structure of the aggregates generally remains quite porous. Spongy
microfabrics are most frequently associated with mull layers common to the A horizons of soils
such as those of the Chernozemic and Brown Forest groups and were believed to be derived
entirely through the activity of diverse faunal populations, especially earthworms and
potworms. This type of fabric arrangement is regarded as superior to all others from an
ecological and management standpoint. Kubiena (1938) also paid attention to forms of moder
humus. However, because of the non-coherent nature of these materials their classification was
considered at the elementary fabric level as some variation of the agglomeratic related
distribution pattern. This approach is understandable since moder humus comprises a loose
mixture of partially decomposed plant remains, mineral fragments and numerous droppings of
small arthropods.

Kubiena’s terminology for classification of biologically generated soil microfabrics provided
a basis for further development by other workers. Up to the time of publication of his text on
‘Fabric and Mineral Analysis of Soils’ in 1964, Brewer had not given particular emphasis to
faunal processes as a basis for classification, however, their influences upon the soil were
recognized. Fecal pellets were described as a special kind of pedological feature and when
deposited in a recognizable channel or chamber they comprised the inner material of some
varieties of pedotubules. Unfilled channels and chambers have been attributed in some
instances to the burrowing action of animals.

Barratt (1964) attempted to clarify some of the confusion that existed in the classification of
humus types and humus forms. Subdivisions of forms of mull and mor humus were described
micromorphologically. Class divisions were based on the manner in which organic matter was
incorporated (material composition) as well as on the organization (arrangement) of material.
Terminology for the initial classification reflected terms used for subdivisions of various humus

Soil micromorphology and soil fauna

475

forms. Later Barratt introduced new terminology recognizing reorganization and composition
of fine (-col) and coarse ( -skel ) materials. The terms pelleted and spongy humical and
mullicol most frequently served to distinguish fabrics derived from discrete and welded casts
from various soil animals, some of which are capable of intimately incorporating fine mineral
components within the humus forms. Fecal pellets were recognized as an important component
in all but raw humus forms.

Stoops and Jongerius (1975) also devised a classification based on spatial arrangement of
fine and coarse materials. Aggregated materials such as fecal pellets were considered as
discrete entities and divided into fine (f) or coarse (c) material depending upon size. Thus,
fabrics comprising discrete fecal pellets all falling into one size group would be classified as
monic while fecal pellets of finer size arranged in the intergranular spaces of coarse skeletal
particles would be classified as enaulic. Kinds of materials were recognized by using suitable
suffixes as for example recognizable plant fragments as phyto -, well decomposed humus as
humo-, clay minerals as argio-, etc. As with Barratt’s classification, unique kinds of fabrics
could be assigned to activity of specific fauna or faunal groups.

Drawing on concepts by previous workers interested in fabrics of soil organic matter, Bal
(1973) proposed that major emphasis be placed on the soil organic component in his concept of
the humon. The humon is defined as “the collection of observable organic bodies in soil which
are characterized by specific morphology and spatial arrangement.” Excrement or modexi and
comminuted plant material were considered to be important components of the humon and
their specific morphological classification was based on size, shape, composition and
distribution. Bal thus proposed a micromorphometric system for identification of fecal casts but
one lacking emphasis on mode of origin. As Bal pointed out, characteristics of modexi are not
always unique for a single animal species and knowledge of populations is also an essential
element required for assignment of mode of origin. This is especially evident when aging of
excrement has progressed to the point where individual modexi are no longer recognizable. In
some instances the genetic origin of the fabric type may remain in doubt, since similar features
may also be formed by non-biological processes. Biochemical substances (Martin, 1946), frost
processes (Post and Dreibelbis, 1942; Fox, 1979) and wetting and drying (Russell, 1973) are
examples of processes to which granulation of soil material has been attributed in the past.
Formation of organo-clay complexes in mull layers has been attributed to biochemical
processes active outside of, as well as within, the intestinal tracts of soil animals (Satchell,
1967).

A micromorphological Classification for Western Canadian Soils

Recent investigations of western Canadian soils developed under grassland, tundra and
alpine plant communities with a significant component of grass, forbs and shrub species
, ; revealed A horizons with strong granulation at the macro and/or micro levels of fabric
5 reorganization. These soils are in various Soil Orders in the ‘Canadian System of Soil
c Classification’ but all have well developed Ah horizons. In order to accommodate these fabric
arrangements into a suitable micromorphological classification system, Brewer and Pawluk
,( (1975) further developed a scheme published by Stoops and Jungerius (1975) that recognized

special related distribution patterns between fine (f-matrix ) and coarse (f- member ) material.
Brewer (1979) later introduced the concept of fabric sequences which allowed grouping of
I fabric types that exhibited unique genetic relationships. In this regard the fabric sequence that
uS best accommodates the granular character of our soils is the granic sequence. The granic

Quaest. Ent., 1985, 21 (4)

476

Pawluk

sequence comprises four fabric types: granic, granoidic, granoidic porphyric , and porphyric.
The granic fabric type is used for microstructures comprising units that are discrete and
unaccommodated. Such an arrangement is commonly associated with discrete fecal pellets.
Granic units of fabric partially coalesced or fused at their edges are referred to as granoidic
fabric type (Kubiena’s spongy fabric). Granoidic fabric types commonly grade to porphyric
types i.e. coalesced units become more densely packed, individual units are no longer
recognizable and form a vughy or porous groundmass of coherent soil material. Composition of
encompassing material is defined through the use of appropriate prefixes: humi - well
decomposed humus; mull - organo-clay complexes of the mull humus form; phyto - partially
decomposed plant remains; ortho - mineral grains; matri - soil matrix constituents (largely
inorganic). A mull with spongy fabric is thus designated as mullgranoidic fabric type. Since
surface soil horizons of many of our most agriculturally productive soils show mixed granular
microstructures with a variable degree of coalescence, the granic sequence is an especially
useful concept for descriptive purposes and is used throughout the remainder of this discussion.

Objectives

Surface layers of soils, when viewed microscopically along a pedogenic gradient from the
northern Arctic to the southern Parklands within the Interior Plains region and westward to the
Alpine and Interior Grasslands of British Columbia, reveal interesting ecological and
micromorphological relationships. In all samples it appears that soil animals play a major role
in structuring and regulating soil microfabric development. Yet, the relationship that exists
between humus form, synecology and microfabrics is poorly understood. While no attempt can
be made here to fully develop a meaningful understanding of the dynamics,
micromorphological features will be presented in an attempt to illustrate some of the resulting
features within the soil fabrics and problems related to discerning their genesis will be raised.

OBSERVATIONS ON SOIL MICROSTRUCTURES FOR SURFACE SOIL LAYERS
FROM DIFFERENT BIOCLIMATIC REGIONS

Northern Tundra Region (Table 1.1)

Surface soils examined from the ridge area on Devon Island showed comminuted plant
material associated with mineral grains and fine (20-50 nm) humigranic units in a granic
fabric arrangement. The humigranic units were relatively uniform in size but somewhat
irregular in shape. Some were fecal pellets of microarthropods most likely Collembola and some
were melanized plant fragments and cellular tissue. Numerous loosely bound fecal pellets of
200 ium size and comprised of smaller fecal pellets, melanized plant materials and silt size
skeleton grains were similar to those reported in the literature for Enchytraeidae. Plant
material appeared to be darker and more strongly humified in these latter fecal pellets. The
composition is an expression of the feeding habit of the larger fauna. Uniformity in size and
shape of fecal pellets reflects the limited diversity in faunal population of these soils. The lack
of clay mineral constituents in the fecal material at least in part results from the low content in
the soil.

Southern Tundra Region (Table 1.2)

Three well drained soils were studied in the southern Tundra region. Humus forms were
essentially similar at all three sites. The moder humus form was well developed and comprised
a dominance of humified comminuted plant fragments and fecal pellets from various fauna.

Soil micromorphology and soil fauna

477

The majority of fecal pellets were of 30-50 /urn size, irregular in shape and made up of
amorphous humic materials. These were believed to be the droppings of Collembola. Larger
humic fecal pellets of 200 /im size in the upper layer frequently contained loosely bound fecal
pellets of smaller size as well as humified plant fragments. Similar fecal pellets reported in the
literature (Kubikova and Rusek, 1976) were considered to be droppings of Enchytraeidae. Well
developed humic fecal pellets of 350-600 /im size resembled droppings of Diplopoda. In some
instances the larger fecal pellets were either disintegrating or were being destroyed by
microarthropods that left their droppings in the voids. Mull-like fecal pellets of 350-400 /tm
size formed a thin horizonal zone in the lower moder layer. These fecal pellets were generally
smooth, lobate and dense, closely resembling those of Diplopoda. The occasional large humus
fecal pellets (900 /tm) in this zone were similar to those of Diptera larvae or earthworms.
Alignment of unassociated coarse mineral fragments suggested incorporation into the humus
layer by frost action.

The immediate underlying soil layer appeared to be a ‘proto’ mull or mull-like moder. Some
smaller mullgranic units of 50-90 /tm size strongly resembled Collembola fecal pellets although
others appeared smoother and more rounded. Fecal pellets of Enchytraeidae were also very
common in this layer. Strong coalescence made it difficult to discern the original nature of most
units. Units in the underlying layer became much larger and made up of loosely packed
mullgranic material 450-500 /im in size with smaller units of 40-200 /urn size in the voids. The
smaller units were fecal pellets of Collembola and Enchytraeidae. Origin of the larger units was
difficult to discern. In some cases smaller fecal pellets were observed within the larger units.
This probably reflects microarthropods feeding on the larger units. The well-rounded
moderately accommodated weak mull-like granoidic units in the A(B) were relatively large
(1-2 mm) and their origin is unknown. The relatively high amounts of amorphous material
found in the fine fraction suggests possible binding associated with freeze-thaw processes.

Forest-Tundra Transition Region (Table 1.3)

The raw soil humus of the lower leaf mat contained a high percentage of partially
decomposed comminuted plant fragments together with small (50-90 /tm) cylindrical humic
fecal pellets most likely of Collembola. There were zones in which the larger fecal pellets
(120-200 fim) of Enchytraeidae were concentrated. The underlying forms of moder humus
contained an abundance of humic fecal pellets of 50-200 /urn size that probably at least in part
reflected the activity of Collembola and to a greater extent Enchytraeidae. Lesser amounts of
partially decomposed comminuted plant fragments as well as occasional large (850 ^m) fecal
pellets likely of Diptera larvae were also evident. Humus fecal pellets of 400-600 size, that
were smooth and usually quite dense, were probably casts of Diplopoda. The Ah horizon had a
mull-like fabric arrangement comprised of coalesced fecal pellets dominantly 50-90 /um size
and somewhat lesser amounts of units 180-240 /im in size. The units were usually relatively
| compact mull-like material, smooth in form and frequently lobate or round. However, some
| units had characteristics of fecal pellets of Collembola and Enchytraeidae (Rusek, 1974) and
i they were the most likely contributors to this fabric type. Occasional larger units present
! (350-600 /xm) were probably fecal pellets of Diplopoda. Fecal pellets of Collembola were also
observed in aggrotubules.

Quaest. Ent., 1985, 21 (4)

478

Pawluk

Boreal Forest Region (Table 1.4)

The form of upper moder humus within the organic layer comprised partially decomposed
plant fragments and fecal pellets of variable size. Fecal pellets (25 /um) of Acari were
associated with decomposing plant fragments. Most fecal pellets were relatively uniform, fairly
smooth in outline and of 90-125 /um size. There were also zones of units 50-60 /urn in size. The
variation in size distribution between zones of fabric may reflect the presence of different
species of Collembola. Relatively few irregular shaped, loosely structured units of 125-250
size probably were Enchytraeidae droppings while larger units of 600-750 /um size were likely
produced by Diptera larvae or small earthworms. Rare large earthworm casts (1.8 mm) were
found at the mineral surface contact. Fungal hyphae were abundant throughout the humus
layer. This layer showed sharp demarcation from the underlying mineral soil. Mixing of
organic and mineral material is minimal and likely reflects the general lack of larger fauna in
the population.

Transition Aspen Parkland Boreal Forest Region (Table 1.5)

Forms of moder humus contained an abundance of fecal pellets of highly variable size and
shape ranging from 35 to 950 /um in size as well as variable admixtures of partly decomposed
plant fragments and few mineral grain f-members. Humic fecal pellets, irregular in shape and
of 30-50 /um size, dominated, probably reflecting the presence of Collembola. Many well
rounded and lobate units of similar size were likely droppings of Acari. There were zones of
concentration of smooth round fecal pellets of 90-125 /um size that strongly resembled major
units believed to be dropping of other species of Collembola in many of the northern soils.
Other animal origin cannot be discounted, for example because of their similarity to droppings
reported for Isopoda (Kubikov and Rusek, 1976). There was a significant volume of smooth,
loosely bound fecal pellets of 350-600 /um size that comprised both humus and mull in the
upper layer and mull in the lower layer. These appear to be droppings of Diplopoda. A few
large (940 /um) fecal pellets of Diptera and/or small earthworms were also present. The lower
H layer was made up of partially coalesced fecal pellets dominantly 25 to 40 /um in size but also
contained a significant amount of fecal material in the 120 to 180 /um size range. A lesser
amount of fecal material 400-600 /um in size was also present. Most fecal material showed
evidence of breakdown that probably reflected aging of collected fecal casts from animals
similar to that active in the upper layer. The ‘zone of mixing’ that intergrades to the Ah had a
greater dominance of mull fecal pellets of 200-550 /um size typical of that for Diplopoda. Fine
mull fecal pellets of 90-125 /um size, very similar in structure to the humic units of the H layer
were also evident.

The Ah layer had greater dominance of larger fecal pellets of 600-750 /um size. In most
samples these fecal pellets comprised closely packed smaller units of 45 /um size some of which
were made up entirely of humus and others of mull-like material. However a minor portion of
the larger units frequently contained mineral matrix material brought up from the lower solum.
These fecal pellets closely resembled those of Diplopoda and it appears that they may have
played a very significant role in mixing organic and mineral materials in these soils. Large
earthworm fecal pellets were relatively rare, however small earthworm casts maybe confused
with those attributed to Diplopoda. Observed banded fabric reflects freeze-thaw processes but
whether these same processes contribute to formation of well developed microstructural units is
as yet uncertain.

Soil micromorphology and soil fauna

479

Prairie Parkland Region (Table 1.6)

Humigranic and mullgranic units within fabrics of the Ah horizon were largely fecal pellets
varying in size from 30 /um to 2 mm. Most of the finer (30-45 /xm) fecal pellets probably
reflected the presence of Acari and Collembola. Largely humic in composition, they were
concentrated in the upper zones of the horizon. Some humigranic units were also melanized
plant fragments. The larger mullgranic units (350-750 /um) were diverse in size and degree of
compaction and probably reflected the presence of a wide variety of fauna including Diplopoda,
Enchytraeidae, Isopoda, small earthworms and Diptera larvae. Contributions from large
earthworms (1-2 mm) were much less common and frequently consisted of matrix material
from underlying horizons. While the humus form was generally mull-like, the presence of
diverse, discrete, poorly homogenized units of soil material suggests an immature or ‘proto’
stage of mull development.

Alpine Region (Table 1.7)

Humigranic units in the moder layer appeared to be largely fecal pellets of Enchytraeidae
and microarthropods with a signficiant component of melanized comminuted plant fragments.
Very few large fecal pellets (450-550 /um), likely those of Diplopoda, were evident as well. The
Ah showed an increase in dominance of weak mull units of 20-400 /u m size that at least in part
reflected the activity of Collembola, Acari, and Enchytraeidae. Diplopoda were likely
responsible for the very rare large fecal pellets of 600 /um size. Well rounded compact fecal
pellets of 90-125 /um size were commonly present. While their origin is doubtful these casts may
be formed by specific species of Collembola. Biological influence at depth was more difficult to
discern because of the presence of orthogranic units of similar structure that reflected the
presence of amorphous constituents such as perlite and volcanic glass. The lack of well
developed mull probably reflects the low clay content.

Intermontane Prairie Region (Table 1.8)

The mull-like moder Ah comprised at least in part an abundance of fecal pellets of
microarthropods (35 /um), few Enchytraeidae (90-200 /um) and very few Diplopoda (450-600
um). As with most other soils dense, smooth droppings of 90-125 nm size were commonly
present. All fecal material appeared humic in composition under low magnification. Plant
fragments frequently had Acari fecal pellets within their decomposing structure. Frequently
mineral grains were observed to have organic cutans that probably resulted from deposition of
relatively mobile humic substances by wetting and drying and/or freezing and thawing
processes. Clay content was low and relatively ineffective in stabilizing humic material.

CONCLUSIONS

All soils from the Northern Tundra to the Parkland regions had microfabrics of humus-rich
layers that were considerably modified through faunal activity. The animals acted in several
ways. They were responsible for comminution of plant fragments and reorganization of humic
and fine mineral material into discrete microstructural units. In some instances their ingestion
appeared to enhance mull fabric development through formation of organo-clay complexes.
Their channels modified soil porosity and often remained filled with fecal material as
pedotubules.

Quaest. Ent., 1985,21 (4)

480

Pawluk

Among all the fauna present microarthropods appeared to be most ubiquitous in their
influences upon soil microstructures. Fecal pellets of Acari were usually associated with
partially decomposed plant fragments but those of Collembola appeared to be more broadly
distributed and of much greater abundance. Fecal pellets of Collembola were found at all sites
and dominated in the moder humus forms of the northern Tundra region and Intermontane
prairie. In mineral soils collembolan fecal pellets usually occupied voids or old root channels
and consisted of enclosing soil material which may have been humus, mull or mineral matrix
varying with the niche they occupied. Enchytraeidae were also widely distributed
geographically and not all of their fecal pellets were distinguishable from those of Collembola
(Hale, 1967). They contributed significantly to fabrics of Alpine, Parkland and
Forest-Parkland transition regions. Along with fecal pellets of other larger animals such as
Diplopoda and larvae, Enchytraeidae also contributed significantly to the formation of mull
fabrics found in the upper soil layers. Earthworms did not appear to play as dominant a role in
mull formation in these soils as they do elsewhere (Kubiena, 1953), even though small
earthworms are plentiful in some Parkland soils. In some samples it is difficult to distinguish
between casts of small earthworms and Diplopoda. Fecal pellets of Collembola and larger
animals incorporated organic and mineral constituents that vary in dominance with the degree
of soil mixing. Fecal pellets were observed to comprise pure humus, pure mineral matrix as well
as mull at different stages of formation. No organisms appeared to be capable of producing
mull through a single ingestion. Rather, large animals appeared to feed on fecal pellets of
smaller animals which in turn, ingested fecal material of the larger animals. At each stage the
fecal material served as substrate for the growth of microorganisms that were being harvested.
Repeated turnover of humus and mineral matrix material by the faunal community appeared
to enhance the rate of stable mull formation. Thus the synecology within the various soil
systems may be more important than individual species numbers in determining the degree of
and rate of mull formation.

It was difficult to assign humus types to the majority of the observed soils since various
humus forms were identified in each of the pedons albeit in different proportions. The observed
upper organic-rich layers at all sites have the properties described for moder (Kubiena, 1953)
humus form. However, a thin well developed mull humus layer was observed in the Southern
Tundra and Forest-Tundra Transition region. The Alpine and Montane Prairie sites had moder
humus forms although very weak mull fabric i.e. organo-clay complexing, was evident
especially in some of the larger fecal pellets. The lower humus layer of the Parkland-Boreal
Forest Transition soil had a humus form made up entirely of coalescing humic fecal pellets
ranging in size from 30-200 fim and had a mull-like arrangement i.e. a mull-like moder
(Kubiena, 1953) fabric. On the other hand, fecal pellets of variable size and largely comprising
mull material were present in association with mineral grains and partly decomposed plant
fragments as discrete units in moder-like arrangement in the upper Ah of the Parkland soil.
This humus-form has also been referred to as mull-like moder (Barratt, 1964) but may best be
regarded as a ‘proto’ mull formation since the soil materials were not completely homogenized.

The most serious problem we have in describing our soils arises from the need to assign a
genetic origin for the various fabric sequences. Observed fabrics for upper mineral layers of
Tundra soils strongly suggests that mull fabric arrangement (Kubiena’s spongy fabric) can
arise from processes other than faunal activity such as freezing and thawing. How significant
these contributions are to maintaining the tilth of our soils is largely unknown. A better
understanding of synecological and soil microfabric relationships is also required if we are to

Soil micromorphology and soil fauna

481

take maximum advantage of the natural processes within the soil ecosystem to sustain the

productivity of the resource base of our land.

REFERENCES

Babel, U. 1973. Die Verwendung humusmikromorphologischer Merkmale zur Untersuchung
standortskundlicher Fragen, pp. 223-240. In: Rutherford, G.K. (Editor). Soil Microscopy.
The Limestone Press, Kingston, Ont. 857 p.

Bal, L. 1973. Micromorphological analyses of soils. Paper No. 6. Soil Survey Institute
Wageningen, The Netherlands. 174 p.

Barratt, B.C. 1964. A classification of humus forms and micro-fabrics of temperate Grasslands.
J. of Soil Sci. 15: 342-356.

Brewer, R. 1964. Fabric and mineral analysis of soils. John Wiley & Sons, Inc., New York. 470
P-

Brewer, R. 1979. Relationship between particle size, fabric and other factors in some
Australian soils. Aust. J. Soil Res. 17: 29-41.

Brewer, R. and S. Pawluk. 1975. Investigations of some soils developed in hummocks of the
Canadian sub-Arctic and southern Arctic region. I. Morphology and micromorphology.
Can. J. Soil Sci. 55: 301-319.

Dudas, M.J. and S. Pawluk. 1969. Chernozem soils of the Alberta Parklands. Geoderma. 3:
19-36.

Fox, C.A. 1979. The soil micromorphology and genesis of the Turbic Cryosols from the
Mackenzie River Valley and Yukon Coastal Plain. Ph.D. Thesis. Univ. of Guelph, Ont. 196
P-

Hale, W.G. 1967. Collembola, pp. 397-409. In: Burges, A. and F. Raw (Editors). Soil Biology.
Academic Press. 536 p.

Howitt, R.W. and S. Pawluk 1984. Dynamics of a Gray Luvisol. Can. J. of Soil Sci. (in press).

Jacks, G.V. 1963. The biological nature of soil productivity. Soils Fertil. 26: 147-150.

Kubiena, W.L. 1938. Micropedology. Collegiate Press Inc. Ames, la., U.S.A.

Kubiena, W.L. 1953. The Soil of Europe. Thomas Murby and Co., London. 317 p.

Kubikova, J. and J. Rusek, 1976. Development of xerothermic rendzinas. Academia
Nakladatelstvi Ceskoslovenske Akademie Vecl. Praha. 78 p.

Martin, J.P. 1946. Microorganisms and soil aggregation II. Influences of bacterial
polysaccharides on soil structure. Soil Sci. 61: 157-166.

McLean, A. 1982. Guide to the Lac du Bois Grasslands, pp. 113-129. In: Nicolson, A.C., A.
McLean and T.E. Baker (Editors). Grassland Ecology and Classification Symposium
Proceedings, Ministry of Forests, B.C. 353 p.

Pawluk, S. and R. Brewer. 1975a. Micromorphological and analytical characteristics of some
soils from Devon and King Christian Islands, N.W.T. Can. J. Soil Sci. 55: 349-361.

Pawluk, S. and R. Brewer. 1975b. Investigations of some soils developed on hummocks of the
Canadian Sub-arctic and southern Arctic Regions. 2. Analytical characteristics, genesis and
classification. Can. J. Soil Sci. 55: 321-330.

Pawluk, S. and R. Brewer. 1975c. Micromorphological, mineralogical and chemical
characteristics of some Alpine soils and their genetic implications. Can. J. Soil Sci. 55:
415-437.

Post, F.A. and F.R. Dreibelbis, 1942. Some influence of frost penetration and

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microenvironment of water relationships of woodland, pasture and cultivated soils. Soil Sci.
Soc. Amer. Proc. 7: 95-105.

Rusek, J. 1974. Die bodenbildende funktion von Collembolen und Acarina. Pedobiologia Bd.
15:299-308.

Russell, E.W. 1973. Soil conditions and plant growth. Longmans, London. 849 p.

Sanborn, P. and S. Pawluk, 1983. Process studies of a Chernozemic pedon, Alberta (Canada).
Geoderma, 31: 205-238.

Satchell, J.E. 1967. Lumbricidae, pp. 259-318. In: Burges, A. and F. Raw (Editors). Soil
Biology, Academic Press. 536 p.

Zachariae, G. 1963. Was leisten Collembolen fur den Waldhumus? pp. 109-124. In: Doeksen,
J. and J. van der Drift (Editors). Soil Organisms. North-Holland, Amsterdam. 453 p.
Zachariae, G. 1964. Welche Bedeutung, haben Enchytraein im Waldboden? pp. 57-68. In:
Jongerius, J. (Editor). Soil Micromorphology. Elsevier, Amsterdam. 540 p.

Soil micromorphology and soil fauna

483

Table 1. Site Characteristics and Surface Soil Micromorphological Features in some Selected
Western and Northern Canadian Soils.

1. Northern Tundra Region

Location:

Landform:

Truelove Lowlands, Devon Island

a) Gravelly beach ridge deposits

b) well drained soils on ridge top

Vegetation:

Dry as integrefolia and Saxifrage opppositefolia are dominant
with some lichens on top of hummocks.

Soil Structure:

Surface layer is an Ahk horizon 10 cm thick with characteristics of
a moder humus form. The layer is black (10YR2/1, m) in color
and loamy sand in texture. The soil lacks structure and is very
friable. Very fine and fine roots are plentiful. The soil material is
base saturated with pH 7.2. Organic matter content is 9 percent.
The surface layer is underlain by a Bkj and Ck horizon. (For full
description see Pawluk and Brewer, 1975a).

Classification:
Microstructure of Surface
Layer:

Orthic Static Brunisol.

The moder Ahk horizon has dominantly an
ortho-humi-phytogranic fabric near the surface that grades to an
ortho-humigranic fabric with depth (Fig. la). Humigranic units
range from 20-80 /urn in size and are irregular in shape (Fig. lb).
Some loosely bound units of 200 fim size that comprise fine
humigranic, orthogranic and phytogranic units are also evident
(Fig. lc). Some spores and limestone nodules are present. Matrix
is low in clay size mineral components most of which is calcitic in

nature.

2. Southern Tundra Region

Location:

Landform:

Tuktoyaktuk, N.W.T.

a) Hummocks of slumped till developed on thermokarst knoll.

b) Well drained soil in midslope position.

Vegetation:

Betula glandulosa and Salix arctica with a dominance of
feather-mosses and lichens as ground cover.

Soil Structure:

The soil has a 2 cm thick root mat of raw humus with nonvascular
plants overlying a 7 cm thick humus layer. The upper moder
humus form is somewhat fibrous and matted comprising both
semi-decomposed and decomposed plant material. The underlying
3 cm layer has a mull-like moder humus form. The underlying 10
cm thick A(B) horizon is dark brown (10YR3/2, m) clay loam,
with friable moderate to strong granular structure. Very fine, fine
and medium roots are abundant. The soil is slightly acidic (pH 5)
and has 7 percent organic matter in the mineral layer. The A(B)
has characteristics of weak mull. (For full description see Brewer
and Pawluk, 1975; Pawluk and Brewer, 1975b).

I Classification:

Brunisolic Turbic Cryosol.

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Pawluk

Microstructure of Surface
Layer:

3. Forest-Tundra Transition
Region

Location:

Landform:

Vegetation:

Soil Structure:

Classification:

The upper moder humus form has an ortho-phyto-humigranic
fabric (Fig. 2a). The humigranic units range from 30 to 200 /im in
size with a strong distribution around 50 ^m. Occasional
humigranic units of 350 to 600 u m size are randomly distributed.
The lower moder layer has a thick band of mullgranic units
350-400 fim size (Fig. 2b) as well as larger humigranic units up to
900 /urn size. The underlying mull-like moder has a
phyto-mull-humigranoidic fabric type (Fig. 2c) with a dominance
of units of 90-100 /im size some of which are well rounded
although larger units (200 size) are still present. The A(B)
horizon has an upper zone of mullgranic and mullgranoidic fabric
with mull units showing strong bimodal size distribution in the
40-120 nm and 400-600 ^m (Fig. 2d) size ranges. Smaller units
frequently occur within the larger units (Fig. 2e). The lower zone
of the horizon has a moderately accommodated mull-matrigranic
fabric with well rounded units of 1-2 mm size (Fig. 2f).
Accommodation and coalescence increase with depth.

Inuvik, N.W.T.

a) Glacial till flutings

b) well drained site.

An open canopy of Picea mariana, P. glauca, Betula papyrifera
above an understory of Rosa sp., Salix sp., and Ledum
decumbens. The groundcover is dominantly Vaccinium vitis-idaea,
v. uliginosum, with various mosses and lichens.

The soil has an undecomposed leaf mat (LF) 8 cm thick overlying
a 3 cm thick largely decomposed moder H layer. Very fine, fine
and medium roots are abundant. The organic layer is underlain by
a weak mull or mull-like moder Ah horizon 3 cm thick, dark
brown (7.5YR3/2, m) in color, silty clay loam in texture and with
friable fine to medium granular structure. A 12 cm thick Bm
horizon lies below. The soil is moderately acidic (pH 4.5 - 5.2)
with 12.2 percent organic matter in the Ah. (For a complete
description see Brewer and Pawluk, 1975).

Brunisolic Turbic Cryosol.

Soil micromorphology and soil fauna

485

Microstructure of Surface
Layer:

4. Boreal Forest Region

Location:

Landform:

Vegetation:

Soil Structure:

Classification:
Microstructure of Surface
Layer:

The lower leaf mat is primarily humi-phytogranic grading to
ortho-phyto-humigranic in the H layer (Fig. 3a). Humigranic
units are variable in shape and 50-200 /urn in size. Fungal hyphae
are abundant. Some large (850 pm) humigranic units are also
present (Fig. 3b). The Ah horizon largely comprises a mullgranic
fabric that grades to a matrigranoidic fabric with depth (Fig. 3c).
Minor amounts of humigranic and phytogranic units and zones of
mullgranoidic fabric are also evident. Basic fabric units are 50-90
pm in size although some units of 180-240 /urn size are also present
(Fig. 3d). Many aggrotubules comprising units of the same size
and composition are present (Fig. 3e).

Breton, Alberta

a) Undulating ground moraine.

b) Well drained soil adjacent to stream channel.

Canopy of Populus tremuloides and Picea glauca with
groundcover comprising a mixture of various mosses, lichens and
grasses.

The surface organic layer comprises an undecomposed leaf litter
(L) 1 cm thick underlain by a loose to matted semi-decomposed
(F) layer 4 cm thick and a well decomposed matted humus (H)
layer 1.5 cm thick. Acidity ranges from pH 5 to 6. Very fine and
fine roots are plentiful and fungal hyphae are abundant. An Aeh
forms the transition to a platy grayish brown (10YR5/2, m)
eluviated Ae horizon. (For a complete description see Howitt and
Pawluk, 1984).

Orthic Gray Luvisol.

The F layer is dominantly phyto-humigranic fabric with partially
decomposed plant fragments and humigranic units of variable size
(Fig. 4a). Orthogranic f-members are rare to few. The H layer has
a few mullgranic units and orthogranic units are more prominent
as well. Phytogranic units diminish in importance. Fungal hyphae
dominate throughout (Fig. 4b). Fabric units are of variable size
(35 /um-1.2 mm) but show strong bimodal size distribution with
dominance in the 90-125 /urn size range and much fewer in the
600-750 pm size range (Figs. 4c and 4d). All units are largely of
the humigranic type but develop a weak mull character near the
lower boundary. There is a sharp separation to the underlying Aeh
horizon which has a weakly banded mull-matrigranoidic vughy
porphyric fabric type. Fine humigranic units occur only in the
aggrotubules. Small humigranic units are also frequently observed
in decomposing plant tissue (Fig. 4e) within all layers.

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486

Pawluk

5. Transition Aspen

Parkland Boreal Forest
Region

Location:

Landform:

Vegetation:

Soil Structure:

Classification:
Microstructure of Surface
Layer:

Ellerslie, Alberta.

a) Glacial Lake Edmonton Plain.

b) Imperfectly drained site.

Open stand of Populus balsamifera and P. tremuloides with a
well developed shrub layer of Cornus stolonifera, Rosa sp.,
Symphoricarpos albus and minor admixtures of other shrubs. The
herb layer comprises Rubus pubescens, Mitella nuda, Mertensia
paniculata, Cornus canadensis along with a wide variety of other
plants.

The surface layer comprises organic horizons 18 cm thick that are
characteristic of the moder humus type. The upper partially
decomposed litter (LF) is made up largely of aspen leaves. This
layer grades into a dark red brown (5YR2/2, m) well decomposed
loose, fluffy humus, below. The leaf litter has a near neutral pH
and abundant roots of varying size. The organic layer is underlain
by a mull-like Ah approximately 35 cm thick. The Ah is black
(10YR2/1, m) silty clay in texture, and strong granular in
structure. The soil is quite firm and also contains an abundance of
roots. Weak mottling is evident in the underlying B horizon. (For a
complete description see Sanborn and Pawluk, 1983).

Gleyed Black Chernozemic.

The F layer has mull-humiphytogranic fabric comprising
comminuted plant fragments and an abundance of humigranic
units ranging in size from 35 /urn to 950 pm. Units of 45 pm size
dominate but units of 250-950 pm size (Fig. 5a) are also common.
Large (400-600 pm) mullgranic units are concentrated in
horizonal zones that resemble mull-like moder (Fig. 5b). The
transition to the H layer shows a decrease in phytogranic units and
stronger coalescence of units of 45 pm size to form a
humigranoidic component. The H layer is characterized by
humigranoidic fabric of coalesced units dominantly 25-40 pm and
commonly 120-180 pm in size (Figs. 5c and 5d). Phytogranic and
orthogranic components are relatively few. The fabric grades into
a humi-mullgranoidic weakly banded fabric in the upper Ah
comprising moderately well accommodated, partially fused granic
units of variable size (Fig. 5e). Mullgranic fabric is better
developed at a depth of 3 cm but gives way to mullgranoidic
porphyric fabric with depth. Dominance of larger mullgranic units
generally increases with depth but size of units remains quite
variable.

6. Parkland Region

Soil micromorphology and soil fauna

487

Location:

Landform:

Vegetation:

Soil Structure:

Classification:
Microstructure of Surface
Layer:

7. Alpine Region

Hay Lakes, Alberta

a) Undulating ground moraine.

b) Well drained soil on upper knoll.

Open stand of Populus tremuloides with a dense ground cover of
grasses Festuca stipa, Koelaria and Poa. A wide variety of shrubs
and forbes with Rosa dominating are also evident.

The near neutral (pH 6.9) surface layer is a well developed black
(10YR2/1, m) mull Ah horizon approximately 75 cm thick. The
soil is loamy texture (22 percent clay), friable and strong granular.
Organic matter content is 9.2 percent. The underlying Bm horizon
contains numerous krotovinas and earthworm channels. (For more
detailed description see Dudas and Pawluk, 1969).

Orthic Black Chernozem.

Fabric of the Ah horizon is dominantly humi-mullgranic (Fig. 6a)
with phytogranic units more prominent near the surface and with
some orthogranic units throughout. Humigranic units are small in
size (30-60 pm) while mullgranic units vary from 250 pm to 2 mm
in size (Fig. 6b and 6c). There are occasional larger matrigranic
units as well.

Location:

Landform:

Vegetation:

Soil Structure:

Classification:
Microstructure of Surface
Layer:

Sunshine Basin, Banff National Park, Alberta

a) Saddle adjacent to ridge, comprising colluvium/sandstone

b) Moderately well drained site on rise of land.

Phyllodece glanderliflora and Antennaria lanata plant
communities comprising a variety of alpine grasses and forbes.

The upper layer is a very dark brown (10YR3/2, m) moder
grading to mull-like moder Ah horizon 1 1 cm thick, with silt loam
texture and very friable weak fine granular to amorphous
structure. Very fine and fine roots are abundant. The soil is
moderately acidic (pH 4.8) with approximately 18.9 percent
organic matter content. The horizon is underlain by a transition to
the Bm. (For full description see Pawluk and Brewer, 1975c).
Orthic Sombric Brunisol.

The moder humus layer has a humi-phytogranic fabric that grades
to phyto-humigranic fabric below. Humigranic units have a size
mode ranging from 20-180 pm (Fig. 7a). A few units of 550 pm
size are also evident (Fig. 7b). The humus layer is underlain by an
organic-rich Ah horizon that has humi-phyto-ortho-mullgranic
fabric. The mullgranic units range in size from 20-400 pm (Fig.

7c) and show only weak organo clay complexing. There is a
dominance of mullgranic units in the size range of 90-250 (Fig.
7d) in the upper zone but becomes better graded with depth. Glass
shards, phytoliths and diatoms are common.

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488

Pawluk

8. Intermontane Prairie
Region

Location:

Landform:

Vegetation:

Soil Structure:

Classification:
Microstructure of Surface
Layer:

Lac du Bois (above Kamloops) British Columbia.

a) Morainal drumlin.

b) Well drained.

Middle grassland comprising Agropyron spicatum, Koeleria
macranthe, Astragalus miser along with other grassland species.
The surface is a very dark gray to black (10YR3/1 [d]-2/l [m])
mull-like moder loam with weak fine granular to amorphous
structure and very friable consistence (a turf-like feel). This layer
is underlain by a Bm horizon. (For a complete description of site
see McLean, A. 1982).

Orthic Dark Brown Chernozem.

The fabric of the Ah is dominantly ortho-humigranic and
granoidic with a minor chlamydic component (Fig. 8a).
Humigranic units have a strong modal size in the 25 /xm range
(Fig. 8b) but few larger units 90-200 pm size and very few units of
450-600 /urn size are also present (Fig. 8c). Very few phytogranic
units are also found. Mullgranic units are notably absent although
under high magnification humigranic units appear to have a weak
mull-like character.

Soil micromorphology and soil fauna

489

Fig. 1. Fabrics of moder humus from an Orthic Static Brunisol, Devon Island, Northwest Territories, Canada, (a).
Ortho-humigranic fabric (x30m); largely fecal pellets of microarthropods, (b). Humigranic units comprising fecal pellets
20-80 jim size (x 1 50m) likely from collembolans and/or enchytraeids. (c). Loosely bound humigranic units 200 jim in size;
probably enchytraeid cast (I50xm).

Quaest. Ent., 1985,21 (4)

490

Pawluk

Fig. 2. Fabrics from the humus-rich layer of Brunisolic Turbic Cryosol, Tuktoyaktuk, Northwest Territories, Canada-
(a). Ortho-phyto-humigranic fabric (x30m); humigranic units show strong modal distribution of 50 urn and appear to be
largely droppings of collembolans and enchytraeids. (b). Large fecal pellets (400-600 /im) (x30m) of dipteraous larvae
and/or diplopods. (c). Phyto-mull-humigranoidic fabric (x30m); dominance of fecal material 90-100 /im in size
comprising humus and mineral constituents, believed to be droppings of collembolans. (d). Larger mullgranic units
(400-600 Mm) in the A(B) horizon (x30m). (e). Smaller mullgranic units (40-120 Mm) associated with larger units in the
A(B) horizon (x30m), probably fecal pellets of collembolans and/or enchytraeids. (f). Large mull-matrigranic units (1-2
mm) (x30m); possibly fecal pellets but more likely formed through frost processes.

Soil micromorphology and soil fauna

491

Fig. 3. Fabrics from the humus-rich layer of Brunisolic Turbic Cryosol, Inuvik, Northwest Territories, Canada.- (a).
Ortho-phyto-humigranic fabric of the H layer (x30m). Majority of humigranic units are 50 Mm in size and are likely fecal
pellets of collembolans; larger casts of 90-120 Mm size are likely those of enchytraeids. (b). Large fecal pellets (850 Mm) of
humic material (x30m) likely droppings of dipterous larvae, (c). Mullgranoidic fabric of the Ah horizon (x30m). Basic
units are 50-90 Mm size with some units 180-240 Mm size also evident. The smaller units are likely fecal pellets of
enchytraeids and/or collembolans. (d). Same as (c) (x50m). (e). Aggrotubule with fecal pellets (x50m).

Quaest. Ent., 1985,21 (4)

492

Pawluk

^ r*cs from the organic layer of an Orthic Gray Luvisol, Breton, Alberta, Canada - (a). Phyto-humigranic fabric
o t e ayer (x30m). Fecal pellets are dominantly 50 jam size possibly from collembolans. (b). Abundant fungal hyphae
m 'mate y associated with fecal pellets (50-60 jam) and plant fragments (xl50m). (c). Zone of humic fabric with a
ommance o ecal pellets 90-125 jim size (x30m). Regularity in the shape of the units suggests casts of collembolans
l x3(^U^ j! Cr an‘ma*s cannot be discounted, (d). Zone of humic fabric with a dominance of fecal pellets 600-750 jum size
m . eir presence likely reflects the activity of dipterous larvae and/or small earthworms, (e). Fecal pellets (25 jtm)
of acarmes in decomposing plant tissue (xl50m).

Soil micromorphology and soil fauna

493

Fig. 5. Fabrics from humus-rich layers of Gleyed Black Chernozem, Ellerslie, Alberta, Canada- (a).
Mull-humi-phytogranic fabric of the F layer (x30m). Small humigranic units (35 Mm) are fecal pellets of microarthropods,
the larger units (350-400 Mm) are likely fecal pellets of diplopods. (b). Mullgranic units (400-600 *im) in the upper H layer
are likely diplopod casts (x30m). (c). Humigranoidic fabric of the lower H layer (x30m). Units of fabric comprise entire
and decomposing small fecal pellets (25-40 nm) of small arthropods and larger fecal pellets (120-180 #xm) possibly of
isopods and/or enchytraeids. (d). Discrete and decomposing fecal pellets of collembolans and/or enchytraeids in c) under
high magnification (units 25-40 Mm size). (xl50m). (e). Humi-mullgranoidic fabric of the Ah with units dominantly
450-500 Mm size (x30m). Units are porous and appear to comprise smaller fecal pellets reorganized through frost
processes.

Quaest. Ent., 1985,21 (4)

Fig. 6. Fabrics of Ah horizon from an Orthic Black Chernozem, Hay Lakes, Alberta, Canada.- (a). Humt-mu lgramc
fabric from the Ah (,30m). Fecal pellets vary in size from 30 am to 400 am. Smaller untts are " - (b) Sma

humigranic and larger mullgranic units of fabric from the Ah (,50m) ppl. Note uneven dtstr, button of clay and humus
plasma, (c) Same as (b) in plain light.

Soil micromorphology and soil fauna

495

Fig. 7. Fabrics of humus-rich layers of Orthic Sombric Brunisol, Sunshine Basin, Alberta. Canada, (a).
Phyto-humigranic fabric in moder humus layer (x30m). Humus fecal pellets range in size from 20-180 jim and probably
reflect activity of collembolans and enchytraeids. (b). Fecal pellets comprising humic material and silt grains in same layer
as a). Large units (450-550 #xm) are probably casts of Diplopoda (x30m). (c). Humi-ortho-mullgranic fabric in Ah.
Mullgranic units range from 20-250 ^m (x30m). (d). Mullgranic units in Ah 90-250 jim in size (x30m) possibly
collembolans and/or enchytraeid fecal pellets.

Quaest. Ent., 1985, 21 (4)

Fig. 8. Fabrics of Ah horizons of Orthic Dark Brown Chernozem, Lac du Bois, British Columbia, Canada - (a).
Ortho-humigranic fabric with Chlamydic component in Ah (x30m). (b). Humigranic units dominantly 25 mhi size
(xl50m) probably fecal pellets of microarthropods, (c). Fecal pellets 450-600 nm size probably of diplopods (x30m).

SOIL MICROSTRUCTURES - CONTRIBUTIONS ON SPECIFIC SOIL ORGANISMS

J. Rusek

Laboratory of Soil biology
Institute of Landscape Ecology
Czechoslovak Academy of Sciences
C. Budejovice
CZECHOSLOVAKIA

Quaestiones Entomologicae
21:497-514 1985

ABSTRACT

The soil zoological approach to the soil micromorphological studies is described. The
ecological methods, e.g., synecological analysis of soil animal communities, succession of soil
animals, autecology, etc. are determined as basic methods for evaluating soil thin sections
from soil-zoological point of view. The role of soil animals in formation of soil microstructure
is divided into three basic categories: A) disintegration of dead organic matter, B) formation
of zoogenic microstructural soil matrix, and C) tunnelling and burrowing activities of soil
animals. The role of different groups of soil animals in disintegration of dead organic matter
is described and the characteristic features are documented on soil thin section figures. The
characteristic microstructural features of humus development during the succession are
described. The short term processes of decomposition ( disintegration ) in humus profile relate
to the long term development of humus form during the succession as does ontogeny to
phytogeny in the animal kingdom in HeckeTs biogenetical law. The droppings of different
groups of soil animals are described from the morphological point of view and their location
in the soil profile is given.

RESUME

L’auteur dicrit la mithode pido-zoologique d'etude de la micromorphologie des sols. Les mithodes icologiques,
telles que I'analyse synicologique des communitis animales des sols, la succession des animaux des sols, I'auticologie,
etc., sont considiries comme des methodes de base pour revaluation des coupes minces de sol d'un point de vue
pedo-zoologique. Le role des animaux des sols dans les processus de formation de la microstructure des sols se divise en
trois categories fondamentales: A) disintegration de la matiere organique morte, B) formation de la mat rice
microstructurale zooginique, et C) percement de tunnels et fouissage par les animaux endogis. L'auteur dicrit le role de
differents groupes d’animaux des sols dans la disintegration de la matiire organique morte et en prisente les traits
caractiristiques sur des figures de coupes minces de sols. II dicrit aussi les traits microstructuraux caractiristiques du
diveloppement de I’humus duranl la succession. Les processus de dicomposition (disintigration) d court terme ayant lieu
dans la couche d’humus sont en rapport avec le diveloppement d long terme du type d'humus au cours de la succession de
la meme faqon due I’ontoginie I’est d la phyloginie chez les animaux, tel que le repporte la loi bioginitique d Heckel.
Finalement, l’auteur dicrit les excriments de diffirents groupes d’animaux des sols d’un point de vue morphologique et
indique leur emplacement dans le profit du sol.

INTRODUCTION

Micromorphological methods of soil investigation were originated and developed by
Kubiena as a soil biological approach to pedological problems. They were used in soil biology in
the 40’s and 50’s by Kubiena (1943, 1948, 1955) and his collaborator Kiihnelt. Primarily

498

Rusek

through the contributions of pedologists, geologists, and to a lesser extent by the soil biologists,
method, theory and nomenclature of soil micromorphology were further developed in the 60’s
and 70’s, when soil micromorphology became an independent branch of pedology. In spite of
important publications using soil micromorphological methods, the number of soil biologists
using these methods has been and is still very low and does not reflect their present and future
importance. This international meeting of soil zoologists, soil micromorphologists and
pedologists is an important step in collaboration among specialists of these ecological branches.
Only by such an interdisciplinary collaboration is it possible to obtain new and untraditional
views on soil and on complicated, dynamic soil processes.

Soil micromorphology has already helped to solve some practical problems in soil biology. It
is possible to use it for monitoring man’s impact on the environment, for solving practical
questions connected with soil fertility and recultivation, for solving theoretical problems of soil
development, et cetera.

The literature about soil micromorphology contains some contradictory results. My
contribution summarises and discusses both my own and published results dealing with the role
of soil animal groups in forming soil microstructures. A wider examination of these problems
enables better understanding of ecological patterns in formation of microstructure and in soil
development generally. For this reason my contribution also includes soil micromorphological
methods from a soil zoological viewpoint. One part is devoted to the diagnosis of the tracks of
soil animal activities in the soil. Some unsolved or controversial questions will also be pointed
out, to stimulate work in some new directions.

My own results are from soils in the temperate, subpolar, alpine and mediterranean zones
and from some subtropical and tropical soils in Cuba.

METHODS

Methods of preparing thin sections of soil and their morphometric evaluation are well
described in the book edited by Jongerius (1964). Methods for evaluation of soil thin sections
from the soil zoological point of view are more complicated due to the difficulty of determining
the origin of the zoogenous microstructures.

To determine the zoogenous microstructural components of the soil we must start from the
coenological analysis of the zooedaphon in the soil under study, from the food requirements of
the dominant species of the soil meso-, macro- and megafauna, from the shape and size of
faecal pellets obtained in the laboratory, from newly captured animals, and from direct
observation of some dominant species in the field.

The coenological analysis of soil animal communities enable determination of the dominant
species in the soil under study. In the second step we identify the predators, phytophagous and
microphagous species which do not play a direct role in processes of soil microstructure
formation. For questionable taxa, it is necessary to analyse the gut content to establish the roles
of such species in forming the soil microstructure. It is important to point out that populations
of some soil animals have synchronised food consumption and that such animals ( e.g
Collembola) do not feed during certain life periods (ecdysis). For such animals, it is necessary
to analyse the gut content repeatedly. Some soil animals appear in high numbers only in some
parts of the year (cf. Rusek, 1984), also important to remember in evaluating the role of such
animals in the soil forming processes.

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499

Diptera

Chironomidae
Ceratopogonidae
Psychodidae
Coleoptera
Dryops rudolfi

□

Agriotes brevis

Fig. 1. Distribution of different groups of soil insects in a swampy meadow (a), wet meadow (b) and dry meadow (c) in a
periodically inundated area in South Moravia. Height of d - 1000 specimens, nr2. Larvae of some groups of Diptera and
Dryops rudolfi play an important role in processes that form soil microstructure in the swampy meadow.

To obtain droppings of soil animals known to affect processes of soil formation (gut filled
with brown or black particles of dead organic matter in different stages of disintegration and
mostly mixed up with mineral particles), immediately after extraction in the Tullgren
apparatus the animals are placed into glass jars with wet filter paper on the bottom. It enables
us to identify the zoogenous microstructures in soil thin sections with particular species of soil
animals. In a further phase, laboratory rearings of the species forming the soil microstructure
are carried out to prove their food requirements. In the future we will have enough experience
to limit this long procedure to the coenological analysis of particular groups of soil fauna and of
other dominant decomposers for which a role in the processes of formation of microstructure
are still not established. We may also obtain valuable results by direct observation of soil
animals in the field, as is pointed out by Kubiena (1964), Zacharie (1965), Bal (1970) and
others. But the importance of many of the soil animals cannot be established by field
observation.

SUCCESSION OF SOIL TYPES DURING SUCCESSION OF WHOLE ECOSYSTEMS

The coenological composition of soil fauna and the presence of different groups and species
of decomposers in the soil has a crucial effect on the rate and forms of disintegration of dead
organic matter. The composition of soil fauna coenoses, together with some other factors,
determines what humus form, soil microstructure and soil type will develop in the ecosystem. In

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my contribution (Rusek, 1978) the connections between successional development of plant
communities, soil animal communities and soil types were shown. Soil animals play an active
part in the development of soil and whole ecosystems. Soil animal communities are developing
and changing during succession, and in association with them the succession of humus forms
proceeds as well (Rusek, 1978).

Each soil type has its own, characteristic soil fauna (Fig. 1) a fact which enables us to use
soil animals for soil diagnostic purposes (Ghilarov, 1965). It is known that the most developed
humus form, the mull, is formed by earthworms. But only some ecological types of earthworms,
the endogeic and the anecic ones, form the mull. The epigeic type of earthworms form typical
moder. The zoogeographical distribution of earthworms plays an important role in mull
distribution. Mull cannot be formed in areas where the aceic and endogeic earthworms are
missing; such areas include the Arctic and parts of the boreal zones, as well as initial and little
developed soils. In these areas or soil types, only less developed forms of humus ( e.g ., raw
humus, microarthropod moder, arthropod moder, etc.) occur. These ecological and
zoogeographical rules and dependences in soil and humus development must be kept in mind in
evaluating thin sections of soil.

During soil succession many important changes in composition of species and of ecological
groups of soil fauna occur. In the first developmental stages usually only microarthropods (
Collembola, Acarina) play an important role in processes of formation of soil microstructure,
and the microarthropod moder is formed by them, is formed by them. Some soils reach only
this developmental stage as a climax. These soils occur most commonly in the Arctic and in the
alpine zones. In temperate, subtropic and tropic zones the humus develops to more complex
forms, and determination of its micromorphological components is more difficult because of the
great diversity of soil animals taking part in its development.

ROLE OF SOIL ANIMALS IN PROCESSES OF FORMATION OF SOIL
MICROSTRUCTURE

We may divide processes of soil microstructure formation into three basic categories from
the soil zoological point of view:

(a) disintegration of dead organic matter

(b) formation of zoogenic microstructural soil matrix, and

Disintegration of dead organic matter

The main source of dead organic matter used in processes of formation of the zoogenous soil
microstructure is the plant litter. Before disintegration, the litter is intensively invaded by soil
microflora and soil microfauna, and only after a certain period is it attacked by larger soil
animals and disintegrated step by step. Some species of microarthropods (Collembola,
Oribatei) and enchytraeids skeletonize the leaves between the veins only, causing Fensterfrass,
whereas larvae of some Mycetophilidae, Lycoridae and other Nematocera also eat the thinner
ribs, causing Lochfrass (Figs. 2, 3). Diplopods ( Glomeris spp., Julus spp., etc., Marcyzzi,
1970), isopods and some earthworms bite off larger pieces of leaf tissue together with the
thinner ribs. The large midrib of oak leaves is mined by Rhisotritia minima (Berlese Oribatei)
[after Bal, (1968, in Harding and Stuttard, 1974 and Bal, 1970)].

The litter of conifers is disintegrated more slowly and with more difficulty. Parenchymatic
tissues are eaten by phthiracarid mites (Oribatei) which leave typical droppings in the bitten

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501

Fig. 2. Oak leaf partly disintegrated by larvae of Mycetophilidac. Their pellets (on the left side) invaded by nematodes.
Fig. 3. Mycetophilid larvae disintegrating leaf litter.

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off hollows inside the needles. The needles in the litter layer are also disintegrated by larvae of
Tipulidae and Mycetophilidae, caterpillars of Adela spp. (Bal, 1970) and by some earthworms
(Zachariae, 1966).

Wood disintegration has been studied by many authors, but we only have little data about
the zoogenous micromorphological processes in rotting wood (cf. Babel, 1975). Fallen twigs and
dead roots are invaded by phthiracarid mites which feed on the rotten wood. In thin sections of
soil, we can see large hollows filled by ovoid droppings of these animals (Fig. 4). Often, the
periphery of the hollow near the bark is covered by droppings of bark beetles (Scolytidae)
secondarily eaten by phthiracarids (Fig. 5). I have observed a few invasions of Collembola
( Mesaphorura spp.) in the wide, opened hollow of twigs. In such openings, the phthiracarid
pellets are mixed with dark Collembolan droppings containing small mineral particles (Fig. 4).

The wood of the tree stumps and logs is disintegrated first by xylophagous larvae and some
adults of beetles such as Scolytidae, Curculionidae, Buprestidae and Cerambycidae. When the
rotting processes have been advanced, the wood is attacked by larvae of Nematocera
(Tipulidae, Mycetophilidae, Lycoridae, etc.) and some Lucanidae, Cetoniidae and Dynastidae.
The tracks and pellets of these animals are of typical shape, composition and size, but the
micromorphological diagnostic characters have not yet been described. After some years a
typical soil fauna invades the rotten wood. The pellets of the xylophages are then disintegrated
and mixed step by step with mineral particles. Microarthropods, enchytraeids,
macroarthropods and some earthworms contribute to this process. In subtropical mountain rain
forests in Cuba, larvae and adults of Passalus sp. (Coleoptera: Passalidae) play an important
role in wood disintegration. After passing through the gut the pieces of wood in the excrement
are invaded by special microflora and the droppings are then again eaten by larvae of the same
species or other xylophagous animals. In the tunnels of these animals it is easy to distinguish
light coloured pellets after the first passage through the gut and the brownish or black ones
which passed through two or more times. These droppings then become a food source for a
diversified community of soil animals of different size.

Zoogenic formation of the soil matrix microstructure

Through the feeding activity of soil animals, the plant litter is disintegrated and converted
into new structures which may be stable or which are further converted by aging or feeding
activities of other soil animals, into other microstructures characteristic of the soil matrix. Like
the successional development of soil types, the development of humus forms also occurs in
successive steps. This successive development has its own regularities connected with the
ecology of soil animals from the decomposer ecological group. We may follow the progressive
development of humus forms during litter disintegration and during incorporation of the new
microstructural elements into the soil matrix. These short term decomposition processes in the
humus profile relate to the long term humus form development during succession as does
ontogeny to phylogeny in the animal kingdom in Haeckel’s biogenetical law. Also this short
term development of humus forms in the soil profile has its own regularities which may be
observed in, for example, forest soils.

The simplest humus form in xeric succession is the microarthropod moder formed by
droppings of Collembola, Oribatei and some small nematoceran-larvae ( Diptera) (Rusek,
1978). The next developmental step is the arthropod moder formed mainly by the larger
animals belonging to the group of macrofauna (Diplopoda, Isopoda, larvae of Diptera and
Coleoptera), by the enchytraeids and by the small epigeic forms of earthworms living in the

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503

litter ( e.g ., Dendrobaena rubida, Eisenia foetida, etc.). The arthropod moder is
microstructurally heterogenous in comparison with the microarthropod moder. The mull-like
moder has more complicated structure. The highest form of humus is mull and it develops only
when succession reaches a level at which conditions enable high densities of anecic and
endogeic earthworms.

Litter disintegration in a forest soil usually starts with the feeding activity of Collembola,
Oribatei and small larvae of Diptera. Their pellets belong to the microarthropod moder and
they are readily distinguishable in the uppermost litter layer (Fig. 6). “Later” and deeper in the
same litter layer are the larger pellets of enchytraeids, diplopods, larger Diptera larvae, etc.
belonging to the moder. Microarthropod droppings are also formed in this layer, but they are
almost completely comsumed by the macroarthropods and incorporated into their faecal pellets.
In some larger pellets they are easily visible in soil thin sections (Fig. 8). The macroarthropod
droppings can be secondarily disintegrated by aging into the small, original pellets of
microarthropods (Fig. 8). When anecic and endogeic earthworms are present in the soil in high
densities, the picture of processes of formation of arthropod moder can be completely obscured
by the mull production of these lumbricids. The droppings of macro- and microarthropods are
then totally disintegrated and mixed with mineral particles in the guts of these animals (Fig. 7).

The nomenclature of humus forms relates to the whole humus profile; the name of the
humus form is derived from the prevailing microstructural elements in the profile. It is difficult
to decide what humus form occurs in many samples of certain soil profiles. It is proposed here
to identify the humus form in each subhorizon of the L- and F- horizons.

Disintegration of larger droppings by the soil mesofauna (enchytraeids, Collembola, etc.) is
of great importance in the microstructural forming processes. It is most remarkable in the
upper part of the H-horizon, where the large, spongeous droppings of earthworms are eaten by
enchytraids and transferred into their small droppings (Zacharie, 1965). The same is true for
some Collembola (e.g., Onychiurus spp., Tullbergiinae gen. spp.,) ( Folsomia spp.) (Fig. 8). The
macroarthropod droppings in the F-horizon are disintegrated in the same manner as described
by Zachariae (1964) for enchytraeids and by Dunger (1983) for Collembola.

Tunnelling and burrowing activities of soil animals

Only soil animals from decomposer ecological groups play a part in the above described
disintegration processes. Tunnelling and burrowing are also done by animals from other
ecological groups such as herbivores, predators, etc. These have usually well sclerotized bodies,
strong mandibulae, head capsules and urgomphi, feet adapted to life in soil, well developed
muscles, worm-like shapes, etc. They aerate the soil profile by their tunnelling activities when
moving through the soil in search of food, reproductive or hibernation sites, etc. In autumn, the
wireworms (Elateridae) injurious to cultivated plants, move 50-60 cm deep to hibernate. They
migrate to the soil surface the following spring during which time they make long horizontal
channels in the uppermost part of the soil, searching for the roots of the host plants (cf. Rusek,
1972). There is a great diversity of actively tunnelling soil animals within the soil macro- and
megafauna, but some species of the soil mesofauna may also make active microtunnels in the
soil matrix (e.g., some Collembola from the family Onychiuridae, oribatid mites, enchytraeids).
Many of the tunnellers mix the organic matter with mineral particles and translocate the
droppings between soil horizons up to the soil surface. During such activities the organic matter
may be translocated by earthworms deep into the mineral horizon and the mineral soil
components transported to the surface. Soil material is translocated by earthworms, especially

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Fig. 4. Soil thin section from a moder rendzina. Pellets of oribatid mites (o) and Collembola (c) inside the small twig, and
droppings of enchytraeids (e). Fig. 5. Disintegration of twigs in a moder, by bark beetles and oribatid mites: droppings of
these animals are indicated by (s) and (o) respectively.

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505

Fig. 6. Oak leaves (a) partly disintegrated by Collembola (0. Enchytraeidae (b), oribatid mites (c). Below the leaves
droppings of nematoceran larvae (Diptera) (d) and of the epigeic earthworm Dendrobaena rub, da (r). Modcr rendzina
Bohemian Karst. Fig. 7. Spongeous droppings of an cndogeic earthworm, mull. Fig. 8. Droppings of epigeic earthworms
subsequently disintegrated by enchytraeids (e) and Collembola (c).

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the anecic ones, and also by groups such as ants and some other Hymenoptera, termites dung
beetles, some crickets and other insects. The cast-forming activity of anecic earthworms, ants
and another animals is well known and of great importance in microstructural and soil forming
processes. The zoogenous microstructural cavity system has been analysed in soil thin sections
only by a few workers ( e.g ., Babel and LeNgoc, 1977) and deserves more attention in the
future. It is easier to analyse these activities using thick soil sections or on ground block sections
than using thin sections of soil.

Because of their macromorphological impact, the burrowing and tunnelling activities of
vertebrates were not mentioned in connection with soil microstructure processes. The activities
of some groups of the invertebrate soil macro- and megafauna also extend to a
macromorphological level during soil succession.

STRUCTURE OF ANIMAL DROPPINGS IN THE SOIL MATRIX

The most important contribution of soil animals to formation of the microstructural fabric is
their excrement, also called droppings, pellets or faecal pellets. These are the prevailing
primary aggregates of many humus horizons. Each group or even species of soil animal
produces droppings of characteristic shape, composition, size and colour (cf. Bal, 1973).
Location and accumulation within the soil profile are also important features aiding in the
determination of the origin of droppings. Some droppings are very stable for a long time,
especially in rendzina soils, but usually they change with age or through feeding activities of
secondary decomposers. Many taxonomically different groups of soil animals produce similar
pellets, which consequently may be misinterpreted in soil thin sections. As was stated in the
methodological section, the determination of droppings must start from the coenotical analysis
of the soil fauna. The diagnostic features of droppings of the most important soil forming
animals groups are described below. The droppings of well known groups as well as groups for
which we lack information are described briefly.

Droppings of Oribatei

Pellets of oribatid mites (Oribatei, Acarina) are very distinctive and, in most, are easily
recognizable microstructures in the soil matrix. Their characteristics have been described by
many authors (e.g., Zachariae, 1965; Bal, 1970; Rusek, 1975): egg-shaped or sphaeric, with
smooth surface, very compact and without mineral particles inside, light brown coloured and up
to 200 x 140 Mm in size, depending on the species and the instar of the mite. Most characteristic
are the smooth surface and the missing mineral particles. These two characters are conditioned
by the structure of these mites. They have a very narrow pharynx through which mineral
particles and larger pieces of food cannot enter the gut (Taraman, 1968). In the ventriculus the
ball of food particles is covered by a thick peritrophic membrane (Fig. 9) which also covers the
droppings, giving them their smooth surface. Taraman (1968) mentions that the faecal pellets
of some oribatid species are grayish or black; their colour may shift from yellow to black during
aging, due to the action of microorganisms. The same author notes that pellets of Steganacarus
magnus fed on wood tissue do not have a smooth surface. These facts may explain why oribatid
excrements were not recognized in soil thin sections from places where the macrohumiphagous
Oribatei live in high densities.

The oribatid droppings are usually found in groups between the leaves in the L-layer, inside
coniferous needles (Fig. 10) or in feeding cavities in rotten wood (Fig. 4). Often groups of

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507

droppings of different size are together in one hollow, indicating the moulting cycle of the
feeding animal. Macrohumiphagous species from the oribatid family Phthiracaridae are most
important in the processes of formation of soil microstructure, but we may also find species in
other families contributing to these processes. Oribatid mites occur in all soil horizons.

Droppings of Collembola

Collembola are one of the most abundant representatives of soil mesofauna. They belong
together with Acarina and some smaller groups of Tracheata to the group of microarthropods.
Quite contradictory data have been published about the importance of Collembola in processes
of formation of soil microstructure. Zachariae (1963) is of the opinion that Collembola do not
play an important role in the disintegration of organic matter and in the processes of formation
of soil microstructure. But previously Kubiena ( e.g ., 1955) has pointed out the leading role of
Collembola in forming some mountain soils (e.g., pitchmoder rendzina). Also Bal (1970),
Dunger (1983) and other authors have shown the importance of Collembola in litter
disintegration. Kubikova and Rusek (1976) have established that in a xeric protorendzina
profile the droppings of Collembola predominate. Rusek (1975) describes the pellets of
Collembola, Oribatei and Enchytraeidae and the differences between them.

The droppings of Collembola are usually compact, 30-90 pm in diameter (over 100 /urn in
larger species), irregularly round, with rugged, irregular surface, usually containing mineral
particles, and usually black. The remains of organic matter inside them do not contain larger
parts of plant tissue. They clearly differ from the smooth, egg-shaped and light brown oribatid
droppings.

Collembola are one of the most ecologically diversified groups of arthropods and this fact
has given rise to a lack of understanding of their function in the soil by some authors.

There are Collembola living atmobiotically on higher plants and some of them are even
important pests (Fig. 11) (e.g., Sminthurus viridis). The epigeic forms are living on the soil
surface and in litter and some species are important litter decomposers (e.g., Tomocerus spp.
(Fig. 12), Orchesella spp. (Fig. 13), Isotoma spp., Hypogastrura spp.). The hemiedaphic
species live in the litter and F-horizon, whereas the euedaphic ones are in the F- and
H-horizons. Some of the hemiedaphic as well as euedaphic species contribute to the processes of
formation of soil microstructure (e.g., Folsomia spp., Onychiurus spp., Mesaphorura spp.,
Megalothorax minimus , etc.).

In the Collembola strongly developed feeding specialization exists. We recognize
Collembola with sucking mouth parts (Neanura spp., Micranurida spp., etc.), predators
(Friesea spp., Cephalotoma grandiceps), fungivores (Pseudosinella spp., Paratullbergia
callipygos, etc.), macrophytophages (Sminthurus spp., Bourletiella spp. (Fig. 11), etc.),
detritivores and other specialists. It is no wonder that any one who observed a phytophagous or
microhumiphagous species arrives at wrong conclusions about the roles of Collembola in soil
forming processes.

I am not implying that all species of Collembola play a role in soil formation. The same
situation occurs in almost all groups of soil arthropods, and we must always distinguish
ecological groups.

As already mentioned, the Collembola have a leading role in forming the soil microstructure
in some arctic, alpine and weakly developed soils. Sometime the whole soil profile of these soils
is formed primarily by collembolan droppings (Fig. 14). In more developed soils Collembola
take part in disintegration of leaf litter (Fig. 6) and in secondary disintegration of macro- and

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Fig. 9. Hypochtonius sp. (Oribatei), v-ventriculus; p-peritrophic membrane with a ball of food particles inside. Semi-thin
section prepared by Smrz. Fig. 10. Abies alba needles disintegrated inside by phthiracarid mites (Oribatei). Their pellets
are of typical shape (o). Fig. 11. Bourletiella lutea (Collembola) feeding on living plant tissues does not contribute to the
soil microstructures.

Soil microstructures

509

rig. 12. Tomocerus minor, an epigeic species of Collembola.- Feeding on the leaves in the L-layer contributes to the soil
microstructures by its small cylindrical pellets (a group of them on the right side). Fig. 1 3. Orchesella cincta (Collembola)
Lakes part in leaf litter disintegration. Fig. 14. Pellets of Collembola predominate in some alpine soil types. SchneetSlchen
rendzina, West Tatra mountains.

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megafauna droppings (Fig. 8). The small collembolan pellets can be found in lumbricid
channels, as well as inside their large, spongeous excrements in which the Collembola bite
narrow hollows and channels. Collembolan droppings are often confused with pellets of
enchytraeids.

Droppings of Enchytraeidae

The enchytraeids are intermediate in body length between the soil mesofauna and
macrofauna. In the size of their excrements, they are close to the mesofauna. The droppings are
described in many papers ( e.g ., Zachariae, 1964; Babel, 1968; Rusek, 1975). The pellets of
some enchytraeid species resemble those of Collembola; in other species, they differ distinctly
from the collembolan ones in their shape, size, and arrangement and location in the soil profile.
They are the leading microstructural components in some soils (Babel, 1968). In the lower
L-layer, the enchytraeid pellets are often in two parallel rows, forming channels between the
leaves (Zachariae, 1964). In the F- and H-horizons enchytraeids are secondary decomposers of
the larger excrements of soil macrofauna (Fig. 8). They make narrow channels in the large,
spongeous earthworm excrements (Zachariae, 1964).

The enchytraeid droppings are 120-200 nm long, of extremely irregular shape, and with
irregular surface. They contain mineral particles and pieces of plant tissues and many are
divided into primary components (pellets of microarthropods, plant and mineral particles). In
the deeper soil horizons most contain mineral particles larger than the collembolan pellets, and
the collembolan pellets are smaller than are those of the enchytraeids.

Droppings of Diptera, Coleoptera and larvae of other insects

Larvae of Diptera belong partly to the soil mesofauna (Lycoridae, Mycetophilidae,
Chironomidae etc., and partly to the macrofauna (Tipulidae, Bibionidae, etc.). Most form
sphaerical, cylindrical or spindle-like droppings belonging to the moder humus form. They
contain large pieces of plant tissues mixed sparsely with mineral particles, which may
sometimes be missing. Their length ranges from 100 jam to 1 mm (Bibionidae, Lycoridae,
Mycoridae, Mycetophilidae, and even more (Tipulidae). The droppings of Bibionidae are well
described by Szabo et al. (1967). They contain leaf residues, some algal filaments, structureless
organic substances and mineral particles, and they reach 0.3 to 0.4 mm in diameter and are up
to 1 mm long. The droppings of Tipulidae larvae (Fig. 15) are egg-shaped and contain large
pieces of plant material mixed in many cases with large mineral particles. Their surface is
covered by a peritrophic membrane. They are concentrated in the litter layer and some species
also are in the uppermost H-horizon. The droppings of litter feeding mycetophilid larvae are
concentrated in the L-layer. They are sphaerical and contain small pieces of leaf tissues,
70-200 jam in diameter and are not very stable.

Also some larvae of Coleoptera contribute to the soil microstructures. Droppings of some
groups are sphaerical, some more than 5 mm in diameter ( Melolontha spp.); others resemble
large droppings of Enchytraeidae, e.g., those of Dryops rudolfi (Figs. 16, 17). The droppings of
the last species are an important part of the microstructure in the temporarily inundated soils in
South Moravia (Rusek, 1973, 1984).

Bal (1970) described droppings of Adela sp. caterpillars disintegrating conferous needles.
They are cylindrical, solid, often contracted in the middle, with pieces of plant tissues. Their
size reaches 510 x 260 jam and they are deposited in small groups.

Soil microstructures

511

Fig. 15. Diplopod (d) and tipulid (t) larvae pellets in a moder rendzina in Bohemian Karst. Fig. 16. Dryops rudolfi larva
(Coleoptera: Dryopidae) from a periodically flooded swampy meadow in south Moravia. Gut filled with black particles of
dead organic matter and mineral particles.

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Fig. 17. Droppings of Dryops rudolfi larvae with residue of disintegrated leaves of Glyceria maxima in L-layer. Fig. 18.
Droppings of diplopods (a) in a mull-like moder rendzina in Bohemian Karst.

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513

Droppings of Diplopoda and Isopoda

The droppings of litter-consuming diplopods are characteristic microstructural elements in
many soil types. They are, for example, the dominant droppings in moder rendzina and in the
upper part of mull-like rendzina (Kubikova and Rusek, 1976). The diplopod droppings range in
size from 0.5 to 4 mm. Many contain large pieces of litter fragments, droppings of smaller soil
animals and many also a great quantity of mineral particles. The internal structure is not very
compact (Fig. 18). In most species, the droppings are covered with a peritrophic membrane.
The droppings of large julids and glomerids are egg-shaped or sphaerical; those of small julids
are elongate (Babel, 1975).

Droppings of isopods, which also consume litter are very similar. These droppings are
relatively rare in thin sections of soil due to the special ecological requirements of most isopod
species. The animals may be slightly more abundant in very small specific areas. The size and
internal structure of their pellets is almost the same as in diplopods. They are cylindrical and
some have a longitudinal cleft.

Droppings of Lumbricidae

The microstructures of lumbricid droppings are well known, and they are described in
almost all contributions dealing with the role of soil fauna in the formation of soil
microstructure ( e.g Kubiena, 1958; Zachariae, 1965; Babel, 1975). The structure, size and
internal composition of lumbricid droppings depend on the ecological group which produces
them. The epigeic forms produce microstructures belonging to the moder form of humus, the
endogeic and anecic ones produce mull-like or mull excrements. The epigeic group usually
produces cylindrical or irregular droppings containing plant material of different stages of
degradation (brown to black in colour) mixed with some mineral particles ( Dendrobaena
rubida, Eisenia foetida, Eisenia lucens, etc.) (Fig. 6). The droppings of endogeic and anecic
groups are spongeous, with very small pieces of organic matter well mixed with a great quantity
of mineral particles of variable size (Fig. 7). Most of these droppings are usually subsequently
disintegrated by the mesofauna (Fig. 8). They may occupy the whole humus horizon and the
upper parts of the mineral horizon, and within lumbricid channels they may extend deep into
the C-horizon.

REFERENCES

Babel, U. 1968. Enchytraeen-Losungsgefiige im Loss. Geoderma, 2: 37-63.

Babel, U. 1975. Micromorphology of soil organic mattter. pp. 369-473. In: Gieseking J E.
(Editor). Soil Components. Volume 1. Organic Components. Springer Verlag, New York,
Heidelberg, Berlin.

Babel, U. and B. LeNgoc. 1977. Regradierung von Mergelboden nach Schafweidenutzung.
Mitt. Dtsch. Bodenkundl. Gesellsch., 25: 313-320.

| Bal, L. 1970. Morphological investigation in two moderhumus profiles and the role of the soil
fauna in their genesis. Geoderma, 4: 5-35.

: Bal, L. 1973. Micromorphological analysis of soils. Lower levels in the organization of organic
soil materials. Soil Surv. Inst., Wageningen, 174 pp.

Dunger, W. 1983. Tiere im Boden. Die Neue Brehm-Biicherei. A. Ziemsen Verlag-Wittenbcrg
Lutherstadt, 280 pp.

| Ghilarov, M.S. 1965. Zoological methods in soil diagnostics. Publ. Office “Nauka . Moscow,

K Quaest. Ent., 1985, 21 (4)

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279 pp.

Harding, D.J.L. and R.A. Stuttard. 1974. Microarthropods, pp. 489-532. In: Dickinson, C.H.
and G.J.F. Pugh (Editors). Biology of plant litter decomposition. Academic press, London
and New York, Volume 2.

Jongerius, A. (Editor). 1964: Soil micromorphology. Elsevier, Amsterdam, 540 pp.

Kubiena, W.L. 1943. L’investigation microscopique de l’humus. Z. Weltforschirt., 10:
387-410.

Kubiena, W.L. 1948. Entwicklungslehre des Bodens. Springer-Verlag, Wien.

Kubiena, W.L. 1955. Animal activity in soils as a decisive factor in establishment of humus
forms, pp. 73-82. In: Kevan, D.K.McE. (Editor). Soil zoology. Butterworths Sci. Publ.,
London.

Kubiena, W.L. 1964. The role and mission of micromorphology and microscopic biology in
modern soil science, pp. 1-13. In: Jongerius, A. (Editor). Soil micromorphology. Elsevier,
Amsterdam.

Kubikova, J. and J. Rusek. 1976. Development of xerothermic rendzinas. A study in ecology
and soil microstructure. Rozpravy CSAV, Academia, Praha, 79 pp. + 16 pits.

Marcuzzi, G. 1970. Experimental observation on the role of Glomeris spp. (Myriapoda
Diplopoda) in the process of humification of litter. Pedobiol. 10: 401-406.

Rusek, J. 1972. Die mitteleuropaischen Agriotes- und Ectinus- Arten (Coleoptera, Elateridae),
mit besonderer Beriicksichtigung von A. brevis un den in Feldkulturen lebenden Arten.
Rozpravy Ceskosl. akad. ved, Academia, Praha. 90 pp. + 3 Tab.

Rusek, J. 1973. Dryops rudolfi sp.n. und seine Larve (Coleoptera, Dryopidae). Acta Ent.
Bohemoslov., 70: 86-97.

Rusek, J. 1975. Diebodenbildende Funktion von Collembolen und Acarina. Pedobiologia, 15:
299-308.

Rusek, J. 1978. Pedozootische Sukzessionen wahrend der Entwicklung von Okosystemen.
Pedobiologia, 18: 426-433.

Rusek, J. 1984. Zur Bodenfauna in drei Typen von Uberschwemmungswiesen in Siid-Mahren.

Rozpravy CSAV, Academia Praha, 128 pp. + 8 tbls.

Szabo, J., T. Bartlay, and M. Marton. 1967. The role and importance of the larvae of St.
Mark’s fly in the formation of a rendzina soil, pp. 475-489. In: Graft, O. and J.E. Stachell.
(Editors). Progress in Soil biology. Braunschweig and Amsterdam.

Taraman, K. 1968. Anatomy, histology of oribatid gut and their digestion. Bioloski Vestnik, 16:
67-76.

Zachariae, G. 1963. Was leisten Collembolen fur den Waldhumus? pp. 109-124. In: Doeksen,
J. and J. van der Drift. (Editors). Soil organisms. North Holland Publishing Comp.,
Amsterdam.

Zachariae, G. 1964. Welche Bedeutung haben Enchytraeen im Waldboden? pp. 57-68. In:
Jongerius, A. Soil micromorphology. Elsevier Publ. Comp., Amsterdam.

Zachariae, G. 1965. Spuren tierischer Tatigkeit im Boden des Buchenwaldes. Forstwiss.
Forschungen. Heft 20: 1-68.

Zachariae, G. 1966. Die Streuzersetzung im Kohlgartengeibiet, pp. 490-506. In: Graft, O. and
J.E. Satchell. (Editors). Progress in Soil biology. Braunschweig and Amsterdam.

SOME IMPACTS OF FUNGAL-FAUNAL INTERACTIONS IN SOIL

Dennis Parkinson

Kananaskis Centre for Environmental Research
The University of Calgary

Calgary, Alberta T2N 1N4 Quaestiones Entomologicae

CANADA 21:515-516 1985

Soils support complex, heterogeneously distributed communities of soil animals and soil
microorganisms. While accurate methods are available for comprehensive qualitative studies of
the diverse taxa of the soil fauna, such work is impossible for the soil microflora (in spite of the
fact that much attention has been given to microbiological methodology). Nevertheless,
available information ( e.g ., Satchell, 1971; Persson et al. , 1980) indicates that, at least in
temperate ecosystems, the microflora (and fungi in particular) have much greater biomass and
contribution to total respiratory metabolism (Reichle, 1977) in soil and litter than do the fauna.

Decomposer activity of the microflora in organic debris is determined by such factors as
temperature, moisture regimes, resource quality and quantity, and by the inoculum potential
and competitive abilities of the decomposer organisms (Visser, 1985). But activities of the litter
fauna (litter grazers, microbivores, detritivores and predators) must be superimposed on those
of the microflora and affect:

1 . the community structure of the litter microflora.

2. the patterns of decomposition of organic matter.

3. the retention and release of nutrients (attendant on organic matter
decomposition).

These effects are brought about in three major ways (Visser, 1985) i.e.

1. by comminution, mixing and channelling of litter and soil: this not only
causes increased surface area for microbial colonization but also can lead
to a decrease in species richness of fungi and a diversion of their
“uccessional patterns.”

2. grazing on the microflora: while many studies have indicated that faunal
grazing removes only a small proportion of the microbial biomass, selective
grazing (when it occurs) can affect microbial community structure and,
possibly, organic matter decomposition rates. Grazing also affects nutrient
cycling a) by “tying up” nutrients in this faunalbiomass, and, b) by
accelerating nutrient release into the soil solution.

3. dispersal of microbial propagules: apart from very specific fungal-faunal
relationships (Ingold, 1971) fauna carry (superficially and in their faeces)
the cells and spores of a wide variety of saprophytic microbes into new
substrates. Therefore microbial community structure and hence organic
matter decomposition rates may be substantially affected.

Faunal activities, particularly organic matter comminution, channelling into organic
subtrata and soil, and defaecation can significantly affect the micromorphology of the organic
, layers of soils and, in some cases, the upper mineral horizon e.g., by reduction of particle size of
j organic matter (with consequent effects on pore volume), by channel formation, and by the

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movement of organic matter into the mineral horizons.

REFERENCES

Ingold, C. T. 1971. Fungal spores, their liberation and dispersal. Clarendon Press, Oxford. 302

pp.

Persson, T., E. Booth, M. Clarholm, H. Lundkvist, B. E. Soderstrom, and B. Sohlenius. 1980.
Trophic structure, biomass dynamics and carbon metabolism of soil organisms in a Scots
Pine forest. In: Persson, T. (Editor). Structure and Function of Northern Coniferous Forests
- An Ecosystem Study. Ecological Bulletins (Stockholm) 32: 419-459.

Reichle, D. E. 1977. The role of soil invertebrates in nutrient cycling. In: Soil Organisms as
Components of Ecosystems. Ecological Bulletin (Stockholm) 25: 145-156.

Satchell, J. E. 1971. Feasibility study of an energy budget for Meathop Wood, pp. 619-630. In:
Duvigneaud, P. (Editor). Productivity of Forest Ecosystems. UNESCO, Paris.

Visser, S. 1985. The role of the soil invertebrates in determining the composition of soil
microbial communities, pp. 297-317. In: Fitter, A.H., D. Atkinson, D. J. Read and M. B.
Usher (Editors). Ecological Interactions in Soil: Plants, Microbes and Animals. B. E. S.
Special Publication No. 4.

EARTHWORMS IN SOIL FORMATION, STRUCTURE AND FERTILITY

Clive A. Edwards

Chairman, Department of Entomology
The Ohio State University

Columbus, Ohio 43210 Quaestiones Entomologicae

U.S.A. 21:517-522 1985

ABSTRACT

The importance of earthworm activity has been recognized since ancient times and their
role in pedogenesis and soil fertility has been studied since the late 1800's. Earthworms
contribute to soil structure and formation through burrowing, comminution of organic matter
and by formation of aggregates. Earthworm guts are important sites of microbial action
whereby nutrients are made available to plants. Earthworm burrows are conspicuous aspects
of soil structure and contribute to soil aeration and drainage. Earthworm populations can be
extremely dense in soils with abundant organic matter, although species diversity of
earthworm assemblages is relatively low. Empirical data suggest that introduction of
earthworms can improve impoverished soils. However, important information about
taxonomy, distribution and biology of North American species is lacking. Few ecological
studies have examined relationships between earthworms and other organisms in the soil.

RESUME

L’ importance de I’activite des vers de terre est connue depuis les temps anciens, et leur role dans la pedogenese et la
formation des sols est le sujet d’etudes depuis la fin du XIXieme siecle. Les vers de terre contribuent h la structure et d la
formation des sols en fouissant, en pulverisant la matiere organique et en formant des agregats. Le tube digestif des vers
de terre est le site important d’une action microbienne par laquelle les nutriments sont liberes des tissus vegetaux. Les
galeries de vers forment un aspect frappant de la structure du sol et contribuent h son aeration et h son drainage. Les
populations de vers peuvent etre extremement denses dans les sols riches en matieres organiques, mais la diversite des
especes en est relativement faible. Des donnees empiriques suggerent que I’introduction de vers de terre peut ameliorer les
sols appauvris. On manque cependant d’informations essentielles sur la taxonomie, la repartition geographique et
I’histoire naturelle des especes de vers nord-americaines. Peu d’etudes ecologiques ont examinees les rapports entre les
vers de terre et les autres organismes du sol.

INTRODUCTION

The great importance of the soil biota in soil pedogenesis and in the maintenance of
structure and fertility is not always fully appreciated by soil scientists. Earthworms are
probably one of the most important components of the soil biota in terms of soil formation.
Although they are not numerically dominant, their large size makes them one of the major
contributors to animal biomass, and their activities are such that they are extremely important
in maintaining soil fertility in a variety of ways.

Aristotle was the first to draw attention to their role in turning over the soil and he aptly
called them “the intestines of the earth”. However, it was not until the late 1800’s that Charles
Darwin, in his definitive work, “The Formation of Vegetable Mould Through the Action of
Worms”, defined the extreme importance of earthworms in breakdown of dead plant and
animal matter that reaches soil and in the continued maintenance of soil structure, aeration

518

Edwards

drainage and fertility. His views were supported and expanded subsequently by other
contemporary scientists such as Muller (1878) and Urquhart (1887) and many others.

Earthworms belong to the Order Oligochaeta which contains about 3,000 species, although
considerable numbers of these are aquatic in habit, and there is considerable controversy on
their systematics. They are found in most parts of the world, except those with extreme
climates, such as deserts and areas under constant snow and ice. Some species of earthworms,
particularly those belonging to the Lumbricidae, are extremely widely distributed (‘peregrine’)
and often, these species when introduced to new areas, become dominant over the endemic
species; this situation probably applies to parts of the northern United States and Canada,
particularly those close to major waterways. However, the endemic earthworm fauna of North
America has not been well studied. Endemic species include those in the Acanthodrilidae with
its most abundant genus Diplocardia , members of the Sparganophilidae and species in the
Megascolecidae of which the most common genus is Pheretima.

EARTHWORMS AND SOIL

Soil Formation

Earthworms are extremely important in soil formation, principally through their activities in
consuming organic matter, fragmenting it and mixing it intimately with mineral particles to
form aggregates. During their feeding, earthworms greatly promote microbial activity which in
turn also accelerates the breakdown of organic matter. Different species of worms do not all
affect soil formation in the same way. Some species consume mainly inorganic fractions of soil,
whereas others feed almost exclusively on decaying organic matter. They can deposit their feces
as casts either on the soil surface or leave them in their burrows, depending on the species
concerned, but all species contribute in different degrees to the comminution and mixing of the
organic and inorganic components of soil, and decrease the size of not only organic but also
mineral particles (Joshi and Kelkan, 1952; Shrikhande and Pathak, 1951). During passage
through the earthworm gut, the different kinds of particles become mixed intimately and form
aggregates, which improve both the drainage and moisture-loading capacity of the soil. These
aggregates are usually very stable and improve many of the desirable characteristics of soils.
There have been various suggestions as to the possible ways in which earthworms form
aggregates, such as by production of gums (Swaby, 1950), or calcium humate (Meyer, 1943),
by plant residues (Ponomareva, 1953) or by means of polysaccharide molecules (Parle, 1963).
Various authors have estimated that up to 50% of the aggregates in the surface layers of soil are
formed by earthworms (Kubiena, 1953).

Earthworms move large amounts of soil from the deeper strata up to the surface. The
amounts moved in this way range from 2 - 250 tons per ha per annum, equivalent to bringing a
layer of soil between 1 mm and 5 cm thick to the surface every year, creating a stone-free layer
on the soil surface. Earthworms also affect soil structure in other ways. Some species make
‘permanent’ burrows, whereas others move randomly through soil leaving cracks and crevices of
different sizes. Both sorts of burrows are important in maintaining both soil aeration and
drainage. Moreover, earthworm burrows are usually lined with a protein-based mucus, which
helps to stabilize these cavities, and many of the species with permanent burrows cast their
feces aroung the lining of their burrows, the cast material usually containing more plant
nutrients in a readily available form than the surrounding soil. There is good evidence that
earthworm activity increases the porosity and air to soil volume (Wollny, 1890; Hopp, 1974;
Edwards and Lofty, 1977). Burrows are also important in improving soil drainage , particularly

Earthworms in soil formation, structure and fertility

519

since those of some species, such as Lumbricus terrestris L. penetrate deep into soil (Edwards
and Lofty, 1978, 1982) and can even pass through layers of clay. The burrows and pores also
increase the infiltration rate greatly (Slater and Hopp, 1947; Teotia et al., 1950; Carter et al .,
1982), and there are numerous reports of water penetrating the surface soil between two and
ten times faster when earthworms were present than when they were not (Stockdill, 1966;
Wilkinson, 1975; Tisdall, 1978). This effect on infiltration can be of two kinds; firstly, the
presence of large surface-opening holes which are not usually taken into account by soil
scientists when conventional models of infiltration are developed (Edwards et al ., 1979), and
secondly, the crevices also created by earthworms, but which are much smaller, not only
increase infiltration but also aid in water retention.

Finally, earthworm activity makes a significant contribution to soil aeration (Stockli, 1928;
Kretzschmar, 1978), by creating channels, particularly in heavy soils which allow air to
penetrate into the deeper layers of soil, minimizing the incidence of anaerobic layers.

Organic matter breakdown and incorporation into soil

Although all species of earthworms contribute to the breakdown of plant-derived organic
matter, they differ in the ways in which they breakdown organic matter. Their activities can be
of three kinds, each associated with a different group of species. Some species are limited
mainly to the plant litter layer on the soil surface, decaying organic matter or wood, and seldom
penetrate soil more than superficially. The main role of these species seems to be comminution
of the organic matter into fine particles which facilitates microbial activity. Other species live
just below the soil surface most of the year, except when very cold or very dry, do not have
permanent burrows and ingest both organic matter and inorganic materials. These species
produce organically enriched soil materials in the form of casts, which they deposit either
randomly in the surface layers of soil or as distinct casts on the soil surface. Finally, there are
the truly soil-inhabiting species which have permanent burrows that penetrate deep into the
soil. These species feed primarily on organic matter but also ingest considerable quantities of
inorganic materials and mix these thoroughly through the soil profile. These latter species are
of primary importance in pedogenesis. All species depend on consuming organic matter in some
form and play an important role in the final stage of organic matter decomposition, which is
humification into complex amorphous colloids containing phenolic meterials, probably by
promoting microbial activity.

There is little doubt that in many habitats, earthworms are the key organisms in the
breakdown of plant organic matter. Populations of earthworms usually expand in relation to the
availability of organic matter, and in many temperate and even tropical forests, it seems that
earthworms have the capacity to consume the total annual litter fall. Such a total turnover has
I been calculated for an English mixed woodland (Satchell, 1967), an English apple orchard
j (Raw, 1962), a tropical forest in Nigeria (Madge, 1965), an oak forest in Japan (Sugi and

I Tanaka, 1978) and it seems likely that similar calculations would be valid for other sites
(Edwards and Lofty, 1977).

|| During feeding by earthworms, the carbon:nitrogen ratio in the organic matter falls
t progressively and, moreover, the nitrogen is converted into the ammonium or nitrate form. At
j the same time the other nutrients, P and K, are converted into a form available to plants. Soils
| that have poor populations of earthworms often develop a mor structure with a mat of
, undecomposed organic matter at the soil surface (Kubiena, 1953); this can also occur in
l| grassland and is common on poor upland grasslands in temperate countries and in New

||@waes/. Ent., 1985, 21 (4)

520

Edwards

Zealand in areas where earthworms have not yet been introduced (Stockdill, 1966).

ECOLOGICAL ASPECTS OF EARTHWORMS
Abundance and Diversity

Populations of earthworms vary greatly both in terms of numbers or biomass and diversity.
Populations range from only a few per square meter to more than 1,000 per square meter.
Numbers depend on a wide range of factors, including soil type, pH, moisture-loading capacity
of the soil, rainfall and ambient temperatures, but, most importantly, to the availability of
organic matter. Populations in cultivated land seldom exceed 100 per square meter, or 400 per
square meter in grassland, the larger populations usually being found in woodlands where the
availability of organic matter is seldom limiting, and occasionally numbers as high as 2,000 per
square meter have been recorded, although few earthworms occur in the more acid soils under
coniferous forests. Usually, the largest populations are of lumbricid earthworms which seem to
be able to survive adverse conditions much better than species belonging to the other families.

The diversity of species of earthworms varies greatly and there tend to be species
associations in different soil types and habitats. The associations of species of lumbricids in
temperate countries tend to be less diverse than those from other families in warmer latitudes.
However, even in the most complex system, the diversity of species does not seem to be very
great, rarely exceeds 10 and commonly, there are only 3-5 species. There is some evidence that
species that fill the same eological niche do not normally occur in the same degree of abundance
at a particular site (Edwards and Lofty, 1982).

Needs for earthworm research

In view of the great importance of earthworms in soil formation and maintenance of soil
fertility, although the numbers of publications on earthworm biology and ecology is increasing
rapidly, there still seems an urgent need for greatly expanded research, particularly on some
aspects of earthworm activity.

We still have inadequate knowledge of the basic biology and ecology of even the more
common species of lumbricids. Very few studies have addressed the problems of the detailed
interrelationships between earthworms, micro-organisms and decaying organic matter and its
incorporation into soil. There is good empirical evidence that introduction of earthworms
together with organic matter, into impoverished soil with addition of organic matter and
adjustment of pH, can increse their fertility greatly, but we have little knowledge of the
mechanism of such increases or even the best ways of introducing earthworms.

Most important is the world-wide lack of knowledge of the distribution of earthworms and
populations of the different species. Until we know more of the fundamental biology and
ecology and the activities of the many different species and their role in maintaining soil
structure and fertility, it is impossible to assess their potential role in soil improvement. These
problems are particularly acute in North America where earthworm specialists are rare and
research extremely sparse.

REFERENCES

Carter, A., J. Heinonen and J. deVries. 1982. Earthworms and water movement. Pedobiologia
23, 395-397.

Darwin, C.R. 1881. “The formation of Vegetable Mould through the Action of Worms, with

Earthworms in soil formation, structure and fertility

521

Observations on Their Habits”. Murray, London. 326 pp.

Edwards, C.A. and J.R. Lofty. 1977. “Biology of Earthworms”. 2nd Edition, 333 pp.

Edwards, C.A. and J.R. Lofty. 1978. The influence of arthropods and earthworms upon root
growth of direct drilled cereals. Journal of Applied Ecology 15, 789-795.

Edwards, C.A. and J.R. Lofty. 1982. The effect of direct drilling and minimal cultivation on
earthworm populations. Journal of Applied Ecology, 19, 723-734.

Edwards, W.M., R.R. Van der Ploeg and W. Ehlers. 1979. A numerical study of noncapillary
sized pores upon infiltration. Journal of the Soil Science Society of America 43, 851-856.
Hopp, H. 1974. What every gardener should know about earthworms. Garden Way Publishing
Company, Charlotte VT, U.S.A. 39 pp.

Joshi, N.V. and B.V. Kelkar. 1952. The role of earthworms in soil fertility. Indian Journal of
Agricultural Science 22, 189-196.

Kretzschmar, A. 1978. Quantification ecologique des gaeeries de lombriciens. Techniques et
premieres estimations. Pedobiologia 18, 31-38.

Kubiena, W.L. 1953. “The Soils of Europe”. Murray, London. 317 pp.

Madge, D.S. 1965. Leaf fall and disappearance in a tropical forest. Pedobiologia 5, 273-288.
Meyer, L. 1943. Experimenteller Beitrage zu makrobiologischen Wirkungen auf Humus and

IBoden bildung. Archives Pflanzenernahrung Dungung Bodenkunde 29, 1 19-140.

Muller, P.E. 1878. Studier over Skovjord I. Om Bogemuld od Bogemor paa Sand og Ler.
Tidsskrift Skogbruk 3, 1-124.

Parle, J.N. 1963. A microbiological study of earthworm casts. Journal of General
Microbiology 31: 1-13.

Ponomareva, S.I. 1953. The influence of the ctivity of earthworms on the creation of a stable
structure in a sod-podzolised soil. Trudy Pochvenie Institut Dokuehaeve 41: 304-318.

Raw, F. 1962. Studies of earthworm populations in orchards. I. Leaf burial in apple orchards.
Annals of Applied Biology 50, 389-404.

Satchell, J.E. 1967. pp. 259-322. In: Burgess, A. and F. Raw (Editors). “Soil biology”.
Academic Press, London.

Scrickhande, J.E. and A.N. Pathak. 1951. A comparative study of the physico-chemical
characters of the castings of different insects. Indian Journal of Agricultural Science 21,
401-407.

Slater, C.S. and H. Hopp. 1947. Relation of fall protection to earthworm populations and soil
physical conditions. Proceedings of the Soil Science Society of America 12, 508-51 1.
Stockdill, S.M.J. 1966. The effect of earthworms on pastures. Proceedings of the New Zealand
Ecological Society 13, 68-75.

Stockli, A. 1928. Studien uber den Einfluss der Regenwurmer auf die Beschaffenheit des
Bodens. Landwirtschaft Jahrbuch Schweiz. 42, 1 .

Sugi, Y. and M. Tanaka. 1978. Number and biomass of earthworm populations, pp. 171-178.
In: Kira, T., Y. Ono and T. Hosokawa (Editors). “Biological production in a warm
temperature evergreen oak forest of Japan”. J.I.B.P. Synthesis 18 University of Tokyo Press.
Swaby, R.J. 1950. The influence of earthworms on soil aggregation. Journal of Soil Science l,
195-197.

Teotia, S.P., F.L. Duley and T.M. McCalla. 1950. Effect of stubble mulching on number and
activity of earthworms. Nebraska Agricultural Experiment Station Bulletin 165, 20.

Tisdall, J.M. 1978. Ecology of earthworms in irrigated orchards, pp. 297-303. In: Emerson,
W.W., R.D. Bond and A.R. Dexter (Editors). “Modification of Soil Structure”. Wiley,

Quaest. Ent., 1985,21 (4)

522

Edwards

Chichester.

Urquhart, A.T. 1887. On the work of earthworms in New Zealand. Transactions of the New
Zealand Institute 19, 119-123.

Wollny, E. 1890. Untersuchungen iiber die Beeinflussung der Fruchtbarkeit der Ackerkrume
durch die Tatigdeit der Regenwiirmer. Forschungen Gebeit Agrik Physik Bodenkunde 13,
381-395.

Wilkinson, G.E. 1975. Effect of grass fallow rotations on the infiltration of water into a
savanna zone soil of Northern Nigeria. Tropical Agriculture ( Trinidad ) 52, 97-103.

ASPECTS OF THE BIOLOGY AND SYSTEMATICS OF SOIL ARACHNIDS,
PARTICULARLY SAPROPHAGOUS AND MYCOPHAGOUS MITES

Roy A. Norton

Department of Environmental and Forest Biology

State University of New York

College of Environmental Science and Forestry

Syracuse, New York 13210

U.S.A.

Quaestiones Entomologicae
21:523-541 1985

ABSTRACT

Among members of the class Arachnida, the saprophagous and mycophagous mites are the
most diverse and abundant in soil systems, and have the greatest effects on soil structure and
fertility. In most soils, the oribatid mites are the only arachnids which directly affect soil
structure; they ingest particulate food and produce discrete fecal pellets, the possible
functions of which are discussed. Current research is finding an active gut microflora in both
saprophagous and mycophagous oribatid mites. For those species in which the diet varies
seasonally, it is suggested that the gut microflora also changes, reflecting the new substrates.
Particulate-mycophagy is a very old feeding habit, and was probably the ancestral one of the
earliest known [ Devonian ) oribatid mites. Saprophagy may have originally developed as a
mechanism for obtaining non-surface microbial tissue or exometabolites from decaying higher
plant materials. The use of the plant structural parts as food may have been made possible by
changes in gut microflora. The mite suborder Prostigmata also contains important soil
mycophages, but these feed intracellularly and contribute little to soil structure. They are
more “ r-selected ” than oribatid mites and may show numerical responses to temporary
increases in food supply.

The descriptive taxonomy and systematics of North American saprophagous and
mycophagous soil mites are in a poor state. Currently no “ user-friendly ” monographs are
available for any major group, and higher classifications based on the methodology of
phylogenetic systematics are only beginning to be proposed.

RESUME

Parmi les membres de la classe des Arachnides qui vivent dans les sols, les miles saprophages el mycophages sonl les
plus divers el les plus abondants, et sonl ceux dont les effels sur la structure el la fertility des sols sont les plus marques.

1 Dans la plupart des sols, les mites oribatides sont les seuls arachnides qui affectent directement la structure du sol; elles
| ing'erent des aliments particulaires et excrttent des boulettes fecales. dont les fonctions possibles sont examinees. Les
I recherches actuelles rev&lent une f lore active du tube digestif chez les mites oribatides autant saprophages que
! mycophages. Chez les esp'eces dont la didte varie saisonnikrement, il est possible que la microflore du tube digestif change
\ aussi, refletant ainsi les divers substrats. La mycophagie particulaire est une trks vieille habitude alimentaire qui
| remonte probablement aux plus anciennes mites oribatides connues (du Devonien). La saprophagie pourrait d I'origine
I etre apparue comme un mecanisme per melt ant d’obtenir des couches sous-jacentes de tissus microbiens ou des
1 exometabolites provenant de plantes superieures en decomposition L' utilisation de parties structurales des plant es
1 comme nourriture a pu etre rendue possible par des changements dans la microflore du tube digestif. Les mites du
. sous-ordre Prostigmata incluent aussi d'importants mycophages du sol. mais ceux-ci se nourrissent intra-cellulairement
et ne contribuent it peu pris pas d la structure des sols. Elles sont davantage soumises d la r-silection que les mites

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oribatides et peuvent voir leur nombre augmenter h la suite d’un accroissement temporaire de la quantite de nourriture
disponible.

La taxonomie descriptive et la systematique des mites nord-americaines saprophages et mycophages des sol est dans
un etat lamentable. Presentement il n’existe aucune monographic d’utilisation facile pour aucun des groupes majeurs. et
des classifications superieures basees sur les methodes de la systematique phylogenetique commencent a peine a faire
leur apparition.

INTRODUCTION

The subject of biology and systematics of the Class Arachnida is large even if one’s attention
is restricted to soil dwellers. Representatives of this class usually dominate the arthropod fauna
of soils; numbers in the hundreds of thousands per surface m2 are common (Petersen, 1981a).
The soil arachnid fauna can be conceptualized and categorized from many viewpoints, but the
one used here is that of function, particularly predation, saprophagy and mycophagy.

Predation is the most ancient and general feeding strategy of arachnids and predaceous
species abound in soil and litter. Various groups of cursorial spiders constitute a major fraction
of the larger (length > 1 cm) predaceous soil arthropods; pseudoscorpions and harvestmen,
along with other spider groups, are intermediate (1 mm - 1 cm) in size. By far the greatest
diversity of small arthropod predators in most soil systems is contributed by representatives of
two suborders of mites, the Mesostigmata (temperate region soil dwellers are mostly predators)
and the Prostigmata (in part). The latter group also includes taxa which are external parasites
of other arthropods. Little will be said here regarding these predators and parasites, since their
influence on physical and chemical attributes of soil is at best indirect, through their regulatory
interactions with other soil animals. This is not meant to infer that they are unimportant to the
proper functioning of soil systems; we simply have little empirical information. Progress has
been made in understanding the role of spider predation in agroecosystems (Riechert, 1984),
where they are generalist background regulators of insect populations, but their role in
soil/litter systems is virtually untested, except for the work of Clarke and Grant (1968) and
Kajak and Jakubczyk (1977), who demonstrated higher densities of centipedes and
saprophages such as collembolans after removal or exclusion of spiders and other large
predators. Even the detailed study by Moulder and Reichle (1972) on the significance of spider
predation in the forest floor fell short due to lack of information on prey dynamics. The role of
predation in regulating soil arthropod and nematode (see Karg, 1983) densities remains, for the
most part, unknown. An exception is the important role of predaceous Tydeidae (Prostigmata)
in regulating bacteriophagous nematode populations in desert soils ( e.g .,, Whitford and Santos,
1980).

It is the arachnid groups generally considered to be saprophagous and mycophagous
(fungivorous) which are of greatest interest from the standpoint of soil structure and fertility,
and these will be stressed in the discussions which follow. Saprophages, those feeding directly
on decomposing leafy or woody vegetation (macrophytophages of Schuster, 1956) have the
greater significance in producing structure in organic horizons, particularly by comminution
activities and production of feces. Among the Arachnida, mites of the suborder Oribatida
(Cryptostigmata) perform this function on a scale which is small in size, but not necessarily in
overall effect. Fungivorous mites abound in soils, and along with Collembola form the dominant
mycophages in most terrestrial ecosystems (Seastedt, 1984). Two general fungal-feeding
strategies are apparent: the fungivorous oribatid mites are particulate feeders, and produce
discrete fecal pellets which contribute to soil structure. The fungivorous prostigmatid
(trombidiform) mites feed intracellularly by means of stylettiform chelicerae and contribute

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525

little to soil fabric.

Due to the scope of the topic, available, this paper cannot be considered a complete review of
current problems in arachnid biology and systematics. Only saprophagous and
microphytophagous (bacterial and fungal feeders) arachnids will be dealt with in any detail. In
particular, the oribatid mites are emphasized, for several reasons: 1) they are usually the
dominant arachnid group in terms of numbers and biomass (Petersen, 1982a, 1982b); 2) they
are apparently the most important group of soil arachnids from the standpoint of direct and
indirect effects on the formation and maintenance of soil strucure; 3) they are the best known
biologically, even if “best” is not very good; and 4) they are the soil animals with which I am
most familiar. Little will be said of the mite suborder Astigmata; their occurrence in soils is
sporadic and their overall importance is likely to slight (Luxton, 1981a; see O’Connor, 1982
and Woodring, 1963). Even with these restrictions my comments will be selective. From the
standpoint of biology, I will deal particularly with certain aspects of nutrition, followed by brief
remarks on population dynamics, and then review how the feeding activity of these animals
affects soil structure and fertility. Lastly, I will offer some comments on systematics of soil
arachnids, again with emphasis on non-predators.

BIOLOGY

Some Aspects of Oribatid Mite Nutrition

The diet and method of feeding of saprophagous and mycophagous soil mites greatly
influences the effect they have on soil structure and fertility. Such information was rather
scattered and anecdotal before Schuster’s (1956) classic paper, which described feeding habits
of a wide variety of European oribatid mite. Since that time numerous authors have approached
the subject, generally using one or more of three methods: laboratory food preference tests, gut
I content analysis of field-collected specimens, or the study of gut enzyme complements [see
studies and reviews by Wallwork (1967), Lebrun (1971), Luxton (1972), Pande and Berthet
j (1973), Behan and Hill (1978), Behan-Pelletier and Hill (1983), and especially Harding and
Stuttard(1974)].

Each of these techniques has associated problems. Laboratory preference tests are limited
by our ability to discern and supply the food items available in nature and to keep other, often
unknown, variables within preference, or at least tolerance, ranges of the mites being tested.
Simple choice experiments rarely fail to demonstrate preferences, yet these preferred foods may
not, in some circumstances, be the ones chosen in nature, even if they are available (Mitchell
and Parkinson, 1976). Also, even heavy feeding on a given food does not mean that
reproductive or developmental success is necessarily possible (Luxton, 1972; Saichuae et al.,
1972; Mitchell and Parkinson, 1976; Stefaniak and Senniczak, 1981).

Gut content analyses have the disadvantage that less easily digestible materials will be
overestimated in dietary quantification (Behan-Pelletier and Hill, 1983). In some situations,
i readily digestible materials may be supplying most of an animal’s requirements, yet rarely be
seen in the gut by conventional methods. For example, it is unknown to what extent oribatid
mites are predatory or necrophagous. Muraoka and Ishibashi (1976) and Rockett (1980) have
! demonstrated active feeding on living and dead nematodes by brachypyline oribatid mites and
| the importance of this in nature needs investigation. Stefaniak and Seniczak (1981) found that
cannibalism could occur in oribatid mites in association with poor nutrition, and
Behan-Pelletier and Hill (1983) found unusually high quantities of small animal remains in
I guts of acid-bog dwelling oribatid mites. They also suggested that the presence of such dietary

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components might be related to the overall poor nutrient content of other available foods. The
role of bacteria or other Monera, scraped from soil substrates, as food for oribatid mites is
virtually unknown. Luxton (1972) demonstrated attractiveness of bacteria to some species, and
Behan-Pelletier and Hill (1983) found high percentages of Monera in some oribatid guts, but
whether they are truly food sources, or function as symbiotic sources of enzymes necessary to
complement the mites’ endogenous enzyme system, or both, remains to be elucidated (see
below).

Studies of gut enzymes (Luxton, 1972; Zinkler, 1971, 1972) have been enlightening, but
they are typically done on whole-body homogenates and give no information on the origin of
any given enzyme. Luxton (1972, 1979) suggested that enzyme complements of oribatid mites
correspond to the exhibited feeding habits. Species which are principally saprophagous (in
particular the xylophagous, family Phthiracaridae) possess enzymes capable of hydrolyzing
structural carbohydrates of higher plants ( e.g ., cellulase, xylanase, pectinase), whereas those
ingesting primarily fungal tissue can hydrolyze the fungal storage sugar trehalose and perhaps
chitin, which is found in hyphal walls. Only panphytophagous (unspecialized) species possess
both enzyme systems. Two other authors (Zinkler, 1972; Dinsdale, 1974), however, failed to
find a cellulase in phthiracarid mites. There is mounting evidence that many carbohydase
enzymes, including cellulase and chitinase, are produced by a diverse and very active microflora
(Seniczak and Stefaniak, 1978; Stefaniak and Seniczak, 1976, 1981). These studies indicate
that the composition of the gut microflora in both panphytophages and mycophages (and the
enzymes produced) depends on the food ingested, and is not identical to the microflora of the
surrounding environment. Their floral lists indicate that a subtraction process occurs. Those
microorganisms capable of continued (or enhanced?) enzymatic production in the gut may be
those which are resistant to digestion. Dinsdale (1974) has demonstrated that in Phthiracarus
sp. (which had few demonstrated gut microflora) enzymes acting on the glucosidic bonds of
simple storage carbohydrates were closely associated with the gut mucosa, and protein
digestion appeared to be intracellular, with evidence of pinocytosis. One can speculate that
these enzymes are endogenous, and that all enzymes acting on structural polysaccharides which
are found in oribatid mite guts are of microbial origin, as they apparently are in all other
arthropods which possess them.

It has been known for many years that saprophagous oribatid mites will only eat material if
it has been previously attacked by microorganisms (see Harding and Stuttard, 1974), but the
earlier consensus view was that the microbial tissue itself, or exometabolites from its
decomposition activities, provided the only energy source. The finding of active gut microflora
in saprophagous groups is not surprising, but it is somewhat surprising that fungivorous mites
may have a strong dependence on gut microflora.

The implications and questions raised by Stefaniak and Seniczak’s work are important ones
and relate to the ability of soil mites to adjust to forced or opportunistic changes in diet. There
is growing evidence (e.g., Anderson, 1975; Swift, Heal and Anderson, 1979) that in some soils
many or most oribatid mites do not adhere strictly to saprophagy or mycophagy throughout the
year. Feeding is catholic, and gut contents of a given species can vary with site and season. At
times even those species normally associated with strict xylophagy (Phthiracaridae) can be
fungivorous, and those with chelicerae seemingly specialized for mycophagy, such as Eupelops,
can be saprophagous (Wallwork, 1967; Anderson, 1975; Behan-Pelletier and Hill, 1983).
Anderson (1975), working in a British deciduous forest, considered mycophagy to be the
dominant feeding activity by oribatid mites soon after leaf fall in autumn. Within a rapidly

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degraded litter layer, fungi were less easily available during the following season, and feeding
activity changed to mixed mycophagy/saprophagy and then to saprophagy prior to the
following leaf-fall. Such patterns were apparent even within populations of a given species.

A seemingly conflicting pattern was found, however, in a Canadian aspen forest soil
(Mitchell and Parkinson, 1976) where the litter layer was more stable. The dominant taxa were
primarily fungivorous and there were one or two seasonal peaks in feeding rate related to
leaf-fail. Perhaps most interesting was the fact that overall feeding rate was related to general
microbial activity, and opportunistic switiching to saprophagy apparently did not occur at this
site. Nor did Behan-Pelletier and Hill (1983) find seasonal patterns in diet composition during
a six-month study of feeding by oribatid mites in an Irish acid peat bog, other than
opportunistic use of pollen. For the most part, species were panphytophagous. It seems that the
existence of seasonal patterns in oribatid mite feeding depends on site characteristics, especially
the rate of early decomposition. Where major diet changes do occur, as in Anderson’s (1975)
site, one can speculate that they are made possible by passive access to a rich variety of
symbiotic microflora ingested with the food. Different bacterial floras (and their respective
enzyme complements) may preferentially develop in the gut during the part of the season when
the appropriate food enters the diet. Stefaniak and Seniczak (1976, 1981) have demonstrated
such a relationship between food type and the composition of the gut microflora under both
saprophagous and mycophagous feeding regimes. Perhaps in less active sites, such as that
studied by Mitchell, changes in available microflora are insufficient to necessitate a switch, or
perhaps climatic or other factors intervene when fungal availability is low, resulting in lower
ingestion rates rather than dietary changes. The ability of any oribatid mites to change diet has
been suggested to be adaptive from the standpoint of increasing survival probability in variable
or different environments (Wallwork, 1958; Luxton, 1972), but it is interesting that the two
most widely distributed (both geographically and ecologically) and successful species known,
Tectocepheus velatus (Michael) and Oppiella nova (Oudemans), are apparently strictly
mycophagous.

Saprophagy and Mycophagy in Other Soil Mite Groups

Compared to oribatid mites, little is known of the feeding biology of other mycophagous or
saprophagous mite taxa. Members of the Uropodina (belonging to the predominantly
predaceous suborder Mesostigmata) exhibit these feeding habits, (Krantz, 1978), but they do
not constitute an important fraction of the soil fauna in most areas of North America; in the
tropics they are much more abundant, often outnumbering oribatid mites. Like their
predaceous relatives, they are predominantly liquid-feeders (Karg, 1963; Ahtias-Binche, 1977,
1981) so their influence on soil structure is probably minimal.

A different type of mycophagy is exhibited by some members of the suborder Prostigmata.
The apparent majority of fungal-feeding soil Prostigmata to the families Eupodidae, Tydeidae,
Tarsonemidae, Scutacaridae, and Phgmephoridae (Evans et al ., 1961; Karg, 1963; Krantz and
| Lindquist, 1979; Kethley, in press). Although present in soils of most ecosystems, these mites
; are especially abundant and diverse in herbaceous systems (where oribatid mites are usually
I not dominant) such as prairies, oil-fields and arctic sites (Petersen, 1982a) and also in desert
soils (Santos et al ., 1978). In a study of an oil-field soil in Ohio (D. Dindal and R. Norton,

' unpublished) representatives of more than 100 species of Prostigmata were collected over
several years, most belonging to the aforementioned families. Although there is little supporting
Information, their intracellular style of feeding would seem to preclude any direct impact on

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soil structure, in contrast to the comminution and fecal production exhibited by
particulate-feeding oribatid mites. They are not, therefore, strictly ecological equivalents,
despite a common general food source. As with oribatid mites, mycophagous Prostigmata may
demonstrate feeding preferences in laboratory tests ( e.g ., Kosir, 1975). As might be expected,
no active gut microflora has been reported from these mites. Fungivory is also known in another
group of mites commonly associated with the Prostigmata, a group often referred to as the
“Endeostigmata” or Pachygnathoidea. At least some members of this group are considered
early-derivatives of the lineage which gave rise to oribatid mites (e.g., O’Connor, 1984). Thus,
it is not surprising to find particulate-mycophagy in some of these taxa, such as Terpnacarus
(Theron, 1979) and Grandjeanicus (personal observation). This brings up the question of the
evolutionary development of the various feeding strategies discussed to this point.

Comments on the Evolution of Mycophagy and Saprophagy in Acariform Mites

Intimate asssociations of arachnids and soils are as old as soils themselves (Kevan et al.,
1975). When terrestrial vegetation first began to flourish in Devonian times soil mites were
already present, and in forms not very different from some which exist today. It is fortunate
that some representatives of these ancient lineages have survived, because they give insight into
not only the systematic relationships of early soil mites, but also their possible feeding habits.
The earliest of the mite fossils, Protacarus crani from the Devonian Old Red Sandstone
formation of Scotland (Hirst, 1923), is very similar to extant members of the “Endeostigmata”
(Krantz, 1978), a loosely defined group which is currently thought by some (including myself)
to have given rise independently to both the Prostigmata and the Oribatida-Astigmata lineages.
Some extant members of the group are particulate-mycophages, as noted above. These have
modified mouthparts with specialized setae (rutella) used in conjunction with the chelicerae to
shear off particles as food is pulled toward the mouth (Grandjean, 1957; Theron, 1979), and
appear to be part of the lineage which includes oribatid mites (also with rutella). Other
“Endeostigmata,” lack rutella (a few of these possibly secondarily so) and feed on spores or
pierce roots to obtain nourishment (Theron, 1979). Although the mouthparts of Protacarus are
not well described, they appear to be rather stylettiform and if so, are consistent with this
feeding type. The earliest known fossil oribatid mite, also of Devonian age, is apparently a
member of the extant family Ctenacaridae (Shear et al ., 1984) which are
particulate-mycophages (Grandjean, 1954; personal observation). In none of these early
derivative groups (fossil or extant) is there any evidence of saprophagy, which makes Krantz
and Lindquist’s (1979) suggestion, that mycophagy in oribatid mites evolved from saprophagy,
difficult to accept. It is far more likely that sarcoptiform mites (those with a cutting rutellum)
were ancestrally mycophagous, and fed on the rich terrestrial microflora which probably
existed in the primordial organic soils developing concurrently with the growth and
decomposition of the earliest vascular plants.

Saprophagy was apparently derived within the oribatid mites, seemingly associated with
even stronger, more robust development of the rutellum and sclerotization in general. It is
reasonable to speculate that the appearance of saprophagy gave previously mycophagous mites
a mechanism for ingesting non-surface microbial tissue (or easily utilized exometabolites). The
use of the higher plant structural material itself, by means of a symbiotic gut microflora
derived from environmental sources, may have evolved later and perhaps necessitated changes
in gut chemistry to allow or promote the growth of gut bacteria and actinomycetes.

Soil Arachnids

529

In any event, it is clear that mycophagous and saprophagous soil arachnids were not added
to soils as an evolutionary “after thought”, except perhaps some of the
intracellular-mycophagous Prostigmata (see Krantz and Lindquist, 1979). Rather, the soil
system as we know it today, with its complex patterns of energy flow and nutrient cycling, is a
result of a coevolution between mites and other soil animals (the ecologically similar
Collembola are equally as old), the microflora, and the developing terrestrial vegetation.

A Brief Overview of Development and Population Dynamics

Knowledge of developmental biology and population dynamics of saprophagous and
mycophagous mites is essential for determining the magnitude of their relationship to soil
structure and fertility. Here again, we know much more about oribatid mites than the
mycophagous taxa of Prostigmata. Early estimates of developmental times for oribatids are
mostly useless, since they were done with laboratory cultures at high, constant temperatures
(20-30° C) and with constantly high humidity and food supply (see Lebrun, 1971; Luxton,
1981b for reviews). Consequently, grossly underestimated development and generation times
were the rule. Multivoltine life histories were commonly suggested for temperate-zone oribatid
mites despite the fact that annual mean temperatures in in the soil may be only half those of the
laboratory. The complexity and variation of natural abiotic and biotic factors make simple
extrapolations impossible, in light of our knowledge of such factors as high development Qi0
values (Lebrun and Ruymbeke, 1971), variable temperatures (Lebrun, 1977), and food quality
(Saichuae et al., 1972; Mitchell and Parkinson, 1976; Young and Block, 1980; Stefaniak and
Seniczak, 1981) upon developmental time, survival rates, and metabolic rates of these mites.

Improved estimates have resulted from following population age-structure in the field over
time (e.g., Mitchell, 1977; Thomas, 1979; Luxton, 1981b, 1981c). There are difficulties with
this method, however, when oviposition is not temporally circumscribed. Also, some workers
have equated developmental time (egg-adult) with the more ecologically significant generation
time (adult-adult). For example, Weigmann’s (1979) estimate of a one-year generation time in
Platynothrus poltifer (Koch) is possible only if eggs are laid immediately after the adult instar
is reached. As Harding (1973) has shown, however, a long preoviposition period is typical for
this species and what Weigmann probably observed was two principal cohorts in a population
with a one year development time, but a nearly two year generation time.

The recognition of preoviposition periods and the possible presence of cohort structure
suggests a need for reexamination of earlier age-structure data. It is likely that natural
developmental times of oribatid mites in temperate regions (where they are generally most
abundant) take a year or more, and two-year generation times are probably not uncommon.
Longevity is probably relatively high in natural conditions, with iteroparity common (Mitchell,
1977; Luxton, 1981b); this may keep cohort recognition from being absolute.

Mortality factors in oribatid mite populations are poorly known. Lebrun (1969) and
Mitchell (1977) suggested that mortality is concentrated in the immatures; presumably this is
mostly due to predation on these soft-bodied instars and the rigors of the molting process. But
adults are not immune to predation (Riha, 1951; Norton and MacNamara, 1976) and they
seem to be more vulnerable to internal parasaites (e.g., Purrini, 1983 and included references).
Cold-induced winter mortality may not be important (Mitchell, 1977). Adaptations to survive
subfreezing temperatures have been elucidated (e.g., Somme and Conradi-Larsen, 1977; Block,
1980) and include elevation of cryoprotectants (such as glycerol) in the hemolymph and the
cessation of feeding to avoid the presence of ice nucleating agents in the gut. Feeding activity

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during subfreezing winter temperatures may require the surpassing of a snow-thickness
threshold (Aitchison, 1979).

Oribatid mites can generally be considered K-selected organisms (Mitchell, 1977). Their
high diversity, low fecundity ( e.g Luxton, 1981b), increased variability in instar length as
development procedes (Lebrun, 1971; Luxton, 1981b), and high longevity coupled with
iteroparity, create an overall relatively stable community, especially in forest soils. The higher
oribatid mite densitites often noted during winter (see Harding and Stuttard, 1974) may be
artifacts of the sampling process in two different ways. Persson and Lohm (1977) found that
soil compaction during sampling trapped significant numbers of microarthropods, except when
the soil was frozen. Also, as reviewed by Luxton (1981b), about one-third of an oribatid mite’s
post-hatching development time is spent in pre-ecdysial resting stages and Lebrun (1969) has
noted that this can cause substantial under-estimates of population densities calculated from
desiccating-style extractors. Since molting occurs primarily during warmer months,
under-estimation should especially be a problem in this period. It may be that in temperate
regions the best overall estimates of population density are those obtained in late fall or early
winter.

Information on population dynamics of mycophagous Prostigmata (e.g., Heterostigmata,
Tydeidae, Eupodidae) is much less extensive. Information about structure of age-classes is
almost non-existent, but multiple generations per year seem to be likely (Luxton, 198 Id).
Unlike oribatid mites, they may show numerical responses to temporary increases in food
supply. Very high densities are commonly found in litter-bag studies (Crossley and Hoglund,
1962; personal observations), where the compact, moist substrate is conducive to rapid mycelial
growth. At least some of these taxa (although certainly not the “Endeostigmata”) are r-selected
as is apparently so for some of their phytophagous aerial relatives (Krantz and Lindquist,
1979).

The Role of Saprophagous and Mycophagous Mites

The literature on the role of saprophagous and mycophagous mites and collembolans in the
functioning of soil systems is full of contradictions (Cancela de Fonseca and Poinsot-Balaguer,
1983), especially from the standpoint of whether or not their activities are necessary for
expeditious decomposition of annual organic matter input. What is now clear is that these
animals, even when abundant, use a very small amount of the annual energy input to the
soil-litter system, generally less than 1% (Mitchell, 1979. Thomas, 1979; Luxton 1982a). This
is principally due to their small standing crop biomass combined with a low weight-specific
respiratory metabolism (Mitchell, 1979). Even in the absence of significant direct use of
energy, the modern consensus is that they “earn their keep” indirectly through comminution
and relationships with soil microflora. In reviewing standardized results of exclusion
experiments, Seastedt (1984) calculated an average contribution by microarthropods of 23% to
reduction of standing litter crop. Much research has gone into explaining results from these
“black box” experiments, reviewed most recently by Seastedt (1984).

Saprophagous and mycophagous mites, particularly oribatids, influence soil structure by
comminution of organic inputs, the production of fecal pellets and perhaps the prevention of
fungal matting. Burrowing activity is limited to internal tissues of leaves, petioles, twigs, etc.
and mineral particles are rarely ingested, or at least rarely reported (Harding and Stuttard,
1974). The only clear instance of organic-inorganic soil mixing by a mite seems to be that
reported by Robaux et al. (1977), who, under laboratory conditions, found that Tyrophagous

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531

putrescentiae can create mixed microaggregates in clay soil and increase aeration by the
formation of cavities. Direct vertical and lateral translocations of organic matter are probably
insignificant, since soil mites are rather sedentary on a diurnal basis and generally defecate on
or near their food source. The significance of fecal pellet production by soil mites and
collembolans is usually considered to be the increased surface area (relative to uncomminuted
material), increased water-absorbing qualities, higher nitrogen concentration, higher pH and
their small size, which allows illuviation into lower soil horizons. All these actions purportedly
increase microbial activity, especially that of bacteria, as bacterial populations flourish in the
higher pH regimes of the gut, feces and lower horizons. Since bacterial activity is a surface
phenomenon, constantly requiring fresh surfaces (Luxton, 1 98 1 e) the comminution aspect
seems especially important. When feeding occurs on leaf mesophyll, for example, not only is
new surface exposed, but the food particle itself is subjected to decomposition in the gut and in
the fecal pellet eventually formed. Increases in surface area due to fecal pellet formation are
modest, however. Nef (fide Harding and Stuttard, 1974) found a 10,000-fold increase in
surface area of a conifer needle when comminuted by a phthiracarid mite, but reformation into
pellets reduced this to a 4-fold increase.

Although fecal pellets may decompose readily in certain situations (e.g., Jongerius, 1963),
especially when moved downward in the profile, they are often rather long-lived (Grosbard,
1969; Webb, 1977; Bal, 1982) and accumulate, especially in mor soils with few large
invertebrates to actively mix materials. Webb (1977) has noted that high cohesive forces
between pellet particles, especially very small ones, are apparently the cause of this general
recalcitrance, which is not observed with feces of larger arthropods. The most important
function of small fecal pellets may be that they maintain the highest possible surface area for
decomposition. He suggested there is a theoretical lower limit to size of free particles in the soil,
and if not compacted into pellets the bite-sized particles would form even larger aggregates.
Other work suggests that the increased surface area of mite fecal pellets is ineffective in
increasing decomposition of the contained material. For example, the decaying grasses studied
by Grosbard (1969) showed rapid decomposition after being fed upon by mites, but the fecal
pellets decomposed very slowly. Perhaps mite feeding has an ecological cost associated with it,
especially in the absence of soil mixing, or the slower decomposition of feces may serve an
important regulatory function in some situations. The overall impact of comminution by
saprophagous mites on soil structure and decomposition depends to some extent on the
proportion of annual organic input which they ingest. Recent studies (Mitchell, 1979; Thomas,
1979; Luxton, 1 98 1 e) suggest that the figure may be almost insignificant (less than 2%)
although few soil types have been studied. However, a small amount of feeding may go a long
ways toward opening up new substrates for microbial decomposition. Equating mite ingestion
rates with their contribution to decomposition processes maybe as fallacious as equating the
metabolic contributions of soil animals to their importance in soil systems.

For two decades the concensus has been that the real importance of saprophagous and
mycophagous mites and other microarthropods has been in their interactions with the soil
microflora (the “catalytic” effect of Macfadyen, 1961). Most recently these interactions have
been viewed in relation to nutrient elemment cycling. While microarthropod bodies have been
implicated as potentially important sinks and sources of nutrients (Crossley, 1977; Luxton,
1979; Wallwork, 1983), their low standing crops make the amount insignificant in relation to
quantities immobilized by microorganisms, particularly fungi (Seastedt, 1984). The principal
significance of mycophagous mites seems to lie in their ability to extract limiting nutrients {e.g.,

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nitrogen) from fungal standing crop and, with their death or excrement, make them available
for rapid reuse in further mycelial or bacterial growth, with concomitant organic substrate
decomposition (Whitford and Santos, 1980; Seastedt, 1984). As with comminution, the impact
of soil mites will be some function of the amount of fungal tissue consumed. Estimates are rare,
but consumption may be quite low in relation to fungal standing crop (Mitchell and Parkinson,
1976). The relationship of soil mites and other microarthropods to nutrient dynamics is
complex (Seastedt, 1984) and whether their feeding helps fungal growth, or suppresses it and
shunts decomposition to bacterial pathways (which may be more rapid and complete), depends
on characteristics of the site, the substrate, and the organisms involved.

Considering their high densities, taxonomic diversity, conservative population dynamics, and
a broad mix of specialized and opportunistic feeding habits, oribatid mites seem to be stable
background decomposers, analogous in a way to most predaceous arachnids in that they seem
incapable of rapidly adjusting to changes in resource availability. Crossley (1977) has
contrasted this K-strategy to the r-strategy of collembolans, which may dominate microbial
feeding at times of rapid growth; the r-strategy may also characterize fungivorous Prostigmata.

To better understand these processes we need more detailed information on feeding
specificities of caprophagous and mycophagous mites, not just what will be eaten, but the
physical and chemical cues which provoke feeding (Cancela de Fonseca and Poinsot-Balaguer,
1983). We also need comparative information, from a wide vaiety of habitats, on the portion of
organic matter input comminuted by oribatid mites and similar estimates on consumption of
microbial standing crops by mycophagous mites. Comparative studies of longevity of fecal
pellets and the extent and role of their bacterial enhancement in subsequent decomposition will
also be important.

Some Additional Areas Needing Attention

Many other aspects of the biology and ecology of saprophagous and mycophagous soil mites
are in need of continued study, and these can be used to illustrate or test many current general
hypotheses. Biotic and abiotic determinants of community structure and microdistribution are
known only in very general terms (see Anderson, 1975; Usher et al., 1982; Wallwork, 1983).
The importance of competition in determining coexistence of similar saprophagous and
mycophagous mites ( e.g ., Anderson, 1978) is an especially timely subject. Are the consistent
size differences observed between coexisting pairs or series of congeneric oribatid mites (Walter
and Norton, in press) due to limiting similarities imposed by exploitation competition (if so,
what is the resource?), or are they simply manifestations of reproductive isolation mechanisms?
Much remains to be learned about the distribution and biology of deep-soil mites, especially the
Prostigmata (Kethley, in press). Gerson (1983) has recently suggested that filtrates from
surface organic substrates may be an important resource for such animals.

Inter- and intra-habitat dispersal is another aspect which is virtually unstudied. We have
some knowledge of dispersal in species restricted to specialized, insular microhabitats (e.g.,
Binns, 1982; Norton, 1980), but knowledge of the potential for colonization (and redistribution
within habitats), is important, especially in studies of perturbation effects. The use of soil
arthropods as indicators of soil conditions and disturbances is in its infancy (Lebrun, 1979) but
suffers from the paradox that many responses are species-specific, yet the diversity and
inadequate state of taxonomy for most groups (see below) makes identification extremely
difficult, even for “experts.” The list of necessary reseach in biology is long, and the challenges
are many, even without entering more basic areas of physiology, functional morphology and

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genetics.

SYSTEMATICS

As has been echoed many times by ecologists and systematists alike, sound systematics is
prerequisite to sound biology and ecology ( e.g ., Wilson, 1971). This does not simply mean
having good species descriptions and monographs available. Well-corroborated hypotheses on
patterns of evolution (phylogenies) are essential in attempting to put biological attributes and
problems in an evolutionary perspective. That this “echo” has most often fallen on deaf ears can
be easily seen in the fact that despite the ubiquity and diversity of saprophagous and
mycophagous soil mites, in all of North America a single research position is devoted to study
of their systematics (Dr. V. Behan-Pelletier, B.R.I., Ottawa).

As with the biological section, I deal here primarily with those soil arachnids of most
interest with regard to soil structure and fertility, and make no attempt to summarize
knowledge of major predaceous groups. Edaphic members of Mesostigmata represent about
120 genera in 30 families (Krantz and Ainscough, in press). Currently no North American
monographic works, comparable to the European works of Karg (1971) or Ghilyarov and
Bregetova (1977), are available. However, a valuable contribution has been made recently by
Krantz and Ainscough (in press), who provides generic keys and references. Dondale (in press),
Edgar (in press) and Muchmore (in press) have provided keys and guides to the literature for
soil spiders, harvestmen, and pseudoscorpions, respectively. Of particular importance is the fact
that the long-neglected erigonine linyphiid spiders (Micryphantidae) which are abundant and
diverse in soil litter, are currently receiving attention (e.g., Millidge, 1983).

Saprophagous and Mycophagous Soil Mites: Descriptive Taxonomy and Monographs

Kethley (in press) has provided a family key and comprehensive reference list for soil
Prostigmata, but no North American monographs comparable to those for the Palearctic fauna
(e.g., Schweizer and Bader, 1963; Ghilyarov, 1978) currently exist, although a few families are
known in some detail, at least at the generic level. Many species-level determinations even in
common mycophagous groups such as Eupodidae, Pygmephoridae, Scutacaridae,
Tarsonemidae, and Tydeidae are impossible. Of the approximately 14,000 described species of
Prostigmata in the world, Kethley (in press, and 1982) suggested that nearly 6,400 (678 genera
in 57 families) are associated in some way with the soil/litter community, and that less than
100 are mycophages. For those familiar with the true diversity of the aforementioned families
in soils, it is obvious we have a long, long way to go in descriptive taxonomy.

Oribatid mites are perhaps the most successful of all soil arthropods (Johnston, 1982). The
approximately 6,500 known species-group taxa represent more than 1,000 genera in about 150
families. As in most other mite suborders, the known species constitute a small fraction of the
extant number. For this group also, there are no monographs for the North American fauna, or
any substantial part of it. In contrast, monographic works on the Palearctic fauna steadily
appear (e.g., Sellnick, 1928, 1960; Willmann, 1931; Bulanova-Zachvatkina, 1967; Kunst, 1971;
Ghilyarov and Krivolutsky,1975; Suzuki, 1978; Niedbala, 1980; Balogh and Mahunka, 1983).
Whereas careful use of these works can be helpful in identifying the nearctic oribatid mite
fauna, special care must be used in assigning species names. The North American literature is
replete with wrongly applied names of European species. Especially helpful have been the
several generic-level world or holarctic monographs of Balogh (e.g., 1965, 1972), but the

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Norton

inadequate state of knowledge of the Nearctic fauna makes the distributional information in
these works misleading. Also, generic concepts in many families, developed primarily in
Europe, are not applicable to the North American fauna.

Ever since their serious initiation at the turn of the last century, in the works of Nathan
Banks, descriptive studies on North American oribatid mites have been the domain of only one
or two productive researchers, and the quality of work has varied tremendously. All of this will
be referenced in a catalogue of oribatid mites of Canada and the continental U.S., which is
nearing completion by Drs. V.G. Marshall, R.M. Reeves and me. It lists approximately 1,000
species-group taxa and will be especially important as a guide to the literature. It does not
substitute for much-needed monographs, however, and as in other soil mite groups, the
taxonomy of North American oribatid mites is not yet “user-friendly.”

One problem with most mite monographs is that they do not deal with immatures. Whereas
in most Mesostigmata and many Prostigmata the immatures and adults are easy to associate,
this is not so for the brachypyline, or “higher” oribatid mites, which are the most abundant and
diverse groups in most soil extracts. If extractions are efficient, immatures are obtained in high
numbers and much information is lost by lumping them as “oribatid nymphs.” The only key
available for immatures is that of Wallwork (1969), and this is necessarily very general and
incomplete. Although the importance of immatures in systematics and ecology has been
stressed ( e.g Grandjean, 1953; Trave, 1964) relatively few researchers make the effort to
describe them.

Supraspecific Classification and Phylogeny

Before concluding, a few statements should be made about trends in the classification of soil
arachnids, and the general philosophical issues behind them. Historically, most classifications
have been based on differences between groups of organisms and little distinction was made
between classification and identification, which should be quite opposite procedures.
Classifications have tended to be pragmatic mirrors of identification keys, but the price paid for
such a simple translation is loss of the evolutionary perspective. Like keys, classifications based
on differences, tell us nothing about evolutionary relationships, which can only be deduced from
patterns of similarities between groups. As an example, let us examine the recent classification
of enarthronote oribatid mites by Balogh and Mahunka (1983) (their Arthronota). Two cohorts
are recognized within this group, the Euarthronota and Arthroptyctima. The first has nine
superfamilies, all with a single family. The families are so separated because of discrete
morphological differences. The Arthroptyctima has two superfamilies, each with a single
family. While this classification could readily be translated into a key, it carries essentially no
evolutionary information. The Arthroptyctima is biphyletic; the character on which the
grouping is based (ptychoidy) is clearly derived by convergence so that the closest relatives of
each of the two superfamilies are not each other, but are in different superfamilies of
Euarthronota (see Norton et al., 1983; Norton, 1984). Also, even though there is much to be
learned about relationships in the families constituting Balogh and Mahunka’s Euarthronota,
some relationships are easily defined (Norton et al., 1983; Norton, 1984) yet completely
masked by their excessively split, redundant classification. Scientists who are seeking an
evolutionary understanding of biological processes, such as the distribution of feeding strategies
and their effects on soil structure and fertility, or adaptations to desert conditions, or whatever,
are thus done a disservice.

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535

NO. OF SPECIES-GROUP TAXA

Fig. 1 . Present distribution of species-group taxa among genera of oribatid mites (worldwide).

The effects of emphasizing differences can also be seen in the distribution of species and
subspecies among genera (Fig. 1). Almost half (47%) of the approximately 1,000 proposed
oribatid mite genera are monotypic and three-quarters have fewer than five species-group taxa.
The trend is also growing; in 1980, the monotypic genera constituted 43% of the total. Clearly
the reason is an emphasis on differences for purposes of identification and results in the same
loss of evolutionary information.

Classifications based on similarities, in particular similarities which are evolutionary
“novelties,” are becoming more popular. Such classifications require more thought and effort.
They are based on the development of testable hypotheses of evolutionary patterns, or
phylogenies, the techniques for which are grouped under the rubric “cladistics” or
“phylogenetic systematics” (see Wiley, 1981). It is not always possible or even necessary to
develop phylogenies when doing descriptive work, but the principle of “classification by

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Norton

similarity” can be adhered to, nonetheless.

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Schools Jap. 56: 121-175, 34 pis.

Swift, M.J., O.W. Heal and J.M. Anderson. 1979. Decomposition in terrestrial ecosystems.
Univ. California Press, Berkeley.

Theron, P.D. 1979. The functional morphology of the gnathosoma of some liquid and solid
feeders in the Trombidiformes, Crytostigmata (sic) and Astigmata (Acarina), pp. 575-579.
In: Piffl, E. (Editor). Proc. 4th Int. Congr. Acarology Akademiai Kiado, Budapest.

Thomas, J.O.M. 1979. An energy budget for a woodland population of oribatid mites.
Pedobiologia 19: 346-378.

Trave, J. 1964. Importance des stades immatures des oribates en systematique et en ecologie.
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organization of communities of soil arthropods. Pedobiologia 23: 126-144.

Wallwork, J.A. 1958. Notes on the feeding behavior of some forest soil Acarina. Oikos 9:
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Wallwork, J.A. 1967. Acari, pp. 363-395. In: Burges, A. and F. Raw (Editors). Soil Biology.
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Wallwork, J.A. 1969. Some basic principles underlying the classification and identification of
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Wallwork, J.A. 1983. Oribatids in forest ecosystems. Ann. Rev. Entomol. 28: 109-130.

Walter, D.E. and R.A. Norton, (in press). Body size distribution in sympatric oribatid mites
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Webb, D.P. 1977. Regulation of deciduous forest litter decomposition by soil arthropod feces,
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Wiegmann, G. 1979. On the life cycle of oribatid mites as observed in the laboratory and in the
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Young, S.R. and W. Black. 1980. Some factors affecting metabolic rate in an Antarctic mite.
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Zinkler, D. 1971. Carbohydrasen streubewohnender Collembolen und Oribatiden, pp. 329-334.
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Quaest. Ent., 1985,21 (4)

BIOLOGICAL AND SYSTEMATIC PROBLEMS INVOLVING SOIL DWELLING

ARTHROPODS

R.L. Hoffman
Biology Department
Radford University

Radford, Virginia 24142 Quaestiones Entomologicae

U.S.A. 21:543-557 1985

ABSTRACT

Study of structure, classification, and way of life of myriapods is still in a threshold
position, and ground is being lost rapidly because current researchers are not only numerically
fewer than in the past, but are less productive. Ironically, at the same time, their research
potential is becoming appreciated, and a rapidly increasing interest in these animals is being
shown by ecologists and other biologists. Regrettably, a number of important major
discoveries in myriapod biology during the past 30 years have been left fallow, after their
discoverers died or turned to other subjects. The present deficiency in alpha and beta
taxonomy has had a negative effect on other areas of research: it is not very useful to
investigate organisms which are nameless and unclassified. The obvious solution to the
problem is to increase the number of systematists and provide the necessary outlets for major
revisionary studies. If necessary, funds should be diverted from well-known but still
intensively studied groups like terrestrial vertebrates and angiosperma, and allocated to
inadequately known and even less-studied organisms of the soil, on which all terrestrial life
ultimately depends.

RESUME

Certes on a souvent dit que la taxonomie, la morphologie et ihistoire naturelle des Myriapodes sont encore dans leur
enfance, et que ces domaines prennent meme actuellement du recul parce que les chercheurs y sont moins nombreux et
moins productifs que par le passe. Ironiquement par contre, les ecologistes et aurtes biologistes montrent un interet
croissant pour ces animaux, dont ils realisent le potentiel en matiere de recherche. II est fort regrettable que plusieurs
percees importantes dans I’histoire naturelle des Myriapodes ayant eu lieu au cours de 30 dernieres annees n’aient point
connues de suite apr'es que leurs decouvreurs soient decedes ou aient change de champ d’etude.

La deficience actuelle en taxonomie alpha et beta a evidemment un impact negatif sur les autres domaines de
recherche: il n’est pas tres utile d’etudier des organismes qui ne sont ni nommes ni classifies. La solution evidente h ce
probleme est d’augmenter le nombre de systematiciens etudiant les Myriapodes et d’offrir les debouches necessaires pour
d’importants travaux de revision taxonomique. Si cela s’avbre necessaire, des argents support ant presentement des
travaux sur des groupes bien connus mais malgre cela encore sur-etudes, tels que les Vertebres et les Angiospermes.
devraient etre redistribues pour soutenir des projets d’etude de groupes mal connus et encore negliges d’organismes du
sol. desquels depend ultimement le maintien continu de toute vie terrestre.

INTRODUCTION

It is common for scientists in virtually any discipline, no matter how sophisticated it may
have become, to deplore deficiency of knowledge in their specialty. I suspect, however, that the
participants in this symposium are, by virtue of their own experience, disposed to accept a
general apologia that existing knowledge about myriapods is strickingly deficient and
fragmentary. It is certainly not an overstatement to note that myriapodology - and such a word

544

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is not even in dictionaries - is presently at the same level of development as was entomology
about 1850, or ornithology about 1800.

Many persons whose knowledge of Myriapoda derives from standard texts published in the
recent past, consider this group to include few orders of insignificant classes, and are surprised
to learn that diplopods alone constitute 15 orders and about 115 families. There is more to the
subject than Julus, Spirobolus, and Lithobius, and the major problem that I had to face on
being invited to summarize current “state of the art” for the four myriapod classes was how to
do it meaningfully in 30 minutes. I have little confidence that such a goal can be achieved, even
with rather superficial coverage. Three areas will be considered: present state of knowledge of
myriapods; problems impeding an improved knowledge of them; and what is presently known
about the impact of myriapods on the formation and characteristics of soil. Even for
inadequately known taxa, this is a large order.

PRESENT STATE OF KNOWLEDGE OF MYRIAPODS

Systematics

To begin with, even the phylogenetic relationships of the classes Diplopoda, Chilopoda,
Pauropoda, and Symphyla to each other and to the hexapod tracheates are far from being
established. Much has been written on this point since about 1887, when R.I. Pocock
established the first “modern” arrangement, aligning Diplopoda, Pauropoda, and Symphyla in
a group Progoneata, and the Symphyla and Insecta in the coordinate Opisthogoneata. The most
extensive recent work, and by far the most authoritative, has been that of S.M. Manton whose
approach to the postulation of phylogeny was based largely upon comparisons of structural and
functional aspects of locomotory systems. Without wishing to denigrate in any way the superb
research conducted and published by Manton (1954-1977) with exceptional illustrations, I feel
that her conclusions were seriously flawed by reliance upon an outdated classification (that of
Attems, 1926, which was actually written near 1920), and by insufficient consideration of
adaptive convergences. In particular, I cannot accept the notion that “myriapods” comprise a
monophyletic entity coordinate to a comparable “hexapod” group as separated solely by a
difference in mode of mandibular articulation. Single character differences between taxa do not
inspire much confidence when they oppose groupings made on the basis of extensive similarities
in numerous character-systems. I prefer to recognize an indivisible spectrum of tracheate
classes, which awards class rank for collembolans, proturans, diplurans, thysanurans, and
pterygotes, and which admits the numerous shared characters of diplurans and symphylids. I
am not aware at the present of any convincing arrangement of these five hexapod and four
myriapod classes into higher groups ( e.g ., superclasses or subphyla). Depending on which
character systems are stressed, any number of classifications could be devised, including one
that sets Diplopoda apart in a sister-group relationship to the other eight combined. The fossil
record has, so far, shed very little light on this problem.

Initially, “myriapods” were studied by general zoologists, then - up to about 1900 - by
entomologists. The primary taxonomic characters of both pauropods and symphylids are chiefly
those of chaetotaxy and subtle modifications of the integument, and except for the advantage of
improved optical equipment, the techniques involved in their study have changed but little in
the past century. Similarly, the study of lithobiomorph and scolopendromorph chilopods still
follows classical procedures of the last century (enumeration of spines, spurs, and sutures). But
a fundamental change occurred in classification of geophilomorph centipedes around 1870,
with Meinert’s discovery that the best familial and generic characters reside in mouthpart

Soil dwelling arthropods

545

structure. This realization instantly rendered all previous work on these animals obsolete, and
mandated the eventual restudy of early geophilid types. A similar revolution in milliped
classification was triggered in 1884, when Robert Latzel made extensive and effective use of
male genitalia to distinguish both genera and species in the central European fauna. Genitalia
had been sporadically described, and even illustrated, since 1832, but Latzel’s consistent and
comprehensive emphasis of these appendages was catalytic. Virtually all millipeds named prior
to Latzel’s time require redescription with respect to genitalic structure.

As the result of these important discoveries by Meinert and Latzel, generation of myriapod
specialists emerged around 1890, some of its members being converted entomologists, some
innocent of any previous tradition. A cadre of six dynamic young men working chiefly in the
1890s built the foundations of our existing classifications of the various myriapod groups,
except for the Pauropoda and Symphyla. Working most of the time in isolation, some of them
adopting inappropriate attitudes about taxonomy and nomenclature, they also provided a
heritage of confusion, duplication, and outright systematic anarchy that by 1950 had attained
epic proportions. Most of these pioneers endeavored to study the world fauna of both chilopods
and diplopods (often the other two classes as well), and if their work was adequate in one class,
it was usually catastrophic in the other. Recitation of the problems generated during this period
would fill a volume, and a large part of modern work consists of tedious corrective surgery.

During this period, about 60 years in duration, of intense descriptive work, emphasis was
placed on alpha taxonomy of the crudest sort, usually the naming of material in regional
collections. Some of the most productive workers seemed to operate on the principle that the
mere naming of taxa, without a word of comment, was the pinnacle of taxonomic achievement.
It was not until the global catastrophe of World War II that this period came to an end,
coincidentally with the demise of most of its major figures.

To illustrate the rather spectacular growth during this period I can provide two illustrations
from the Diplopoda, the group I know best. The first is a table of higher taxa recognized at
various time intervals from 1847 to the present. The figures are not absolute, as they do not
take into account existing taxon names regarded as synonyms by the various authors cited.

Another way to show the same trend is with a line graph (Fig. 1) showing the increase in
number of generic names cumulatively, without prejudice as to their actual status.

The almost explosive increase, beginning in the 1890s, is not much different from that in
other major taxa, but begins much later than most, and represents the astonishing productivity
of three persons: Carl Attems, K.W. Verhoeff, (1926-1932), and R.V. Chamberlin, who among
them proposed no fewer than 1 199 genera. One notices that the curve begins to level off after
1950, but this is purely a result of changing times and not a depletion of undescribed genera.
Actually, two things have conspired to dampen the growth rate. First is a post-war change in
taxonomic philosophy, from sheer mindless description of novelties as an end in itself to a
strong emphasis on clean-up work: restudy of old types, preparation of whatever revisions could
be managed, and so on. Second, and perhaps more compelling, has been the incredible increase
in the costs of publication. (In these days of near-universal page-charges, it is refreshing to
recall that Verhoeff, for instance, was actually paid - so many words per mark - by the
Zoologischer Anzeiger and other German journals. Today only a millionaire could afford to
publish the typical Verhoeffian output of several hundred pages per year.)

I believe that we have so far described about 20% of the actual milliped fauna of the world.
If this figure be true also for the other three classes, a sum total of more than 100,000 myriapod
species must be reckoned with.

Quaest. Ent., 1985, 21 (4)

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Hoffman

2500

>000

1500

1000

500

Fig. 1. Cumulative increase in the number of generic names in Diplopoda, including synonyms & homonyms (1758-1980).

Table I

Increase in the number of higher taxa, Class Diplopoda

Reference

Orders

Families

Genera

Gervais, 1847

Bollman, 1893

Cook, 1895

190

Silvestri, 1897

353

Attems, 1926

621

Hoffman, 1980

115

1701

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547

The only comprehensive treatment of the classfication, structure, and way of life of all four
classes is that of Carl Attems, in the Kukenthal-Krumbach Handbuch der Zoologie (1926),
which was written more than 60 years ago. The taxonomic part is of course hopelessly out of
date, and was seriously flawed even at the time it was written, but for many taxa it still remains
the only existing reference.

For Diplopoda, two recently published manuals are useful. One is a catalog of all generic
and familial group names, with their type species, published from 1758 to 1957 (Jeekel, 1971).
The other is a classification of the world fauna down to the level of subgenera, compiled by me
(Hoffman, 1980). It contains no keys nor descriptions, but does include reference to all
post- 1926 synoptic taxonomic papers.

The enormous order Polydesmida was surveyed by Attems in three big volumes of the
Tierreich series (1937-1940), but these works are chiefly useful from a bibliographic sense,
being mostly compilations severely handicapped by their author’s ultraconservative taxonomic
philosophy. At least all polydesmoids described up to that time are included somewhere, and
Attems’ real contribution was to provide a beachhead for further, more refined studies. In
recent years, some work, reminiscent of the labors of Hercules, has been conducted by a few
hobbyists. The Paradoxosomatidae, largest family of the entire class, has been under study by
C.A.W. Jeekel since about 1950. This author published a provisional classification of the group
in 1968, as well as numerous generic synopses and clarifications of nomenclature, but his
intended goal - a new revision of the entire family - is still a long way in the future. Since about
1955, I have been working in a similar way on the larger chelodesmoid families, e.g., the
Chelodesmidae, Oxydesmidae, and Gomphodesmidae. Although revisions of many genera and
tribes have been published, only the African family Oxydesmidae is now actually at the stage of
preparation for publication. The Chelodesmidae will doubless prove to be the largest family of
Diplopoda: already more than 20 tribes and 200 genera have been defined even though the
fauna of Brasil has scarcely yet been sampled. The family Xystodesmidae, virtually endemic to
North America, is being worked up one genus at a time, beginning with the rich Appalachian
fauna, by R.M. Shelley (e.g., Sigmoria, 1981). But the numerous families of “smaller
polydesmoids” have received essentially no attention and at present nobody has either the time
or interest to study them despite their importance in soil samples from any tropical region.

In the order Chordeumatida, characterized by a large number of mostly small disjunct
families, some progress has been made chiefly on the Nearctic fauna by W.A. Shear, who has
revised the Conotylidae (1971), the Cleidogonidae (1972), Rhiscosomididae (1973), and
Tingupidae (1982). Dr. Shear advises me, however, that in less than a decade so much new
material has accumulated as to render his cleidogonid monograph obsolete. Other students of
this order, notably S.I. Golovatch and J.-P. Mauries, have published descriptive papers on the
Old World fauna but do not appear to be contemplating comprehensive revisions. The study of
this order is greatly impeded by the scarcity of material; a great many species still remain
known only from the type series named decades ago.

The large, mostly tropical species of the order Spirostreptida have been recently, and
adequately, summarized: the Spirostreptidae itself by Krabbe (1982), the Harpagophoridae (in
part) by Demange (1961 et seq.), and the Odontopygidae by Kraus (1960, 1966). These large
I and useful papers go far to setting in order the classification of the three families, but still
| represent only a first step, and none of the many genera involved have yet been the subject of a
i “modern” revision. The cambaloid members of this order remain in a state of substantial
confusion, with little agreement even about the definition of families, but the group is being

il Quaest. Ent., 1985,21 (4)

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Hoffman

studied by Mauries and it is hoped that a revisionary monograph may be forthcoming in a few
years.

Species of the related order Julida remain in a sort of limbo. The family Parajulidae, which
is virtually endemic to North America, was studied for many years by N.B. Causey, but despite
appreciable research nothing useful was published before Dr. Causey’s death in 1979, and no
one has touched the group since. A more optimistic statement can be made about other juloid
families, which are now being investigated by Henrik Enghoff. It is the intention of Dr. Enghoff
to eventually reorganize the classification of the entire order, and toward this end a number of
preliminary studies have already been published.

Lastly, in the order Spirobolida, the family Spirobolidae was monographed in a very
adequate way by W.T. Keeton in 1960. This group is in a good condition for detailed
biosystematic studies of individual genera. The other spiroboloid families - such as have been
adequately defined - remain in complete chaos, and identifications of rhinocricids, pachybolids,
and trigoniulids are virtually impossible to make.

Many families, especially in the Palearctic region, are monobasic or nearly so, and their
revision would entail only careful studies of structural features and comparisons with related
taxa. Omitting such groups, and in summary, less than 10 families of Diplopoda have been
recently treated taxonomically in a way useful to beginners, e.g., with keys, diagnoses,
illustrations, synonymical lists of species, maps, and other features normally taken for granted
by students of most other animal groups.

From a faunistic standpoint, the record is not much better. Checklists are available for
North America (Chamberlin & Hoffman, 1958) and Mesoamerica (Loomis, 1968); both are
not considerably outdated. National surveys are available for Great Britain (Blower, 1955, and
in press), Germany (Schubart, 1934), France (Demange, 1981), India (Attems, 1936), and
Japan (Miyosi, 1959). One of the best-known parts of the world for diplopods is the Union of
South Africa, thanks to the work of Attems (1928, 1934), Schubart (1956, 1958, 1966), and
Lawrence (numerous papers, e.g., 1953a and b, 1967). A few unlikely parts of the world have
been treated faunistically, such as the island of Hispaniola (Loomis, 1936) and Panama
(Loomis, 1964).

Centipeds are probably even more inadequately-known than millipeds. A catalog of generic
names and their type species has been compiled by C.A.W. Jeekel but not yet published, and
there is no classification of the Chilopoda in toto since 1926. The order Geophilomorpha was
treated in the Tierreich series by Attems (1929) and the Scolopendromorpha by the same
author a year later (1930). Aside from being decades out of date, both of these manuals were
largely compiled from faulty literature and were inadequate the day they were published. The
content of both orders has virtually doubled in the past fifty years, with no reliable update. The
enormous and difficult order Lithobiomorpha has not been treated comprehensively, nor has
the much smaller Scutigeromorpha.

Regional papers have been published for Great Britain (Eason, 1964), France (Brolemann,
1935, Demange, 1981), and South Africa (Attems, 1928). The Lithobiomorpha of the Soviet
Union was treated by Zalesskaja (1978) and the North American species of this order were
covered in an excellent series by R.V. Chamberlin (1913-1925). Unfortunately, the good start
embodied in the last-cited reference was promptly subverted by a long sequence of
unsatisfactory “descriptive” papers by the same author during the following 30 years. The often
cryptic synonymy and nomenclature of lithobiids has been clarified over a period of time by
E.H. Eason, who hopes to prepare a world catalog for this large and difficult family. A good

Soil dwelling arthropods

549

start was made toward reclassification of Geophilomorpha by R.E. Crabill during the years
1960-1968, but regrettably no major synthesis was published before his retirement in 1983.
Recent, outstanding work on this order is being published by L.A. Pereira, who expects to
revise initially the family Schendylidae, and eventually other geophiloid taxa as well. The
chilopod fauna of southern Europe, particularly Italy, is being studied by A. Minelli.

No optimistic statement can be made about the classes Pauropoda and Symphyla. At
present, both of them are virtually the exclusive domain of Ulf Scheller. The scarcity of good
material in both groups, and the very fragmentary geographic representation, renders
revisionary studies almost impossible. Scheller’s faunistic studies, however, are models of
excellent presentation and include as much group taxonomy as can be managed. So many
pauropods are cosmospolitan or nearly so, that a world synopsis of this class is necessary for
adequate work, and at present this can be gained only by knowledge of the entire published
taxonomic literature in the original. It is possible that Dr. Scheller will prepare a catalog of the
species of one or both classes.

Concluding this somewhat discouraging summary of myriapod classification at the present
time, a glance at the number of current active specialists cannot fail to give an even gloomier
prospect for the future:

Chilopoda: England, 2; France, 2; Italy, 1; Australia, 1; Argentina, 1; U.S.S.R., 2; total: 9.

Diplopoda: U.S.A., 3; France, 2; Denmark, 1; Germany, 1; U.S.S.R., 2; Japan, 2; Holland,

1; total: 11, two of which are duplicated in the chilopod list.

Pauropoda: Sweden, 1; Austria, 1; Germany, 1.

Symphyla: Sweden, 1.

Most of the foregoing specialists are either teachers or curators; in either case, their research
time is limited (or outright stolen from primary obligations). Nearly half of them are nearing
the end of their productive years. All are innundated with material, and years behind on
projects and gratuitous identification work. At most, only about five persons are relative
newcomers to myriapod taxonomy.

Morphology

What can be said of taxonomy’s sister science, morphology? Outstanding anatomical studies
have been made in recent years by Demange and by Manton. The latter author dealt primarily
with integumental and musculature modification associated with locomotion. Demange
published an outstanding study on thoracic segmental musculature in 1967, with many
profound implications (some of them controversial). I do not know any subsequent researches
extending, confirming, or refuting the findings of these two pioneers. It cannot be said that the
study of even the general aspects of structure of myriapods has been exhausted, and I cite a few
examples. (1). A good comparative study of the head capsule amongst diplopods has not been
published, nor has an attempt been made to homologize head musculature with that of body
segments. (2). Species in several spiroboloid families have paired paramedian dorsal pits on
each segment, of totally unknown function. (3). In the family Paradoxosomatidae, many
species have glands opening through paired pores on the 5th sternum: such glands have not
been mentioned by anybody and I suppose have been overlooked to the present. Obviously their
function likewise remains unknown!

For Chilopoda, at least, the areas of ignorance have been categorized in Dr. John Lewis’s
recent (1981) book on centiped biology; someone seeking structural, developmental, or
ecological problems can find one on nearly every page. Some come at once to mind. (1). Many

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Hoffman

geophiloids have conspicuous sclerotized sternal pits, much used in taxonomy but of totally
unknown function. (2). What is the function of the Tomosvary Organ? (3). What is the
function of coxal pores in lithobiomorphs?

Only within the past two decades has anything been done of note with the neurosecretory
structure of centipeds (or millipeds, for that matter). The same time period has seen the
initiation of work on microstructure of muscles, of sperm cells, of sensory organs. As many as a
dozen papers have been published in these areas. But since something has to be skimmed over
in this review, the cut is in structure: there is much to cover yet.

Embryonic development of millipeds was first studied in the last century by Metschnikoff,
Newport, and Heathcote. Several papers were published by Silvestri ( e.g ., 1950), Pflugfelder
(1932), and most recently and thoroughly, by Dohle (1974). Details of development for many
orders remain unknown, including those for the exceptional group Stemmiulida in which the
young eclose with 19 segments instead of the six common to all other diplopods. Demange has
observed that embryos of most groups reveal little information about phylogeny because many
critical structures do not appear in the early stages. Yet, there is plenty of opportunity for a
student to make a distinguished career in this area.

Much happens after hatching. In many milliped groups, the male genitalia begin to modify
from normal walking legs early in the stadium sequence, becoming larger and more specialized
with each moult. In polydesmoids, however, the final moult changes a small knob-like
primordium into a mature gonopod of often fantastic complexity. Nobody has sectioned
specimens during this diapause period to follow the sequences of events, to determine what
pattern may exist comparable to the mechanisms that direct the reorganization of
holometabolous insects during pupation. Development of the modified posteriormost legs of
male lithobiomorph centipeds has not been studied, either.

Way of Life

The foundations of present knowledge about myripod way of life were laid down chiefly by
K.W. Verhoeff, who studied the Palearctic fauna for half a century. Verhoeff (1926-1932)
worked out the life histories of many kinds of millipeds and centipeds, and discovererd the
interesting phenomenon that occurs in various kinds of julids: non-mating intercalary adult
males (“Schaltstadium”) which moult into a sexually active stage. This subject has been
carefully studied in England by J.G. Blower and some of his students, and in France by F. Sahli
(e.g., 1969). In general, postembryonic development, particularly of julids, occurs in a number
of remarkable patterns, in many taxa with stadia omitted or added. Blower’s group has also
worked on population structure, phenology, and general natural history of various British
millipeds, and provided a fine model for those who might wish to study the fauna of other
regions (e.g., Blower & Gabutt, 1964; Blower & Miller, 1974). Fundamental work on way of
life of Ommatoiulus moreleti, an Iberian julid introduced into South Africa and Australia, is
being conducted by G.H. Baker (1978a-c).

Details about life history have been published for only two North American millipeds (and
no centipeds), and these are not comparable to the precisely executed studies of European
investigators. A few papers have referred superficially to habitat preferences of American
species, contrasting with the careful work of J.-J. Geoffroy (1981) on the French fauna.
Interactions of myriapods with other organisms and with their environment have rarely been
better-accounted than in R.F. Lawrence’s notable book about South African soil fauna (1953).

Soil dwelling arthropods

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Diplopods were considered to practice only the most perfunctory kinds of reproductive
behavior. During the past two decades, publication by Ulrich Haacker (1969), in Germany,
reported fairly sophisticated courtship practised by some julids, the males of which preferred an
attractive secretion from the base of the 2nd pair of legs, which attracted (and distracted)
females which fed upon the material whilst the male inobstrusively effected copulation and
sperm transfer. Haacker (1971) also reported apparently similar glands located middorsally on
the terga of several European chordeumatids, but was not able to observe their actual use. In
other studies (1968) he taped and analyzed the stridulation of South African sphaerotheriids,
produced by males as an element in courtship. Lamentably, this gifted investigator died at an
early point in his career, and nobody has since continued along the trail he blazed so well.
Regrettably, detailed studies of reproduction have not been reported for a single North
American milliped. The considerable body of published field and laboratory observations has
not been organized for second-stage, follow-up work. One facet that merits careful study is the
sociality of platydesmid species, represented most conspicuously in the United States by the
genus Brachycybe. These animals tend to live in large aggregations of all stages, and in such
colonies specimens are often seen in a stellate arrangement, heads together, bodies radiating out
like spokes, for a still-unknown reason. In this genus, large numbers of tiny yellow eggs are
released by the females, then gathered up and brooded by males, an exceptionally rare
occurrence among anthropods. The phenomenon was observed by me in North Carolina in the
summer of 1958; by an astonishing coincidence it was published in the same year by Y.
Murakami for a Japanese species of Brachycybe. Careful studies remain to be made for other
platydesmid genera in North America and the Mediterranean region. Do they share this trait?
How could such deviant behaviour have developed?

Males of many diplopod taxa, particularly polydesmoids, are provided with a complex
arsenal of secondary sexual modifications of legs, sterna, and mouthparts. How such equipment
is used remains completely unknown, and could be elucidated by just the simplest observation
of mating pairs. Some is obviously involved in clasping the female, some, involving internal
glands and their pores, must perform an attractant function. Mauries (1969) described the
mating behaviour of Typhloblaniulus lorifer , in which coupling and positioning of the female is
achieved by intertwining of the bodies, by the female biting the modified 1st legs of the male,
and by the female’s antennae being clasped by a modification of the male’s mandibles. Species
of the allied family Parajulidae occur in abundance over much of North America, adults
exhibit a wider variety of sexual modifications, and yet not a single observation has been
published on reproductive aspects of this big family. There is also a capital problem involving
Aenigmopus alatus , Guatemalan polydesmoid males, which lack gonopods: how does it
accomplish sperm transfer? This species is known so far only from type material, but a precise
locality is known and it should be possible to obtain living specimens.

Prior to about 1957, virtually nothing was known about the mating behaviour of chilopods.
Using infra-red light for observations, H. Klingel solved this riddle and reported his findings in
several papers ( e.g ., 1957, 1960). Apparently little has been done since that time. It is
well-known that the males of numerous American lithobiomorph genera have the last pair of
legs modified in curious ways: a spectrum of knobs, crests, cavities, hair tufts, and pore fields.
Could not some student of behaviour adopt Klingel’s techniques to see what role these strange
modifications play in mating? Do females recognize corresponding males tactily?

It has been known for years that millipeds produce a variety of caustic and/or aromatic
secretions when disturbed, the odours being variously reported subjectively as like camphor,

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Hoffman

almond extract, osmic acid, quinine, creosote, and rotting sponges. A few chemical analyses
were made during the first half of this century, but scientific studies on allomones were really
first initiated by Thomas Eisner about 25 years ago. Eisner investigated not only the chemical
composition of these secretions but their biological functions as well. Aside from the obvious
role of predator deterrents, most of the secretions are markedly fungicidal, suitable for
organisms which live in damp biotopes (Eisner, 1970). The structure of the ozadenes was
worked out by D.W. Alsop in Eisner’s laboratory, but details have, to the best of my knowledge,
not yet been published. Biosynthesis of benzaldehyde and hydrogen cyanide, common
ingredients in polydesmoid allomones, was worked out by Duffey, Underhill & Towers (1974)
in Harpaphe haydeniana, a common species in British Columbia. Substantial progress was
made at the University of Georgia (cf. Duffey, 1977) toward possible chemotaxonomic use of
allomones, but once again, a promising start soon faltered and nothing is currently being done
along these lines. Existing evidence suggests a fairly close correlation between allomone
structure and established taxonomic groups.

Some millipeds are known to be luminescent, a phenomenon especially well developed in
some Californian xystodesmids, reported by Davenport (1952), but with inconclusive evidence
about the cause. Some geophilomorph centipedes emit a phosphorescent secretion from sternal
glands, but to what end remains unknown. Most geophilomorphs are some shade of yellow,
brown or red. The small species of the tropical family Ballophilidae, however, depart from this
norm in their colouration: bright blue, violet, purple, green, and black species are known.
Ballophilids are characterized in part by having the sternal glands open onto a midventral
sternal knob, and in fresh specimens the glands can be easily seen as clusters of intense
pigmentation through the more dilute colouration of the integument. What is different about
ballophilids and their sternal glands? No one has any idea. Not even the crudest histochemical
assay has yet been attempted.

The foregoing enumeration of some areas of ignorance has largely avoided mention of either
pauropods or symphylids. It is hardly necessary to add that virtually nothing is known about the
structure and way of life of members of more than one or two common European species.

IMPEDIMENTS TO DEVELOPMENT OF MYRIAPODOLOGY

I am sure that those who study mites, nematodes, springtails, or pseudoscorpions will be
surprised at little I have said so far: most soil organisms share this heritage of neglect. No doubt
all of us tend to agree that problems such as the following are serious ones:

1. Virtual ignorance of the actual fauna in many parts of the world, especially the tropics,
and frequently there is a burden of inadequate taxonomic and complex nomenclatorial
problems afflicting even the better-known faunas.

2. The likelihood that major parts of the world’s soil fauna will become extinct before it can
even be sampled. Berleseate samples now in dead storage in various museums probably contain
a number of already extinct species: fossils in alcohol.

3. Difficulty of entry into the classification and identification of most groups because the
literature is extensive, fragmentary, widely scattered, and polyglot.

4. The frequent impossibility of obtaining identifications because either there are no
specialists, or, if such exist, they are 35 years behind their unidentified backlog, or, worse,
unable to make an identification without having first to revise the genus, tribe, or family
involved.

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553

I may be forgiven my bias in believing that organisms must be described and placed in a
classification before information about them is meaningful. Taxonomy may be passe in
ornithology and some other mature fields of zoology, but I am appalled to observe how many
people are still investing vast resources of time and money deciding whether a given vertebrate
taxon is a good species, a sibling species, a subspecies, or what, when the majority of arthropods
are still unknown, uncollected, and ignored. Is it a better investment to investigate details in
vertebrates, or get on with the higher classification of other phyla?

Solutions are fairly obvious. Most of the present generation of myriapodologists drifted into
this area accidentally, and remained in active pursuit of research goals primarily as a personal
hobby, with time abstracted from career requirements and family obligations. Even museum
curating is no ideal occupation, if one is primarily responsible for the collections first, routine
identifications second, and perhaps personal research last. If more taxonomists are needed to
handle the job of working up what we have already in museum jars, some better way must be
found to employ their talents on an occupational basis. What graduate student wishes to invest
quite some years in learning the complexities of myriapod lore, if there is no hope whatever for
finding gainful employment in such a specialty? Research on structure, behaviour, and ecology
can be left to academic sectors. These are areas which can be rather quickly comprehended,
pursued, and solved in segments by graduate students. Systematic work, in my view, requires a
far longer time to master, and productivity is linked with continuity. I began the study of
millipeds as an undergraduate, as did several of my friends, but could not do so today simply
because I could not cope with publication problems. If progress is to be made in myriapod
taxonomy not only must career opportunities be guaranteed, but possibilities for publication of
taxonomic monographs must also be improved. Many of the better-known research-support
sources (I may mention the U.S. National Science Foundation) award grants on an egalitarian
basis: as much is given for studies of vertebrates (less than 1% of animal creation) as for
arthropods (more than 90%). Is it possible to redistribute the available largesse on a scale
commensurate with the actual size of the group, and its need for study? I strongly support the
principle of peer evaluation of research proposals, but appeal for reason in the process. I have
known excellent, deserving projects turned down because one or two reviewers felt that the
applicant should have introduced reference to “phenetics” or “cladistics” or some other popular
fad. In work on many groups of arthropods, we are still trying to scramble into the lower levels
of beta taxonomy. We must crawl before we fly, and to impose a requirement for theoretical
biology when there is no existing base for it, seems entirely unrealistic and counter-productive.

MYRIAPODS AND SOIL

Lastly, it is necessary to append a few remarks appropriate to the subject of this conference.
I have investigated the historical background as far as the paper by Shaler, which first
suggested a substantial role of diplopods in soil formation; also the classical texts written or
edited by Kevan, Raw, and Schaller, also recent papers by van der Drift, Gere, and other
European workers. Most publications so far relate to diplopods, and are in two categories: some
subjective field observations lacking quantitative controls; and laboratory experiments not
closely associated with natural conditions.

Two areas of actual soil influence are generally accepted: physical and chemical. The first
involves disruption of the upper layers of soil and the litter accumulation by burrowing
activities of diplopods. Many of these (which may be surface or even arboreal dwellers when

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mature) may spend all of their immature stadia burrowed fairly deeply in the soil itself: the
general collector rarely finds young millipeds in the upper horizons. Scolopendromorph and
geophilomorph centipeds likewise burrow to some extent, or exploit the burrows of other
animals. I think that either exclusively edaphic residence, or vertical circadian movement must
be accounted a substantial influence on the physical makeup of upper soil strata, although I do
not know of any work quantifying the effect. It is well-known, secondarily, that most millipeds
are detritivores and break down a lot of vegetable material (leaves, rotting wood, fungi) simply
by mechanical trituration as they feed upon it. Some earlier authors (Romell, 1935; Eaton,
1943) implicated millipeds as a major factor in mull formation, and certainly captives are able
to reduce a handful of decomposing leaves in short order, as can be confirmed by anybody who
keeps a live spirobolid under observation. But I am often amazed to sift through really large
quantities of leaf litter in apparently optimal situations without finding a single milliped of any
species, and humification proceeds apace. So far as I know, all chilopods are carnivores, and
pauropods and symphylids probably poelomicrophaghes; these groups probably contribute very
little to mechanical litter conversion.

Chemical influences are of several kinds: modification of plant material through digestion;
uptake and concentration of calcium and other minerals; release of nitrogenous compounds
from metabolic excretion; and formation of weak organic acids as the result of death and
protein breakdown. Most of these factors have been alluded to qualitatively in the literature,
but I have nowhere found quantitative studies aside from some experiments on mineral cycling
at Oak Ridge, Tennessee, by Reichle and collaborators (1965).

One possible influence of a chemical nature was suggested by O.F. Cook in 1911, but not
apparently considered by anyone subsequently. Cook, who was by profession an agricultural
botanist, believed that the allomones produced by many millipeds were capable of altering soil
composition by precipitating colloidal substances in the humus. He claimed, from personal
observations, that “ ... African forests have very slight superficial accumulation of dead leaves
and humus. The soil remains relatively open and noncolloidal, and is inhabited by numerous
species of millipeds. In the forests of tropical America ... the underlying soils are generally
much more colloidal than in Africa and the milliped population is generally sparse, or often
lacking altogether ... I pretend no knowledge whatever of this aspect of soil structure and
present Cook’s views here solely to give them circulation.

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Quaest. Ent., 1985,21 (4)

RECENT ADVANCES AND FUTURE NEEDS IN THE STUDY OF COLLEMBOLA
BIOLOGY AND SYSTEMATICS

A. Fjellberg
TromsQ Museum
N-9000 TromsQ

N OR WA Y Quaestiones Entomologicae

21:559-570 1985

ABSTRACT

Some new results from studies of collembolan feeding, reproduction, behaviour and
response to chemical pollution and mechanical habitat disturbance are presented.
Morphological variation in terms of ecomorphosis, cyclomorphosis and epitoky is discussed.
A future expansion into the fields of cytogenetics, physiology and functional morphology is
expected to accelerate taxonomic refinement of current systematics.

RESUME

L' auteur presente des donnees nouvelles sur ialimentation des Collemboles, leur reproduction, leur comportement et
leur reaction h la pollution chimique et aux perturbations mecaniques de leur habitat. II discute de variation
morphologique en termes d’ecomorphose, de cyclomorphose et d’epitokie. L’ expansion eventuelle dans les domaines de la
cytogenetique, de la physiologie et de la morpohologie fonctionnelle devrait accelerer le raffinement taxonomique de la
systematique actuelle.

BIOLOGY

Introduction

As one of the major groups of soil microarthropods, Collembola has received increasing
attention from ecologists and biologists during the few last decades. Collembola biology has
become a very complex and multifaceted field of research. It is impossible in just a few pages to
outline all aspects of current Collembola biology studies. Instead, I will select a few important
aspects of Collembola life such as feeding, reproduction, behaviour, reactions to chemical
pollution and dependence on moisture - probably the one external factor that has the greatest
effect on Collembolan life.

Feeding and nutrition

Apart from a few predaceous species, most Collembola feed upon a variety of organic
material, both detritus and living substances, such as algae, fungal hyphae and bacteria. A
considerable amount of inorganic material may pass through the digestive tract as well. Some
authors regard soil Collembola as rather generalized feeders (Anderson & Healey, 1972;
Greenslade & Greenslade, 1983). Others are of the opinion that they are more or less selective
(Hale 1967). In either event, certain food preferences - depending on species, habitat and time
of the year - are frequently reported (McMillan 1975, Vegter 1983). The coexistence of 15-20
species of Collembola in a small volume of soil may indicate that either there is a surplus of
food, or that the food is partitioned by subject or by microhabitat differentiation of the involved
species. The latter is obviously possible. Even a few centimeters of a soil profile is rarely
uniform from top to bottom, offering a variety of habitats. But it is more difficult to explain

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how nine different species of surface active Isotoma could occur together in a handful of damp
tundra soil in north Alaska, unless there is some kind of food specialization or an excess of food
(Fjellberg, unpubl.)

The uniformity of the mouthparts in large groups of Collembola has been interpreted as an
indication of unspecialized feeding (Greenslade & Greenslade, 1983). However, I think this is
an oversimplification. Although built around the same model, the actual construction of
individual parts of the feeding apparatus varies considerably, even in close systematic groups
(Fjellberg, 1984a, 1984b). And what else other than food or feeding specialization could lie
behind this quick adaptive evolution? Or, to put it another way: if two related species shared
the same food, why should they differentiate their feeding apparatus? I believe we still have a
very crude impression of how and on what individual Collembola species feed in natural
habitats, and how they interact with other members of the community. It is a great challenge to
students of functional morphology and to persons who want to develop techniques for studying
Collembola feeding both in laboratory and in nature.

Reproduction

As soon as the individual can keep itself alive by feeding, it must keep the species alive by
reproduction. In understanding the reproductive biology of Collembola, we need information
about life cycles, recruitment, mortality and other parameters. The size distribution of
individuals in field populations - measured at various times during the year - has been used for
a long time to obtain basic information about life cycles under natural conditions (Agrell, 1941;
Fjellberg, 1975; Addison, 1981). A closer demographic analysis, involving some mathematical
modelling, may also provide information about recruitment and mortality, which are essential
factors in the energy budget of a population (Hale, 1980; Straalen, 1982, 1983). Data about
fecundity, the potential number of eggs produced by a female, has accumulated from a number
of laboratory studies. From these studies, individual fecundity appears to be rather flexible,
influenced by population density (crowding), age of the individual, temperature, substrate, and
other things (Hutson 1978, Snider 1973, Snider 1983). Concerning life cycles and longevity, it
is clear that the long arctic winter arrests development and delays reproduction until
individuals are 1-2 years old (Fjellberg 1975, Burn 1981, Addison 1977, 1981). Mature
specimens may live for several (3-7) years and may reproduce several times. A winter diapause
is demonstrated in several species, and is supposed to be essential for synchronization of the
spring reproduction of adults in the European species Hypogastrura socialis (Uzel). This
species has a rather fixed reproductive pattern. Adults reproduce only once in spring, and die
shortly after. Pheromones appear to be essential for the group behaviour of this species (see
later) (Leinaas, 1983b).

Collembola living in cold environments have clearly adopted an opportunistic reproductive
strategy. Overwintering may occur in any stage of development and life cycles are adjusted to
physical conditions of the habitat, which sometimes gives different life cycles in different
cohorts of the population (Addison, 1977) or in different populations along a microclimatic
gradient (Tamara & Mihara, 1977).

In temperate and warmer regions the generation time is shorter and reproduction runs more
freely, giving a very complex age structure with indistinct cohorts (Petersen 1980, Tanaka
1970).

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Behaviour

The study of behaviour and behavioural ecology of Collembola will probably receive
increased attention in the future. At first consideration, the behaviour of a springtail may seem
odd and bizarre. However, with closer inspection, we usually find that the behaviour is very
reasonable and part of the solution to fundamental biological or ecological problems. Bretfeld
(1970, 1971, 1976a, 1976b) described the rather complex mating systems and sexual
interference in different species of sminthurids. Although the Collembola do not have a direct
copulation and sperm transfer, the mating behaviour is probably part of the isolating
mechanisms between species, just as in other groups of arthropods.

The mass occurrence of Collembola - especially on snow - is frequently noted. Less often
seen, but probably of greater significance, is the aggregated occurrence of Collembola in soil
and litter, where they sometimes form dense colonies of millions of individuals. We have now
gained some insight into the mechanisms regulating this particular behaviour. Verhoef et al.
(1977) and Mertens & Bourgoigne (1977) reported aggregation pheromones in Collembola.
Leinaas (1983b) found a strong dependence on pheromones and phototaxis in the activity of
two species of Hypogastrura. Leinaas & Fjellberg (1984) found a strict sun orientation in
migrating colonies of an alpine Vertagopus species in Norway.

The social behaviour, resulting in smaller or larger aggregates, is partly interpreted as an
adaptation to patchy, ephemeral habitats (Leinaas, 1983b), or - to put it in other words - to the
non-random distribution of food, optimal moisture conditions, breeding sites, overwintering
sites, etc. Obviously, pheromones help to keep the colonies together in favourable spots, as well
as to coordinate their movements when they have to leave the area.

Curious enough, the mass occurrence on snow often results from the disintegration of
colonies living on ground in the snow-free period. Activity on the snow surface probably acts as
a way of dispersal in species that feed and breed in island-like habitats such as tree trunks,
compost heaps, etc. A number of these “snow-fleas” also have a special winter morph with
modification that possibly makes movement on the snow easier (Leinaas, 1983a).

Dependence on moisture

Moisture is probably one of the most important factors influencing the daily life of the
Collembola. Unlike many other arthropods, Collembola have no hard exoskeleton that prevents
evaporation from the body surface. Much research has been done on the water balance of
Collembola, stressing the importance of keeping down the loss of water vapour from the body
surface. Two different strategies seem to be followed. (1). Either the Collembola live in a
constantly damp environment like deep soil or in caves, which makes special adaptations to dry
air unnecessary. Some species of this group have completely lost their ability to control water
loss (Vannier, 1977). (2). Some species have a variety of morphological adaptations to reduce
the transpiration from the body surface, like scales and dense cover of hairs. This is combined
in many species with an ability to actually survive shorter or longer periods with reduced water
content of the body. In dry periods the species Xenylla maritima and Vertagopus westerlundi
become visibly “shrunken”, but are still active (Leinaas & Fjellberg, 1984). An extreme
example is the inactive but reversible anhydrobiotic stage reported from several species in the
Mediterranean region (Poinsot, 1968, 1974). This anhydrobiosis is also combined with the
ability to survive extreme cold, as much as -180° below zero (Poinsot-Balaguer & Barra, 1983).
But so far no arctic species has been demonstrated to switch to this mode of surviving the
winter.

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Some species have behavioural adaptations, regulating their activity to times with high
substrate moisture (Leinaas & Fjellberg, 1984). Certain xerophilic species react the other way,
seeking drier places when substrate becomes too damp (Vegter, 1983; Bauer, 1979).

Effects of pollution and human activities

Increased attention is paid to the ability of Collembola to live in a polluted environment.
Detoxification mechanisms are found in several soil dwelling groups, like molluscs and
earthworms. In Collembola certain poisonous metals are accumulated in midgut cells and leave
the organism during moulting when the midgut cells are also shed (Humbert, 1974, 1977;
Joosse & Bucker, 1979). Thus, the Collembola may cope with fairly high levels of metals
without any immediate lethal effect. However, Joosse & Verhoef (1983) found reduced
metabolic rate and lowered reproduction in Collembola which were experimentally fed on
lead-contaminated food. Also Bengtsson et al. (1983) found reduced growth rate in the
euedaphic Onychiurus armatus fed on Pb + Cu contaminated fungi, giving an equlibrium
concentration of about 200 ppm copper in live specimens. Similar concentrations were found in
a natural population of O. armatus in the vicinity of a brass mill. Copper is essential in
respiration enzymes, and moderate levels actually increased growth rate of O. armatus.

A similar fertilizer effect was reported by Hagvar & Kjondal (1981) who found an increase
in density of some species in litter bags treated by simulated acid precipitation. The authors
indicate a better reproductive success in “acidophile” species in litter bags with lowered pH as a
possible reason for this increase.

Chemical pollution, directly affecting the physiological processes of the individual, is
certainly different from the more technical disturbance of the habitat caused by activities like
logging, grazing and plowing. Certain opportunistic species clearly benefit from man’s
activities. Greenslade & Greenslade (1983) found an increase of /--selected opportunistic
isotomids in disturbed soils in the Solomon Islands. Unfortunately, such a faunal shift has a
negative effect on the more special ^-selected species. And these species are often stenotopic,
rare and need special protection. Work in progress indicates a serious impoverishment of the
endemic Collembola fauna of Hawaii (Bellinger, pers. comm.). If it is correct that arctic
Collembola in general are opportunistic, as Greenslade (1983) puts it, it shall be interesting to
see if they will be better off in man-made and altered habitats than their tropical relatives.

Collembolan species and assemblages have become subjects for testing ecological theories
that were originally developed on vertebrates and better known groups of arthropods. I have
already mentioned MacArthur & Wilson’s (1967) continuum of r- and /^-selection, which has
now got a third dimension, the adversity or ^-selection of Greenslade (1983) who partly used
Collembola to develop the hypothesis. In future, Collembola will certainly receive increased
attention from students of fundamental ecological and evolutionary processes. Such studies will
probably detect and illuminate a number of taxonomic problems. And this brings me to the
second part of this presentation, the systematics.

SYSTEMATICS

Introduction

The majority of collembolan taxonomists have worked in European countries. The various
European schools have developed new analytical approaches to understanding structure and
classification of Collembola. Consequently, the European fauna is fairly well known.

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563

In North America, Collembola has been a much neglected group, probably because so few
of them are pests in agriculture and forestry. The precise identification of North American
species has been difficult due to lack of required literature. The recent, monumental work of
Christiansen & Bellinger (1980, 1981) has altered this situation. However, their work is a
preliminary and partly synoptic presentation of the fauna, serving as a platform for the future
monographic work which is necessary. A few small soil samples from almost any part of North
America will produce one or more undescribed species. Thus the North American fauna is a
great challenge to taxonomists and a tremendous source of primary material for studies in
classification, evolution and phylogeny. In the following paragraphs I describe some of the
biological phenomena underlying the frequently observed intraspecific variation in morphology.

Ecomorphosis

The Collembola have direct development with continued growth throughout life. The
various instars are separated by ecdysis in which the old cuticle is shed. Apart from the small
changes related to increased size, development of hair cover and reproductive organs, more
drastic changes may appear as a response to both intrinsic and external stimuli.

In a number of papers Cassagnau (1955, 1956, 1971, 1974) described ecomorphosis both in
Hypogastruridae and Isotomidae. Ecomorphic specimens had enlarged setae and cuticular
ridges and spines on the last abdominal segments. Also mouth parts, digestive tract and fat
reservoirs of the body were affected. Ecomorphosis was supposed to be a physiological response
to warm and dry conditions in species having their optimum in damp, cool, habitats. However,
recent studies by Najt (1982) give a more complex interpretation. Given enough time, the
classical species Isotoma tigrina ( olivacea auct.) would produce ecomorphic specimens even at
5° C. Sooner or later all individuals would pass through one or more ecomorphic instars.
Contrary to earlier reports, Najt also found that some of the ecomorphic specimens had normal
digestive tracts and would feed. Apparently the onset of ecomorphosis is genetically fixed in the
species and not a simple response to unfavourable microclimate.

Najt’s observations open some very interesting perspectives. A number of isotomid genera
always have abdominal spines and cuticular modifications which are essentially the same as in
ecomorphic species displaying these structures in certain instars only. It seems possible that
there is an evolutionary sequence from normal, “non-ecomorphic” forms to forms displaying
ecomorphic traits in certain instars only (induced or not induced by external factors), to forms
in which ecomorphic traits have become permanently fixed in all instars (all the “spined”
isotomid genera, Anurophorus, Proctostephanus, etc.). Modifications of the digestive tract are
no longer associated with this last stage, but individuals still benefit from the reduced
evapotranspiration (modified cuticle) and may occupy rather xeric habitats.

Epitoky

Another phenomenon giving similar anatomical modifications as ecomorphosis, is epitoky.
Epitoky is linked to the reproductive cycle and is only shown by reproductive specimens. It is
quite common in many genera and affects various parts of the body, generally with reduction or
modification of anal spines, claws, dens and mucro, body setae and skin granules (Bourgeois,
1971, 1973, 1974, 1981). Males of some Vertagopus get enlarged antennae (Fjellberg 1982).

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Cyclomorphosis

A third phenomenon affecting the individual during its life cycle, is cyclomorphosis - the
occurrence of seasonal forms not linked to the reproductive cycle or ecomorphosis (Fjellberg,
1976). Usually this involves the appearence of distinct summer and winter forms. Sometimes a
switch from the one form to the other is associated with a shift in habitat. Certain
Hypogastrura species have summer forms living on tree trunks while the winter forms are
active on snow. The morphological changes are supposed to facilitate movement on the snow
surface, and thus aid in the dispersal of the species (Leinaas, 1983a). Of particular interest are
the clavate tibiotarsal hairs that appear in the European Isotoma nivea Schaffer during winter.
Following traditional systematics, the winter form would be classified as a Vertagopus
(Fjellberg, 1978).

Studies of the above phenomena are in an early stage. We know very little about the
evolutionary significance and what it means to the individual. A number of examples are still to
be detected and described. Among North American Hypogastrura of the nivicola-packardi—
notha groups, there are a number of forms displaying cyclomorphosis. Some of the currently
recognized species are probably just seasonal forms of each other.

Chaetotaxy

In former days Collembola systematics was based on easily visible characters like number of
eyes, presence/absence of certain organs, fusion of body segments, claw structures, position and
shape of sensorial organs, etc. Such characters still rank among the most important, but must
be used with caution as several of them are affected by the phenomena described above.

During the last two decades, several new analytical methods have come into use. One of the
most promising is probably the strict use of chaetotaxy, the mapping of the body hairs. In the
more primitive families of Collembola the hair cover is fairly simple and each seta has a more
or less fixed position. In more advanced groups, hair cover is dense and irregular, and
chaetotaxy is applied to certain parts of the body like furca, mouth region and antennae, or to
“markers” like body macrochaetae or sensillae.

Chaetotaxy often provides clearcut differences among related species. Apparently evolution
of the hair cover is a fairly rapid process with a high degree of parallelism even in species
groups within a single genus (Fjellberg, 1984c). A general trend seems to be reduction in
number and dislocation and differentiation of setae (Bourgeois & Cassagnau, 1972).
Chaetotaxy as a method has developed in a rather individualistic way. Different specialists have
produced more or less independent systems (Gama, 1969; Rusek, 1971; Cassagnau, 1980). An
important aim for future research would be to develop a common system, making possible a
comparison between distant taxa.

Cytogenetics and physiology

During the last few years, biochemical methods, cytogenetics and physiology have come into
use to discriminate between species. Hale & Rowland (1977) used protein electrophoresis and
amino acid chromatography and found convincing differences among forms of the problematic
Onychiurus armatus group. Hart & Allmong (1979) made an electrophoretic analysis of
esterase enzymes in species from various genera and found consistent differences. These
biochemical methods are promising and may also trace populational characteristics which are
not detected by traditional studies. Dalens (1982) found differences in amino acid composition
in two populations of Hypogastrura tullbergi (Schaffer). Such differences were also

Collembola biology and systematics

565

documented by studies of polytene chromosomes in Bilobella aurantiaca Caroli by Cassagnau
(1976), Dalens (1976, 1977, 1978, 1979) and Dallai (1979). The giant chromosomes display a
varying degree of polymorphism, and offer a great potential for mapping populations of
different origin (Cassagnau et al. , 1979; Deharveng, 1982a). Even the existence of sibling
species is indicated by some authors (Dallai et al ., 1983). Although chromosomal studies are
limited to a small section of the family Neanuridae, they provide valuable information to clarify
species concepts and to understand population genetics of Collembola in general.

Future approach

Sitting by the microscope to observe and make drawings is still the same approach that was
used by the pioneers in Collembola systematics a hundred years ago. Although we have better
quality microscopes and benefit from all the accumulated experience, the species identification
of a collembolan is technically much the same as in “good old days”. When the scanning
electron microscope (SEM) was introduced some years ago, many people expected a revolution
in the morphological studies of small organisms - of course with certain consequences to
established classifications. I think it is correct to say that this has not happened in Collembola.
Certainly the SEM has revealed some interesting developmental lines in surface structure of
the cuticle as well as the fine structure and possible function of some sense organs. But, in
routine identification and in ordinary work, the light microscope is superior.

Progress in Collembola systematics will probably not come as a result of new technical
inventions - although much might happen in the fields of biochemistry and cytogenetics. There
are still some parts of the collembolan body which have not - or only rarely - been used in
practical identification and systematic work. The mouth region is one of the most promising.
Apart from the often very complex maxillae and the simple mandibles lying inside the head,
there are the external maxillary lobes, the labium, labrum and associated structures. The
internal mouthparts, especially the maxilla, are commonly used in the families Neanuridae and
Isotomidae, partly also Hypogastruridae (Massoud, 1967; Poinsot, 1965; Fjellberg, 1977,
1984a). The labium has proved very significant in Entomobryidae and other families (Gisin,
1965; Christiansen & Bellinger, 1980; Deharveng, 1981). In a recent paper I drew attention to
the maxillary outer lobe which is particularly useful in species separation in the bulky genus
Isotoma (Fjellberg, 1984b).

As I have argued earlier I think the differentiation of the mouthparts reflects a progressive
food specialization, and I believe that combined studies of structure, function and actual type of
food ingested might produce some very interesting results.

In order to have success today, a taxonomist has to construct a phylogenetic tree - a
cladogram. The necessary tools and methods are only partly developed in Collembola.
Chaetotaxy has been used by a number of workers (Gama, 1969, 1980; Najt, 1974; Deharveng,
1982b) as well as distribution of body tubercles in Neanuridae (Cassagnau, 1983; Deharveng,
1982c). Reduction seems to be a universal principle in Collembola. Number of hairs are
reduced, ocelli are lost, furca becomes shorter and finally disappears. However, several
structures increase in complexity, like the feeding apparatus, the claws, the differentiation and
shape of individual body hairs, and so on.

There is still a lot of work to be done before the relative plesimorphy/apomorphy of the
various character states along a transformation series can be established. In order to achieve
reasonably sound conclusions, it is necessary to include as many species or samples as possible
from the entire geographical area covered by the taxon under study. In this context, the rich

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Nearctic fauna is particularly important as a pool of still unknown or inadequately described
species.

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PTERYGOTE INSECTS AND THE SOIL: THEIR DIVERSITY, THEIR EFFECTS ON
SOILS AND THE PROBLEM OF SPECIES IDENTIFICATION

P.J.N. Greenslade1
CSIRO, Division of Soils
Glen Osmond
South Australia, 5064
AUSTRALIA

Quaestiones Entomologicae
21:571-585 1985

ABSTRACT

Attention is focussed on soil-dwelling pterygote insects that directly influence soil profiles
and soil fabrics, especially on the largest order of insects the Coleoptera or beetles, and is then
further restricted to groups that are important in the tropics and in the southern hemisphere.
This means especially termites (Isoptera) and ants (Hymenoptera, Formicidae). These two
groups of insects affect soil structure by building mounds and excavating nest chambers and
galleries, often raising lower horizon material to the surface. They also influence the nature of
organic matter and its distribution in the soil profile and, consequently, the horizontal
dispersion of plant nutrients. Some of these influences are illustrated by reference to recent
work on the micromorphological and profile-forming effects of termites. Recent Australian
studies are also used to illustrate the pedological effects of ants. In turn, Australian ants
introduce, and are used to exemplify, the general taxonomic problem presented by many
groups of soil-associated pterygote insects in many parts of the world: numerous species, most
of which are underscribed. The magnitude of this problem is examined on a world basis and it
is suggested that the conventional taxonomic process should be inverted: work should proceed
downward, starting from the higher taxa, organizing the species of larger genera in natural
species groups. Species-level taxonomic studies can be deferred, carried out on a provisional
basis or concentrated on critical groups of species.

L’auteur passe en revue les groupes d’insectes pterygotes qui habitent le sol et autres habitats semblables. II se
concentre sur ceux qui influencent directement les horizons et la structure des sols, plus particulierement sur les
Coleopttres, qui forment le plus grand ordre d’insectes; par la suite il restreint encore davantage son analyses aux
groupes importants des tropiques et de I'hemisph&re sud, c’est-d-dire plus specialement les termites (Isopt&res) et les
fourmis (Hymenopteres, Formicidae). Ces deux groupes d’insectes affectent la structure du sol en construisant des
monticules et en creusant des chambres de reproduction et des galleries, amenant ainsi d la surface des matures
provenant des horizons inferieurs. Ils influencent aussi la nature de la mature organique et sa repartition dans le profit
du sol et, par consequent, la dispersion horizontale des Elements nutritifs des plantes. Quelques-uns de ces effets sont
| illustres a I’aide d’exemples de travaux recents sur le mode d’influence des termites sur la micromorphologie et la
formation du profil des sols. De recentes etudes australiennes servent aussi d’exemples pour illustrer les effets des
fourmis sur les sols. De meme, I’exemple des fourmis australiennes sert d illustrer le probl&me general d' ordre
taxonomique que presentent plusieurs groupes d’insectes pterygotes associes aux sols dans plusieurs regions du monde,
c’est-d-dire la presence d’un trts grand nombre d'especes dont la grande majorite n’est pas decrite. L’auteur analyse
I’ampleur de ce probleme h I’echelle mondiale et conclut que le processus conventional de la taxonomie devrait etre
inverse: c’est-d-dire que les travaux devraient progresser du general au particulier, en etudiant d'abord les taxons
superieurs et en divisant les gros genres en groupements naturels d’espices. Quant aux traitements taxonomiques des
I especes, ils peuvent etre soil retardes, soit executes de faqon provisoire. ou encore concentres sur des groupes d’espices

RESUME

Present Address: CSIRO, Division of Soils, GPO Box 639. Canberra. A.C.T. 2601

572

Greenslade

critiques.

INTRODUCTION

The subclass Pterygota consists of insects which, as adults, are winged or are secondarily
apterous. The subclass contains about three-quarters of all the described species of animals and
a very substantial proportion of them are associated with the soil system. This refers to the
actual mineral and organic horizons of the soil profile and also to related decomposition
habitats such as decaying wood and fallen fruits, carrion and dung. Conventionally regarded as
soil animals as well are those that are active on the soil surface, among insects for example,
many carabid beetles and ants. Indeed it is at the air-soil interface i.e., the soil surface and
uppermost part of the soil profile, that biological activity and diversity usually reach their
absolute maxima in terrestrial ecosystems. Much of this diversity, both taxonomic diversity and
diversity of form and function, is due to pterygote insects. This account of pterygote-soil
relationships is inevitably cursory, even with several restrictions of the area which I attempt to
cover. In accordance with the subject of the Symposium, attention is concentrated on those
pterygote groups that influence soil fabrics and the constitution of soil profiles. Until recent
years, by far the greater part of research in soil zoology had been carried out in temperate
regions in the northern hemisphere. Here, therefore, as a gesture towards restoring some
balance, and to complement other contributions to this volume, I emphasize two groups,
termites and ants, that are particularily important in soils throughout the tropics and over
much of the southern hemisphere. I also select out the Coleoptera or beetles, by far the largest
order of insects and whose species interact with the soil in a multitude of different ways.

THE VARIETY OF SOIL PTERYGOTA

Of the 26 orders that form the Pterygota all but seven contain at least some species in which
an active life history stage involves or depends directly on the soil system. The Phasmatodea
(stick insects) live and feed on vegetation (although the eggs are deposited on or laid in the
soil), while Phthiraptera (lice), Siphonaptera (fleas) and Strepsiptera are parasitic. Three other
orders have winged, generally short-lived adults and exclusively aquatic larvae. Similarly the
Odonata (Dragonflies) and Trichoptera (caddisflies) are essentially aquatic but in both groups
are a few species with truly terrestrial larvae living, for example, in rainforest litter far from
standing water. Increasing terrestrialism is seen in the Mecoptera (scorpion flies); the larvae
are mainly aquatic or are found in damp swampy habitats but some are predators and
scavengers on the open soil surface and in litter.

The other 16 orders are of varying importance in the soil system. Zorapterans,
grylloblattodeans, and embiopterans are relatively small and obscure orders and are, as a rule,
of minor functional importance. Zorapterans are small (length < 3mm ) gregarious insects
living in logs in the tropics. Grylloblattodeans are small and are found under stones and logs,
and in the soil in cold wet situations in the northern hemisphere. The mainly tropical
embiopterans construct silken galleries, sometimes in leaf litter, under stones or in crevices in
the soil. Mantodea (preying mantises) are primarily predators on vegetation but there are some
representatives adapted to life on the soil surface.

Members of all the remaining dozen orders contribute significantly to soil processes. The
Isoptera (termites) are the most closely associated with the soil and they, with the ants
(Hymenoptera, Formicidae, are dealt with separately below. The activities and influences of

Pterygote insects and the soil

573

the rest of the Pterygota are far too varied for it to be possible here to do more than note a few
salient points for most orders.

Blattodeans (cockroaches) and dermapterans (earwigs) are omnivorous feeders and
detritivors. Ground-living psocopterans (booklice) and thysanopterans (thrips) feed on a wide
range of unicellular algae, lichens and fungal hyphae and spores amongst litter and on the soil
surface. Orthoptera (grasshoppers and crickets) include many cryptic forms that shelter in
burrows in soil and decaying wood or under debris, and a smaller number of wholly
subterranean species that are highly adapted morphologically to life in the soil (Key, 1970).
Among the Neuroptera (lace wings) there are families whose larvae are terrestrial predators,
notably the ant-lions (Myremeleontidae) some of which construct pitfall traps in loose sandy
soils. The importance of Lepidoptera (butterflies and moths) as soil animals is probably
underestimated. In Australia the larvae of at least six families, especially Tortricidae, feed on
dead leaves in the litter layer, (Common, 1970), very often when the dry condition of these
leaves inhibits attack or decomposition by other organisms, while some cossid larvae feed,
internally or externally, on the roots of trees and shrubs. Ground-living Hemiptera (bugs)
include seed-feeders, a wide variety of predators and fungus feeders. Some aphids and,
probably, most cydnidas are root-feeders. The Diptera (flies) form one of the larger orders of
insects and very many of their larvae occur in moist, nutrient-rich habitats such as decaying
fruits, dung, rotting vegetation, carrion and logs which have a large active microbial popularion
at a moderately advanced stage of decomposition. Some of these larvae are predacous and
many others have more or less specialised relationships with fungi.

A more detailed examination of the remaining order, the Coleoptera (beetles), which
contains about 40% of all known insect species and perhaps a third, or even more, of all animal
species, illustrates the taxonomic complexity of pterygotes associated with the soil. In Table 1
the families of Coleoptera are arranged in four groups. Group 1 consists of families composed
of species that can be regarded as effectively independent of the soil both as adults and larvae.
It covers parasites, aquatic forms, species living entirely on vegetation or in the nests of
vertebrates and social insects. Group 2 includes all families that have representatives
functioning in the soil system, including species whose activities are centred on discrete habitat
units such as logs and vertebrate dung and carrion. Group 3 excludes the latter and is restricted
to families with species living in soil or litter or on the soil surface. Group 4 is further restricted
to families containing species that penetrate the soil profile and are therefore likely to affect soil
properties directly. Table 1 is derived from a summary of the classification of Coleoptera given
by Britton (1970) modified according to subsequent major changes (Britton, 1974). This
classification is essentially that of Crowson (1955, 1967) and it was used also by Richards and
Davies (1960) who provide brief notes on the biology of the families. Several points should be
noted. The Table refers to species of Coleoptera known in 1970. By 1974 the total had risen
from about 280,000 species to 290,000 (Britton, 1974). If this represents a steady rate of
increse, the total should now stand at around 320,000 known species. However this must still
fall very far short of the real total of all species of Coleoptera (see below). Inevitably the
attribution to categories 1-4 in the Table is arbitrary and debatable for many taxa. Families
are placed according to whether they contain representatives in the habitat groups 1-4. The
Carabidae for example are in Group 4 because the family contains species with burrowing
adults while the larvae of many species with surface-active adults live in the upper part of the
soil profile. This does not mean however that all Carabidae fall in Group 4 and indeed there are
many highly adapted arboreal carabid species, especially in tropical rain-forests. It does mean

I Quaest. Ent., 1985, 21 (4)

574

Greenslade

TABLE 1. Degree of association of families of Coleoptera with soil and allied habitats. The
Table shows the number of known species in families which contain representatives variously
associated with the soil. Only the major families (> 2,000 known species) are shown
individually; the fractions represent number of families/number of known species (see text).

Superfamily

Habitat

Totals

Family

Unrelated to
soil (or
uncertain)

Soil System

Ground layer

Soil profile

Cupedoidea

2/26

Sphaerioidea

4/22

Caraboidea

Carabidae

1/25,000

8/30,184

Other families

6/5,059

1/125

Hydrophiloidea

Hydrophilidae

1/2,000

5/2,400

Other families

4/400

Histeroidea

Histeridae

1/2,500

3/2,507

Other families

2/7

Staphylinoidea

Staphylinidae

1/27,000

1 /27,000

10/35,149

Pselaphidae

1/5,000

1 /5,000

Other families

3/49

5/3,100

4/2,800

Scarabaeoidea

Scarabaeidae

1/17,000

6/18,827

Other families

5/1,287

3/1,587

1/300

Eucinetoidea

1/360

2/85

1/61

3/445

Dascilloidea

1/50

2/69

1/65

3/119

Byrrhoidea

2/300

1/270

2/300

Dryopoidea

7/1,008

Buprestoidea

Buprestidae

1/11,500

Artemetopoidea

1/45

2/115

1/1

3/160

Elateroidea

Elateridae

1 /7,000

1/7,000

5/8,208

Other families

4/1,208

1/3

Cantharoidea

Lycidae

1/3,000

7/8,252

Cantharidae

1/3,500

Other families

1/3

4/1,749

Dermestoidea

3/88

1/731

4/819

(continued on next page)

Pterygote insects and the soil

575

Table 1 (continued)

Superfamily

Family

Habitat

Totals

Unrelated to
soil (or
uncertain)

Soil System

Ground layer

Soil profile

Bostrychoidea

1/700

3/1,604

1/70

4/2,304

Cleroidea

7/9,152

Cleridae

1/3,400

Melyridae

1/4,000

1 /4,000

1/4,000

Other families

1/3

4/1,749

Lymexyloidea

1/37

Cucujoidea

46/41,011

Nitidulidae

1/2,200

Coccinellidae

1/5,000

- '

Meloidae

1 /2,000

Tenebrionidae

1/16,100

Other families

16/2,702

26/13,009

8/8,087

Chrysomeloidea

3/41,200

Cerambycidae

1 /20,000

1/20,000

Chrysomelidae

1/20,000

1 /20,000

1/20,000

Other families

1/1,200

Curculionoidea

8/61,264

Anthribidae

1 /2,400

Curculionidae

1 /60,000

1/60,000

1 /60,000

Other families

4/514

2/1,350

1/1,060

Totals

56/19,200

86/258,621

43/225,857

12/137,495

142/282,880

that identification of a carabid species from soil entails its discrimination within a family of
more than 20,000 known species.

The functional complexity of soil-associated Coleoptera has to be considered as well. The
very high proportion of families in Group 2 in Table 1, and the large number of known species
in these families (over a quarter of a million) reflects, in part, the close association of
Coleoptera with dead wood and fungi. For example, many Coleoptera have mycangia,
structures that allow adult beetles to transport fungal spores when they move from one site to
another. The superfamily Cucujoidea is particularly well represented in Group 2. It is
conservatively divided into 45 families but well over 50 can be recognised, ranging from the
Tenebrionidae, a major family of important detritivores, especially in the tropics and

Quaest. Ent., 1985, 21 (4)

576

Greenslade

TABLE 2. Summary of the superfamily Scarabaeoidea (Coleoptera): larval habitats and food

Family

Subfamily

Known

species

(1970)

Larval

habitat

Food

Lucanidae

750

Dead trees, logs, stumps

Decaying wood

Passalidae

490

Under bark of dead trees, inDecaying wood
logs

Geotrupidae

300

Soil, often in excavations
below dung

Dung, fungi, green and
decaying vegetation

Acanthoceridae

120

? Under bark, in litter, soil

? Decaying vegetation

Trogidae

Scarabaeidae

167

Soil below dry vertebrate
carcasses

Carrion

Aclopinae

Hybosorinae

100

Ground layer

Carrion

Aphodiinae1

1,220

Dung, or burrows in soil

Dung, green vegetation at
night, ? roots

Scarabaeinae

2,000

Soil below dung

Dung

Melolonthinae

9,000

Soil

Roots, organic matter

Rutelinae

2,500

Soil

Roots, organic matter

Dynastinae

1,400

Soil, logs

Roots, organic matter

Valginae

200

? Associated with termites

Cetoniinae

2,600

Soil, humus

Organic matter

1 Including Aegialinae

subtropics, to numerous smaller families of small beetles, many of which probably have very
specialised relationships with micro-organisms.

Even if attention is restricted to Group 4 in Table 1 there still remains the majority of the
larger families of Coleoptera, i.e., six families of 7,000 to 60,000 known species. Apart from
their mechanical effects on soils they are important as predators of other soil animals, or as
feeders on roots, dead plant material and/or associated microbial biomass. The Carabidae,
which have been mentioned, are predominantly predators and scavengers while the larvae of
many if not most Elateridae are root-feeders. Although Cerambycidae, Chrysomelidae and
Curculionidae are all typically phytophagous above the soil surface, the Cerambycidae and
Curculionidae (the largest family of animals) contain root-feeding species. In the
Chrysomelidae, larvae of Eumolpinae and Cryptocephalinae are found in the soil and, they
probably feed on roots as well.

Finally the Scarabaeidae and other families in the Scarabaeoidea exemplify a single major
phylogenetic radiation which contributes to the soil, and to the soil system as a whole, in a
variety of different ways. The larvae live almost exclusively in soil and allied habitats (Table 2)
and occupy a low position in the trophic system, feeding mainly on live and dead plant material
and associated micro-organisms. Adult scarabaeoids are large bulky beetles, up to 7.5 cm or

Pterygote insects and the soil

577

more in length and the group includes some of the largest of all insects. They can occur at very
high population densities, for example around 400 scarabaeids per square metre in Australian
and New Zealand grasslands. Without entering into the extensive literature on their population
dynamics and pest status, it can be noted that Scarabaeidae in particular can have important
influences on soil properties and processes. Over much of the Old World tropics and subtropics
they also have a very significant role in nutrient cycling by disposing of large quantities of dung
produced by herbivorous mammals (Bornemissza, 1961). This may be consumed by the larvae
on the soil surface or in burrows excavated and stocked by the parent beetles.

The question of the food of larval scarabaeoids typifies a recurrent problem in soil zoology:
to distinguish between what is ingested and what is digested. Scarabaeid larvae for instance
commonly ingest live and dead roots, soil organic matter, mineral particles of the rhizosphere
and other micro-organisms; for any one scarabaeid species it may be difficult to establish on
what elements of this intake larval nutrition actually depends (Greenslade and Greenslade,
1983). This is complicated by the existence of a continuum, in the Scarabaeidae for example,
from Cetoniinae whose larvae feed on organic matter, to Rutelinae and Melolonthinae feeding
on live roots. The digestive physiology of scarabaeoid larvae is an important topic which is
beyond the scope of this paper but, clearly, the group as a whole is one which merits much more
attention from the point of view of their effect on the soil system (see Table 3 below).

PEDOLOGICAL INFLUENCES OF SOIL PTERYGOTA

Hole (1981) discussed 11 different ways in which animals can affect soils citing many
examples, with references, that involve insects. They need not be repeated here in detail but
three groups are briefly examined (Table 3).

Merely by excavating galleries and burrows in soil all three of the taxa in Table 3 contribute
to effects 1, 2, 4 and 6. Humphreys and Mitchell (1983) suggest that mixing by soil animals
may have a significant effect on the rate of development of texture contrast soil profiles; they
point out that, over time, it allows rainfall to affect a greater thickness of the profile than just
the surface. Ants and termites backfill voids (effect 3) when they remodel their nests or when
those structures are taken over and altered by other ant or termite species, and soil-living
scarabaeid larvae generally fill their burrows behind them. Soil erosion (effect 5) is influenced
by removal of plant-cover ( e.g ., by scarabaeoid larvae) and by deposition of loose soil on the
surface, susceptible to movement by wind or water, when subterranean nests are initially
excavated or when they are cleaned (e.g., by ants). Elevated ant and termite mounds with a
cemented surface or matrix, or a protective gravel cover can reduce erosion locally but may
accelerate it elsewhere by modifying surface run-off of rain water.

Because of their population biomass and their food, ranging from dry dead wood to already
well-decomposed organic matter, both termites and scarabaeoids have important influences in
regulating the nature and mass of plant litter, and the course and rate of decomposition and
hence nutrient cycling (effects 7 and 9). Termites can accelerate processes by disposing of
recalcitrant substances with a high content of lignin, or retard them by locking up material in
long-lasting nest structures (Lee and Wood, 1971). Ants have minor effects here although those
with large thatched mounds, for example some wood ants, Formica spp. in the northern
hemisphere, and the myrmiciine Myremecia pilosula in Australia, have some effect on the
distribution of litter. However, Cowan et al. (1985) concluded that the Australian Camponotus
intrepidus which also has thatched mounds, has a trivial role in pedogenesis. Ants very rapidly

I Quaest. Ent., 1985, 21 (4)

578

Greenslade

TABLE 3. Pedological effects of soil fauna, from Hole (1981), and the roles ofthe pterygote
insects: termites, ants and scarabaeoid beetles.

Effect

Taxon

Termites

Ants

Scarabaeoids

Mixing

Forming voids

Backfilling voids

Forming and destroying peds

Regulating soil erosion

Regulating movement of
water and air in soil

Regulating plant litter

( + )

Regulating animal litter

Regulating nutrient cycling

( + )

10.

Regulating biota

11.

Producing special
constituents

recycle any invertebrate carrion that appears on the soil surface (effect 9) usually finding it
within minutes of its arrival, but this cannot compare with the mass effect of termites and
beetles (such as scarabaeoids) on plant material.

Termites seem to have little direct influence on other biota (effect 10), excluding their
microbial gut flora and the animals, mainly insects, that live with them in their nests. Indeed
much of the success of the order Isoptera must derive from their exploitation of resources that
were hardly used by other animals. In contrast, ants are particularly significant as dominant
predators and competitors on the soil surface and in litter, with profound effects on the rest of
the soil and surface fauna at both ecological and evolutionary levels. Soil-living scarabaeoids
perhaps illustrate the classic influences of soil fauna (according to conventional wisdom) in
comminuting plant material, dispersing soil micro-organisms and controlling their populations
by feeding upon them.

All three taxa have the final effect (11) of producing special constituents. Both ants and
termites make structures from selected soil particles, frequently cemented with salivary
secretions or faecal material, while scarabaeoid larvae leave excreta-filled tunnels behind them.

Three effects of soil insects do not come across clearly in Hole’s scheme. They are effects:
(1), on the rhizosphere ( e.g by scarabaeoid larvae); (2), of soil-nesting and mound-building
ants and termites on the distribution pattern of plant nutrients in the horizontal plane; and (3),
on the composition and structure of A-horizons (although to some extent the last is covered by
Hole’s ‘mixing’ or ‘bioturbation’).

The interactions of termites with soils were reviewed by Lee and Wood (1971). They showed
that two of the most important activities of species that build mounds are the concentration, in
the mounds, of organic matter and hence plant nutrients, and the elevation of lower horizon

Pterygote insects and the soil

579

material to the surface. More recently, Holt et al. (1980) and Spain et al. (1983) have studied
the pedological significance of mound-building termites in northern Australia and their results
are typical of those in the literature. On two soil types (red and yellow earths) Holt et al.
(1980) found a total mound basal area of about 1% of their plots. Assuming the life-time of a
mound from inception to complete erosion to be 25-50 years, they calculated an annual rate of
accumulation of lower horizon soil on the surface of 0.025-0.05 mm per year. This means that
any point in the landscape will support a termite mound once every 1-1,000 years and that in
the 10,000 years since the end of the Pleistocene a 20-50 cm thick uppermost horizon could
develop from the erosion of termite mounds. Since termite galleries commonly extend 1-2 m
into the soil it follows also that over a few millennia entire soil profiles, or all of the upper part,
can be worked and reworked by termites. In this way, termites appear to have a major role in
the formation of the tubulo-alveolar laterites and pisolitic laterites and bauxites that are
frequent throughout the warmer parts of the world. From the micromorphology of these
laterites and bauxites, and their content of plant and termite fragments, de Barros Machado
(1982a, b) concluded that they are formed by capillary impregnation by sesquioxides of the
lining of termite galleries.

Mound-building ants also have received considerable attention on account of their possible
role in raising soil to the surface and in affecting the distribution of plant nutrients, recently for
example from Briese (1982), Cowan et al. (1985), Culver and Beattie (1983), Davidson and
Morton (1981), Humphreys and Mitchell (1983) and Mandel and Sorenson (1982) and these
authors provide many references to earlier investigations. Most of this work however has been
done outside the tropics (in which ants reach their greatest diversity) and has generally involved
only one or a few ant species which construct distinct nest mounds.

Humphreys and Mitchell (1983) recognised two broad types of mound, Type I where subsoil
material is simply deposited loosely on the surface and Type II in which the mound is
chambered, and the material compacted and cemented, to form a much more permanent nest
structure. In fact there is a continuous range of nest types from subterranean nests that just
open on to the soil surface, to entrances that are surrounded by fans, rings or small turrets of
loose soil, through mounds that are increasingly compacted, worked and variously covered with
thatch or gravel, to some very elaborate structures. Examples are the nests of New World
fungus-growing ants (Attini), described by Moser (1963) and Weber (1966) (and see Wilson,
1971), and the ring nests of certain Polyrachis species on red earths and earthy sands in central
Australia. The latter, which have yet to be described in detail, consist of substantial earthern
rings which are covered with dead leaves of mulga ( Acacia aneura) and contain a complex
arrangement of interpenetrating galleries and spouts opening into voluminous atria.

These mound structures have a variety of functions ranging from spoil heaps or middens, to
the control of nest microclimate and flood-defence. When they are thatched or covered with
gravel the covering may act as a protection against rain splash erosion (Cowan et al ., 1985)
and/or as a behavioural boundary (Gordon, 1984).

For some species the longevity of these mounds, for example the large gravel-covered nests
of the meat ant, Iridomymex purpureus , of eastern Australia, is such that their contribution to
pedogenesis is negligible, despite their size (Greenslade, 1974; Cowan et al ., 1985). In other
species, however, the turnover rate is much more rapid and Culver and Beattie (1983) cite King
and Sallee’s (1956) and Smallwood’s (1982) observations that the half life of large Formica
mounds may be 10 years or less while some species relocate their nests several times a year. In
arid Australia there are indications that nest turnover, even for the elaborate ring nests of

Quaest. Ent.. 1985,21 (4)

580

Greenslade

Polyrachis species, is very much more rapid than was hitherto assumed (P.J.M. Greenslade and
W.A. Low, E. and B. Case, unpublished observations). It has been estimated that subsoil is
brought to the surface by ants at rates of up to 0.1 mm per year, for example by Formica
cinerea in North America (Baxter and Hole, 1967), quite comparable with estimates obtained
for termites. These rates are of a magnitude that could be a significant influence in pedogenesis
within the time-span of the Holocene so that it becomes unnecessary to extrapolate over longer
periods of time that include major climatic changes and probable changes in the rate and
nature of biological activities in the soil. Humphreys and Mitchell (1983) point out that,
depending on soil material, rate and depth of mixing, and intensity of rainfall and rainwash,
animal activity in general (including that of ants and termites) can either homogenize soil
profiles or accentuate texture contrasts, leading to duplex profiles.

In semi-arid southern Australia Briese (1982) studied the combined effects of the members
of a moderately diverse assemblage of ants. There was a total of 22 species in a plot of 500m2 of
low, open chenopod shrubland and none of them built large mounds. The turnover rate of soil
attributed to these ants was 0.03 mm a year, again comparable to figures for termites in
northern Australia.

Several investigators have compared the properties of mound and nonmound soils [see for
example Culver and Beattie (1983), Davidson and Morton (1981a, b) Mandel and Sorenson
(1982)]. They found commonly, but not invariably, enhanced levels of plant nutrients in the
mound soils, notably of nitrogen and available phosporus and, where ants allow plants to grow
on mounds, floristic contrasts with surrounding areas. Briese (1982) compared soils from the
nests of six selected ant species with those from control sites. Four seed-harvester or
seed-harvester-omnivore species and one predator showed increased concentrations of nitrogen
and phosphorus in nest over control soils, especially close to the surface. This was related to the
presence of discarded prey fragments, seed husks and other plant material. However, a
non-harvesting ant, an Iridomyrmex species which is a predator-omnivore, does not discard
material around the nest entrance, and there was no nest-enhancement of plant nutrient
concentrations. Levels actually decreased, probably because of the presence of lower horizon
material that had been brought to the surface.

Charley (1971) and Rixon (1970) have described and discussed the significance of the
surface patterning of plant nutrients in the type of shrubland in which Briese studied ants.
Briese added the point that, by concentrating nutrients around their nest entrances, ants
contribute to a mineral mosaic which influences the overall nutrition of the plant community.
This can be extended to other soil-nesting ants and termites. The scale at which they are likely
to influence nutrient patterns is close to that illustrated by Tillman (1982), who argued that
adaptation to and competition for specific ratios of resources, such as nutrients, is a major
factor in the coexistence of plant species and in the control of floristic diversity. Consequently,
the effects of social insects on soils may have wide significance to vegetation.

Briese’s (1982) work takes us back to the problems created by insect diversity. First, when
ant communities are composed of large numbers of species (as is the rule over most of Australia
for example, and the whole of the world’s tropical regions) with differing effects on the soil, the
influence of each species should be assessed independently. When a local ant fauna can consist
of more than 100 species in an area of less than 1 ha, the difficulties are obvious. Second, there
is the problem of identifying the species, essential if one study is to be comparable with another.

Pterygote insects and the soil

581

Fig. 1. Relative apparent taxonomic knowledge of soil Pterygota, from 1, probably very inadequate for most groups, to 6,
good, more than 90% of species described, at least as adults, in most groups.

IDENTIFICATION OF SOIL INSECTS

The accurate identification of species is essential to any biological study in order to allow the
comparison, application and testing of results, just as the consistent and accurate identification
of soil types is essential to any study of soils. Hollis (1980) has edited a multi-authored guide
aiming to provide a list of primary references, enabling non-specialists to set about identifying
insects, including soil pterygotes, from any part of the world. Because of the diversity of soil
biota, however, and especially of soil insects, specific identification often seems to be an ideal
the attainment of which is surrounded by insuperable barriers. In many studies, specific
identification is sacrificed for the sake of statistical validity, and animals identified only to the
level of the family or even the order, a pronounced deficiency of much research in soil zoology.

Australian ants illustrate the sort of problem that the identification of insect species presents
to the soil zoologist. To take only one example, Bolton (1981) revised the African members of
the ant genus Meranoplus which is distributed through the Old World tropics and adjacent
areas. From the taxonomic literature, he concluded that the Australian region had the most
diverse fauna with ca. 25 named taxa. Since 1970, I have collected more than 200 Meranoplus
species in Australia and this can be but a fraction of the total so that less, and probably much
less, than 10% are described. Consequently, for this quite important genus there are hardly any
descriptions of species, no keys for their identification and its study is closed to the
non-specialist. Admittedly, the Australian ant fauna is remarkably diverse; but in other
continental areas other groups of soil insects have radiated in the same way, creating the same
obstacles to research.

New (1984) refers to this as the ‘taxonomic impediment’ to work on insects. The problem
was discussed by Wilson (1980) who considered it capable of being solved. He started with the
then commonly agreed maximum figure of a total of 10 million species of organisms of which
ca 1.5 million had been described. He suggested that if a taxonomist deals with 10 species per

Quaest. Ent., 1985,21 (4)

582

Greenslade

year over a span of 40 years, 25,000 taxomists’ working lives would be required to revise the
biota of the world, a not impossible number given contemporary populations of scientists.
However, basing calculations on the number of host-specific Coleoptera on tropical trees, Erwin
(1982) proposed that there may be up to 30 million species of tropical arthropods. New (1984)
describes “reactions ranging from incredulity to relief that a more realistic figure has been
published.” It is unlikely that many soil zoologists acquainted with the invertebrate fauna of the
litter layer in lowland tropical rain-forests would dispute Erwin’s estimate, even if they
disagreed with the means by which he arrived at it. Indeed it is quite probable that an extensive
survey of ground-layer invertebrates in tropical rain-forests would result in another massive
increase to the estimated total. It should be added that this figure refers to taxonomist’s
morphological species and evades the question of species that can be recognised only with the
biochemical and karyological techniques of the geneticist.

The taxonomic problem is not uniformly spread throughout the different groups of soil
insects or the world’s geographical regions. Some insects that are important in the soil system
are relatively well known, even in the tropics. The prime example is the Isoptera (termites)
although even here much taxonomic study is still needed. At the other extreme lie groups such
as tropical curculionid and staphylinid beetles. There have been a number of attempts to assess
the state of taxonomic knowledge of selected portions of the biota. Examples are surveys of
recorded, and estimates of the uncollected, soil fauna of Canada (Danks, 1979; Marshall et al.,
1982); terrestrial and freshwater Hexapoda {i.e., pterygote and apterygote insects and allied
groups); Myriapoda and Arachnida of New Zealand (Watt, 1983); insects of Australia
(Taylor, 1976); and biota of the British Isles with particular references to insects (Stubbs,
1982). Figure 1 is a very subjective attempt to illustrate geographical variation in the apparent
magnitude of the taxonomic impediment to work on the pterygote of the soil insects. It derives
from surveys such as those mentioned, superficial familiarity with the taxonomic literature and
the impression gained from collecting and sampling a variety of soil insects in the world’s major
biomes. Regional variation is caused by such factors as differences in the diversity of faunas
and in the history of biological investigation in different areas.

By far, the best known soil insects are those of Britain and northwestern Europe, where
probably more than 95% of species are described and a comprehensive range of guides and keys
to adults is available. Even here, however, the specific identification of immature stages is
generally difficult and impossible for many taxa. In New Zealand, more than half the species
are thought to be described, while in Australia, it is estimated that more than half have yet to
be collected. In the humid tropics, of course, the situation is much worse, but precisely how
much we do not know.

Much current taxanomic work is based on revisions of genera, in which all available
representatives of a genus are gathered together and species are described or redescribed and
catalogued. It is then possible to revise the higher classification and to prepare keys to species.
For most of the world’s soil pterygotes it is obvious that this conventional taxonomic process is
quite inadequate. For example, of the postulated 30 million or so arthropod species about 12
million or 40% should be Coleoptera. At the current rate at which Coleoptera species are being
described ( ca . 2-3,000 per year, see account of Coleoptera here) a very long time indeed would
elapse before all were known. Unsatisfactory partial answers are available in that attention can
be restricted to better known taxa and/or the soil zoologist can become his own taxonomist.
Sometimes species can be identified through a combination of voucher specimens and code
numbers, but this system fails in large, inadequately known genera. A possible solution lies in

Pterygote insects and the soil

583

inverting conventional taxonomy. Instead of starting with the description of species, work
should proceed downward from higher taxonomic categories in order to provide guides to
genera and, within large genera, to natural groups of species. In this way, the material with
which the soil zoologist works is reduced to sets of species of manageable size that are relatively
easily recognised and when recognised convey biological information. It is feasible also to link
species’ identities to vouchers and code numbers. The time-consuming production of detailed
species-descriptions, which generally fail to discriminate between sibling or cryptic species, and
are rarely adequate without access to types, can probably be omitted. At least it can be
deferred, carried out on a provisional basis or concentrated on critical groups of species.

REFERENCES

Baxter, F.P. and F.D. Hole. 1967. Ant ( Formica cinerea ) pedoturbation in a prairie soil. Soil
Sci. Soc. Am. Proc. 31: 425-428.

Bolton, B. 1981. A revision of the ant genera Meranoplus F. Smith, Dicroaspis Emery and
Calyptomyrmex Emery (Hymenoptera: Formicidae) in the Ethiopian Zoogeographical
region. Bull. Brit. Mus. (Nat. Hist.) (Ent.) 42, 2.

Bornemissza, G.E. 1976. The Australian dung beetle project 1965-1975. Australian Meat
Research Committee Review No. 30: 1-30.

Briese, D.T. 1982. The effect of ants on the soil of a semi-arid salt bush habitat. Insectes
Sociaux 29: 375-386.

Britton, E.B. 1970. Coleoptera (beetles), pp. 495-621. In: The Insects of Australia. Melbourne
University Press, Melbourne.

Britton, E.B. 1974. Coleoptera (beetles). In: The Insects of Australia , Supplement. Melbourne
University Press, Melbourne.

Charley, J.L. 1971. The role of shrubs in nutrient cycling. In: Wildland Shrubs, their Biology
and Utilization. Utah State University.

Common, I.F.B. 1970. Lepidoptera (moths and butterflies). In: The Insects of Australia. Pp.

765-866. Melbourne University Press, Melbourne.

Cowan, J.A., G.S. Humphreys, P.B. Mitchell and C.L. Murphy. 1985. An assessment of
pedoturbation by two species of mound-building ants, Camponotus intrepidus (Kirby) and
Iridomyrmex purpureus (F. Smith). Aust. J. Soil Res. 22: 95-107.

Crowson, R.A. 1955. The Natural Classification of the Families of Coleoptera. Nathaniel
Lloyd, London.

Crowson, R.A. 1967. The Natural Classification of the Families of Coleoptera. Reprint with
addenda and corrigenda. W.E. Classey, London.

Culver, D.C. and A.J. Beattie. 1983. Effects of ant mounds on soil chemistry and vegetation
patterns in a Colorado montane meadow. Ecology 64: 485-492.

Danks, H.V. (Editor). 1979. Canada and its insect fauna. Mem. Entomol. Soc. Can. 108.
Davidson, D.W. and S.R. Morton. 1981. Myrmecochory in some plants (F. Chenopodiacae) in
the Australian arid zone. Oecologia 50: 357-366.
de Barros Machado, A. 1982a. The contribution of termites to the formation of laterites. Proc.

II, International Seminar on Laterization Processes, Sao Paulo, July 1982.
de Barros Machado, A. 1982b. Termitic remains in some bauxites. Proc. II. International
Seminar on Laterization Processes, Sao Paulo, July 1982.

Erwin, T.L. 1982. Tropical forests: their richness in coleoptera and other arthropod species.

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Coleopt. Bull. 36: 74-75.

Gordon, D.M. 1984. The harvester ant (Pogonomyrme x badius ) midden: refuse or boundary?
Ecological Entomology 9: 403-412.

Greenslade, P.J.M. 1974. Some relations of the meat ant, Iridomyrmex purpureus
(Hymenoptera: Formicidae) with soil in South Australia, soil Biol. Biochem. 6: 7-14.

Greenslade, P.J.M. and P. Greenslade. 1983. Ecology of soil invertebrates, pp. 645-669. In:
Soils, an Australian Viewpoint. Division of Soils, CSIRO. SCIRO, Melbourne/Academic
Press, London.

Hole, F.D. 1981. Effects of animals on soil. Geoderma 25, 75-112.

Hollis, D. 1980. Animal identification, a reference guide. Vol. 3, Insects. British Museum
(Natural History), London, John Wiley & Sons, Chichester.

Holt, J.A., R.J. Coventry and D.F. Sinclair. 1980. Some aspects of the biology and pedological
significance of mound-building termites in a red and yellow earth landscape near Charters
Towers, north Queensland. Aust. J. Soil. Res. 18: 97-109.

Humphreys, G.S. and P.B. Mitchell. 1983. A preliminary assessment of the role of bioturbation
and rainwash on sandstone hillslopes in the Sydney Basin, pp. 66-79. In: Young, R.W., and
G.C. Nanson (Editors). Aspects of Australian Sandstone Landscapes. Australian & New
Zealand Geomorphology Group Special Publication No. 1.

Key, K.H.L. 1970. Orthoptera (grasshoppers, locusts and circkets), pp. 323-347. In: The
Insects of Australia. Melbourne University Press, Melbourne.

King, R.L. and R.M. Sallee. 1956. On the half-life of nests of Formica obscuripes. Proc. Iowa
Acad. See. 63:721-723.

Lee, K.E. and T.G. Wood. 1971. Termites and Soils. Academic Press, London.

Mandel, R.D. and C.J. Sorenson. 1982. The role of the western harvester ant ( Pogonomyrmex
occidentalis) in soil formation. Soil Sci. Soc. Am. J. 46: 785-788.

Marshall, V.G., D.K. McE. Kevan, J.V. Matthews and A.D. Tomlin. 1982. Status and research
needs of Canadian soil arthropods. Biological Survey of Canada.

Moser, J.C. 1963. Contents and structure of Atta texana nests in summer. Ann. Entomol. Soc.
Amer. 56: 286-291.

New, T.R. 1984. Insect Conservation, an Australian Perspective. Junk, Dordrecht.

Richards, O.W. and R.G. Davies. 1960. A.D. Imms: a General Textbook of Entomology. 9th
Edn., Methuen, London.

Rixon, A.J. 1970. Cycling of nutrients in a grazed Atriplex vesicaria community, pp. 87-95.
In: R. Jones (Editor). The Biology of Atriplex.

Smallwood, J. 1982. Nest relocations in ants. Insectes Sociaux 29, 318-147.

Spain, A.V. Rev. Ecol. Bibl. Sol 20:547-566.

Stubbs, A.E. 1982. Conservation and the future of the field entomologist. Proc. Trans. Brit,
entomol. nat. hist. Soc. 15: 55-67.

Taylor, R.W. 1976. A submission to the enquiry into the impact on the Australian environment
of the current enquiry programme. Aust. Senate Official Hansard Rep. (Ref. Current
Woodchip Enquiry), Melbourne 12 August, 1976, pp. 3724-3731.

Tilman, D. 1982. Resource Competition and Community Structure. Princeton University
Press, Princeton.

Watt, J.C. 1983. Hexapoda, Myriapoda and Arachnida, pp. 62-67. In: Brownsey, P.J., and
A.N. Baker (Editors). The New Zealand Biota. What Do We Know After 200 Years?
National Museum of New Zealand, Miscellaneous Series No. 7.

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Weber, N.A. 1966. Fungus-growing ants. Science 153, 587-609.

Wilson, E.O. 1971. The Insect Societies. Belknap Press of Harvard University Press,
Cambridge, Massachusetts.

Wilson, E.O. 1980. Taxonomic exploration (book review). Science 208: 721-722.

| Quaest. Ent., 1985, 21 (4)

SOIL ANIMALS AND SOIL FABRIC PRODUCTION: FACTS AND PERCEPTIONS

Daniel L. Dindal

Professor of Soil Ecology

State University of New York

College of Environmental Science and Forestry

Syracuse, New York 13210

U. S. A.

Quaestiones Entomologicae
21:587-594 1985

ABSTRACT

Soil is composed of microenvironments resulting largely from distinct biological inputs or
activities. These microenvironments in turn foster development of microcommunities which, in
composite, determine patterns of soil micromorphology. Biological processes contribute to the
local dynamics of microcommunities, bring about changes in soil structure, and are major
features of soil function. Various spheres of influence, based upon soil microcommunities
organize biologically mediated interactions between soil structure and soil function.

RESUME

Le sol se compose de microenvironnements qui resultent principalement d’energies ou d’activites biologiques
distinctes. En retour, ces microenvironnements promouvoient le developpement de microcommunautes qui, dans leur
ensemble, determined les differents aspects de la micromorphologie du sol. Les processus biologiques contribuent h la
dynamique locale des microcommunautes, modifient la structure du sol, et sont parmi les plus importants aspects de la
fonction du sol. Diverses spheres d’influence, qui originent des microcommunautes terricoles, organisent, par I’entremise
d’agents biologiques, les actions reciproques entre la structure et la fonction du sol.

INTRODUCTION

The contribution of soil animals to production and maintenance of soil fabrics is a vital part
of soil function., Being a complex system, soil must be investigated in a holistic way so as to
include the influence of soil animals.

Soil litter is the major source of organic input at the interfaces with the abiotic mineral soil.
Soil litter is composed of a variety of plant parts including various species of leaves, seeds and
fruits, other dead plant tissues - woody and herbaceous, fungal fruiting bodies, rhizomorphs and
other microbial tissues. Animal contributions to litter are of all types and sizes: fecal masses
and pellets, nesting materials and various forms of shed skins, egg shells and carcasses. Litter
may appear as a zone of very high entropy, but when viewed in a more microscopic sense, order
abounds, and within a very short time interbiotic and biotic - abiotic organization reaches high
levels of fidelity. Cybernetic information is stored and patterns become more predictable soon
after litterfall. Each litter component, being of different taxonomic origin or morphological
form, provides a unique mass of highly organized elements, compounds and energy sources, and
is a potential microhabitat. Each type of microhabitat, dependent upon its origin or form,
further supports a predictable microcommunity of associated soil organisms predominantly
decomposer microorganisms and invertebrates - the decomposer food web (Dindal, 1971;

588

Dindal

1980). Decomposer microcommunities are structurally and functionally unique (Dindal, 1971;
1978a, b). When we think of the array of litter microcommunities as intricately related to
abiotic soil components, a microecosystem paradigm emerges. Furthermore, the
micromorphology of the soil fabric is an expression of microcommunity structure and function.
Biological mediation, therefore, is vital at interfaces within the soil, and entropy of the soil
ecosystem is continually lessened by these biotic activities. The
microecosystem/microcommunity concept provides an obvious link connecting the abiotic,
vegetative and vertebrate animal characteristics of any macroecosystem; thus a holistic
approach to the understanding of soil fabric production is imperative.

AUTUMN OF LIFE

Litter Fall

An obvious component of soil litter is the mass of deciduous leaves shed annually within a
temperate forest. In a northern hardwood forest in central New York (dominated by sugar
maple, Acer saccharum Marsh, and ash, Fraxinus spp.) all of the leaf canopy was shed in 167
days (October 3, 1979 to January 17, 1980) with 86% of the leaves falling during the first 24
days. Surface area of fallen sugar maple leaves during this autumn was 3.6m2/m2 of soil surface
(Dindal and Dindal, unpubl.). Also, we found the majority of the leaf litter was incorporated on
site by the following April (Tardiff and Dindal, 1980); thus the potential physical and chemical
properties of the previous leaf litter fall are accessible within soil by the following growing
season. Such a rapid rate of input of organic compounds and fiber via leaf material is of
considerable ecological significance related to soil structure and micromorphology.

Earthworms, Lumbricus terrestris L. (Hamilton, 1983) and isopods, Oniscus asellus L.
were the dominant biotic mediators of incorporation of of leaf litter on the New York site. The
remainder of the maple leaves, not consumed by earthworms, were skeletonized by the isopods,
with the removal of 91% of the leaf (mostly mesophyll and palisade tissues) leaving the 9%
fibrovascular bundle leaf structure intact. The obvious increase in surface area resulting from
skeletonization is phenomenal; isopods reprocess leaf tissue into numerous fecal pellets each
with an average surface area of 3.3 mm2. Each pellet is a rectangular solid averaging 1.1 mm
long with a 0.6 mm square end. In addition to the incorporation of these pellets into the soil,
Oniscus also produces a network of 0.33 m2/m2 of fibrovascular bundle “lace” destined for soil
microsites. This network provides potentially active sites with capacity for cation exchange and
a cellulose/lignin matrix onto which soil minerals may be further physically and chemically
bound.

Carrion Deposition

Carrion, invertebrate exuviae and carcasses of vertebrates comprise important components
of soil litter even though they are less obvious or deliberately ignored. Carrion falling to the soil
surface deposit various elements, biochemical compounds and energy sources that support
decomposer food webs. Heterotrophic microcommunities quickly colonize, use and distribute
the structural ingredients of highly proteinaceous vertebrate carcasses. Nitrogenous and
sulphur laden compounds seep into the adjacent soil. Molecules that are naturally recalcitrant,
like chitin which contains nitrogen radicals, scleroproteins with both nitrogen and sulphur
moeities, calcium and magnesium carbonate-protein complexes and even organosiliceous
compounds, can originate from decay of invertebrate and vertebrate carcasses. They are buried,
directly or indirectly, providing unique slow release compounds as well as extraordinary organic

Soil animals and soil fabric production

589

matter substrates on which and from which soil fabric is produced.

Fecal Rain

Animal defecation constantly subjects the earth’s surface to a “rain” of feces. These
nutrient/energy-rich additions to soil sediments are extremely subtle, even more so than the fall
of carrion. Perhaps the holistic effects of this omnipresent phenomenon would never be totally
appreciated unless all organisms evacuated at exactly the same moment! Because defecation is
a natural packaging, recycling process by which biochemical compounds and energy sources
are pelletized, it is very important to soil formation and to dynamics of decomposer food webs.
Fecal structure, chemical composition, pellet size, and rate of deposition are species specific.
These characteristics represent a wide array of variables that are interjected into substrates of
all ecosystems. Fecal pellets or dung balls are, therefore, energy/nutrient dissemules that are
formed, transported and distributed onto the earth’s soils by all kinds of animals.

SPECIFIC BIOTIC PROCESSES AT ORGANIC/INORGANIC INTERFACES

Total functions of decomposer microcommunities in association with unique microhabitats
are responsible for processes such as soil fabric production, translocation, and transformation
which ultimately lead to fabric reorganization and “soil ripening” as per Bal (1982). These are
part of the biological activities referred to by Kubiena (1948) as the ’’principal driving forces of
any soil forming processes.” Several biologically mediated processes warrant more specific
comment.

Slime and Gum Production

Mucopolysaccharides and other carbohydrate complexes are produced by many soil
decomposer organisms within their soil/litter realm. Slimes and gums are exuded as metabolic
byproducts, lubricants for mobility, forms of chemical and physical defense, modes of substrate
attachment, and mechanisms for food-getting and pheromonal dispersal agents. In addition to
their adaptive roles, these compounds may directly or indirectly cause or aid in formation of soil
aggregates causing organic and abiotic materials to adhere forming erosion-stable units. In
turn, this gives specific character to both the micro- and macro-structure of soil, i.e., increasing
organic matter incorporation, water holding capacity, porosity and ion exchange capacities.
Also, the metabolism of the soil ecosystem is enhanced by the subtle monomolecular layers of
slime that are potentially important microsubstrates for soil microbial colonization and
population maintenance.

Coprophagy and Geophagy

Eating soil or mineral materials - geophagy (Jones and Hanson, 1985; Kramar, 1973), and
consuming another individual’s feces (either interspecifically or intraspecifically) - coprophagy
(Hassall and Rushton, 1985; Simmons, 1983; Anderson, 1978; Kenagy and Hoyt, 1980), is not
uncommon in the natural world. With future research, many more examples involving soil
animals are likely to be documented. In the observed examples of coprophagy, a diversity of
organic compounds, already subjected to an initial digestion are further subjected successively
to the digestive processes and gut symbionts of new consumers. Inorganic and organic
substances are forced together very closely within a gastrointestinal microhabitat and
eventually incorporated into the soil matrix. Large fecal masses or pellets are altered

Quaest. Ent., 1985,21 (4)

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Dindal

chemically; they are reduced to smaller and smaller units, increasing in surface area and thus
having particular impacts on the soil micromorphological structure and function.

Insertion of Organic Matter

Both invertebrates and vertebrates exhibit habits that cause many forms of organic matter
to be inserted into soil thus modifying the soil fabric. Mammal burrows filled with organic
material within soil profiles were recognized as Krotovinas by early agronomists. Birds which
nest in ground burrows, such as the burrowing owl ( Speotyto cunicularia ), bank and cliff
swallows ( Riparia r. riparia and Petrochelidon pyrrhonota ) and the belted kingfisher
(Megaceryle a. alycon) all deposit and interject various organic compounds during production
of their annual broods. Along sea coasts and above intertidal lines, crabs regularly bury
carcasses and other organic debris. Dung beetles (Stevenson, 1983; Brussard, 1985), some
spiders like Geolycosa (Shelford, 1913), ants, and enchytraeid and lumbricid worms constantly
bury or intertwine organic matter with soil particles. Dipteran maggots migrate from their
decayed food source, burrow and then pupate within the surrounding soil; most edaphic pupae
die and decay in this buried state (Dillon, 1984; Hall, 1947).

Although each interposition of organic substance may be relatively microscopic when viewed
from the macroecosystem level, the constancy of pattern, the regularity and ultimate sum of
biotic input via these subtle and mundane processes greatly influences soil micromorphology
and structure. Such active processes led Jenny (1980) to classify soil invertebrates functionally
as “mechanical blenders” of soil. The insertion-upwelling activities are perhaps analogous to the
action of the sewing machine where organic compounds are threaded into soil fabric following a
specific spatial and temporal pattern; each stitch, no matter how minor, has its functional and
structural role.

Upwelling of Inorganic Matter

Certain soil animals are responsible for mining and deposition of large quantities of mineral
materials on soil surfaces. Burrowing rodents unearth and build surface mounds that have the
heterogenous physical characteristics and textures of deeper soils. Upwelling not only
influences the below-ground soil fabric but also noticeably shapes the surface landscape,
whether caused by mound-building ants (Werner, 1984) or by fossorial rodents (Cox, 1984).
Less noticeable, but of equal importance, are the excavations of mineral soil by non-mound
building ants and earthworms. In a central New York old field, 78% of the mineral soil
particles excavated by the ant, Lasuis niger neoniger Emery, are within the 1 80-425 /j,m size
range (Dindal, pers. obs.). Possible species specificity of size selection of soil particles and
movement by ants is probably an important factor in soil formation. We observed species
specificity of soil aggregate (fecal casting) size formation relative to several dominant
earthworm species (Dindal, Theoret and Moreau, 1978); also Lumbricus terrestris populations
are highly correlated with presence of 4.0 mm water-stable soil aggregates (Hamilton, 1983).
From these studies specific size relationships of soil aggregates to their biotic source are
suggested (Table 1).

Gradual Comminution

Constituents of ingested plants and animals are radically transformed into complex forms
and new compounds as they pass through the guts of large and small grazers and carnivores.
These materials are microbially primed and again low entropy is facilitated (this time by

Soil animals and soil fabric production

591

Table 1: PROPOSED SIZE RELATIONSHIPS OF EROSION-STABLE SOIL

AGGREGATES TO EARTHWORM SPECIES AND THEIR SYMBIONTS.

SOIL AGGREGATE
SIZE (mm)

SOIL BIOTIC SOURCE OF FORMATION

> 6.4

Symbiotic complex of earthworms, roots and microorganisms

4.0

Lumbricus terrestris L.

2.0

Aporrectodea tuberculata (Eisen)

1.0

Octolasion tyrtaeum (Savigny)

0.5

Lumbricus rubellus Hoffmeister

0.5

Dendrobaena octaedra (Savigny)

0.25

Dendrodrilus rubidus (Savigny)

0.15

Microorganisms

symbiotic relationships) as the food bolus is gradually transposed into feces.

Expelled remnants of ingesta that are packaged in dung pellets provide two surface area
configurations different from the original form of the food. The initial size and surface area of
the fecal pellet are functions of the rectal and cloacal organs and the anal cross-section. Species
specific pellet size determines potential interspecific coprophagic efficiency and provides a
unique microbial substrate within the soil microecosystem. As the pellet breaks down, a second
potential surface area increases dramatically as dung constituents are exposed. These materials
are the function of mastication, peristalsis and digestive activities and represent the maximum
size reduction of food eaten by a given consumer. These secondary particles, which are finely
divided, blend with the surrounding mineral particles and thus reflect the specificity of the
animal species on soil formation.

For example, the surface area of the fragments of herbaceous fabric comprising fecal pellets
of the cottontail rabbit, Sylvilagus floridanus, in central New York is 10 times greater than the
surface of the individual entire ovoid pellet (Figure 1). Such a modification in the vegetation of
the secondary fecal fragments deposited on or in the soil greatly increases the potential for
organic/inorganic interfaces. Microbial and decomposer invertebrate activity, which is vitally
important in the genesis of soil micromorphology, is stimulated.

SUMMARY

Understanding soil fabric production demands a holistic, cybernetic approach; this includes
a multivariate consideration of all physical, chemical and biological intricacies of the soil, both
macroscopic and microscopic, within ecological spheres of influence. The rhizosphere was one
of the First of these ecological spheres of influence to be recognized, illustrating the
microhabitat/microcommunity dynamics related to plant root systems. Phyllospheres have
been conceptualized to study aerial microhabitats and microcommunities on surfaces of living
leaves (Preece and Dickinson, 1971). We described the vermisphere (Hamilton and Dindal,
1983), another ecological sphere of influence within soil, which shows delicate biotic/abiotic

Quaest. Ent., 1985, 21 (4)

592

Dindal

POTENTIAL SURFACE AREA
COTTONTAIL RAl3ErT(c f&wabszois)
FECAL PELLETS

-1984

£A/r/%£ fn=4-)

SURFACS A%£A

Cx±S£)

*ZA± OtZ cm?

D/SSECPED pallets

HERBACEOUS J=A\3F^JO ^ _ «

2 ^biO^cm2

Kentucky slueg-tra^

OR^MARD Gr

■p/^K DELI 0^1

Jowl

Figure 1.

Soil animals and soil fabric production

593

interactions associated with earthworm burrows. Based upon unique soil microcommunities and
related microhabitats emphasized in this paper, I propose the following additional
microecosystem concepts to aid in research, communication and understanding of animal
involvement in soil fabric production:

1. Edaphophyllosphere ( = edaphic phyllosphere) - sphere of influence of the
fallen leaf and vegetative litter as a soil microhabitat,

2. Coprosphere - sphere of influence of vertebrate and invertebrate fecal
material as a soil microhabitat,

3. Necrosphere - sphere of influence of vertebrate and invertebrate carcasses
as soil microhabitats,

4. Nidusphere - sphere of influence of vertebrate and invertebrate nests, nest
sites and burrows as soil microhabitats.

The active result of the structure and function of the specific microcommunities inhabiting
each of these microecosystems governs the immediate soil fabric formation and plays an
ultimate influential role in the characteristic genesis and maintenance of any given soil.

REFERENCES

Anderson, J.M. 1978. Competition between two unrelated species of soil Cryptostigmata
(Acari) in experimental microcosms. J. Anim. Ecol. 47: 787-803.

Bal, L. 1982. Zoological ripening of soils. PUDOC, Wageningen, The Netherlands. 365 pp.
Brussard, L. 1985. A pedobiological study of the dung beetle, Typhaeus typhoeus. PhD Diss.

Agricultural Univ., 6700 AA Wageningen, The Netherlands. 168 pp.

Cox, G.W. 1984. The distribution and origin of Mima mound grasslands in San Diego County,
California. Ecology 65(5): 1397-1405.

Dillon, T.A. 1984. Analysis and ecology of organochlorine contaminant pathways in a carrion
microcommunity. Unpubl. MS Thesis. SUNY College Environm. Sci. Forestry, Syracuse,
NY. 188 pp.

Dindal, D.L. 1971. Ecology of Compost. Publications Office, SUNY College Environm. Sci.
Forestry, Syracuse, NY. 12 pp.

Dindal, D.L. 1978a. Microcommunities defined, pp. 2-6. In Dindal, D.L., (Editor). Soil
Microcommunities. CONF-71 1076. NTIS, Springfield, VA.

Dindal, D.L. 1978b. Soil organisms and stabilizing wastes. Compost Sci. /Land Utilization. J.
Waste Recycling. 19(4): 8-1 1.

Dindal, D.L. 1980. “The Decomposer Food Web”, Script and set of 70 color slides for
educational purposes. The JG Press, Emmaus, PA.

Dindal, D.L., L. Theoret and J-P. Moreau. 1978. Municipal wastewater irrigation: effects on
community ecology of soil invertebrates, pp. 197-205. In Sopper, W.E. and S.N. Kerr.
(Editors). Utilization of Municipal Sewage Effluent and Sludge on Forest and Disturbed
Land. Penn. State Univ. Press, University Park. 537 pp.

Hall, D.G. 1947. Blowflies of North America. Thomas Say Foundation Publ., Baltimore. 447

pp.

Hamilton, W.E. 1983. Impact of landspread sewage sludge on soil organisms and soil structure.

Unpubl. PhD Diss. SUNY College Environm. Sci. Forestry, Syracuse, NY 148 pp.

Hamilton, W.E. and D.L. Dindal. 1983. The vermisphere concept: earthworm activity and

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sewage sludge. BioCycle-J. Waste Recycling 25(5): 54-55.

Hassall, M. and S.P. Rushton. 1985. The adaptive significance of coprophagous behavior in the
terrestrial isopod Porcellio scaber. Pedobiologia 28(3): 169-176.

Jenny, H. 1980. The soil resource, origin and behavior. Springer- Verlag, NY. 377 pp.

Jones, R.L. and H.C. Hanson. 1985. Mineral licks, geophagy and biogeochemistry of North
American ungulates. Iowa St. Univ. Press, Ames. 302 pp.

Kenagy, G.J. and D.F. Hoyt. 1980. Reingestion of feces in rodents and its daily rhythmicity.
Oecologia 44(3): 403-409.

Kramer, D. C. 1973. Geophagy in Terrepene o. ornata Agassiz. J. Herpetology 7(2): 138.

Kubiena, W.L. 1948. Entwicklungslehre des Bodens. Springer, Vienna. 215 pp.

Preece, T.F. and C.H. Dickinson (Editors). 1971. Ecology of leaf surface microorganisms.
Academic Press, NY. 640 pp.

Shelford, V.E. 1913. Animal communities in temperate America. Univ. Chicago Press. 362 pp.

Simmons, K.E.L. 1983. Starlings eating dog faeces. Brit. Birds 76(9): 411.

Stevenson, B.G. 1983. Functional ecology of coprophagous insects. Unpubl. PhD Diss. SUNY
College Environm. Sci. Forestry, Syracuse, NY. 167 pp.

Tardiff, M.F. and D.L. Dindal. 1980. Annual carbon, nitrogen and calcium trends in litter and
surface soil of a mixed hardwood stand, pp. 529-541. In Dindal, D.L. (Editor). Soil Biology
as Related to Land Use Practices. USEPA, Office of Toxic Substances, Wash., D.C.
EPA-560/ 13-80-038.

Werner, M.R. 1984. Mound building ants in an old-field: effects on soil, vegetation and
microarthropods. Unpubl. MS Thesis. SUNY College Environm. Sci. Forestry, Syracuse,
NY 96 pp.

FAUNAL INFLUENCE ON SOIL MICROFABRICS AND OTHER SOIL PROPERTIES

A.R. Mermut

Department of Soil Science
University of Saskatchewan
Saskatoon, SKS7N0W0
CANADA

Quaestiones Entomologicae
21:595-608 1985

ABSTRACT

The influence of animal activity becomes visible by studying thin sections of soil with
microscopes. Several specific soil microfabrics ( organic laminae, lenticular fabric, lamellar
fabric, mesh fabric, cross hatching ) that are directly associated with animal activity are
recognized. However, more research is needed to understand the role of the fauna and specific
animals in the formation of soil fabrics. A restricted number of other micromorphological
features (fecal pellets, inorganic pellets, faunal tubules and chambers and mammilated
metavughs) identified so far are used only to indicate the presence of faunal activity in soils.
Animals may significantly alter the soil characteristics and have an important role in the
chemical decomposition of plant residues and the accumulation of nutrients in the biomass.
Their excrement, together with organic residues, is essential in the formation of soil
aggregates. Faunal turbation facilitates deep rooting, and consequently, higher production of
biomass.

RESUME

L’ influence de I’activite des animaux devient apparente lorsqu'on etudie au microscope des coupes fines du sol.
L'auteur reconnait plusieurs microstructures specifiques du sol qui sont directement associees avec des activites animates
(feuillets organiques, structure lenticulaire, structure lamellaire, structure en maille, double hachure). Toutefois, plus de
recherches seront necessaires pour comprendre le role de la faune dans son ensemble et celui d'animaux specifiques dans
la formation des diverses structures des sols. Un nombre limite d’autres traits micromorphologiques qui ont ete identifies
(boulettes fecales, boulettes inorganiques, petites galeries et chambres d'animaux, et metavughs mamelonnes) ne sont
pour I’instant utilises que pour indiquer la presence d’activite animate dans les sols. Les animaux peuvent alterer d’une
maniere significative les caracteristiques des sols et jouent un role important dans la decomposition chimique des residus
vegetaux et dans i accumulation des elements nutritifs dans la biomasse. Leurs excrements, combines avec les residus
organiques, sont essentiels a la formation des agregats des sols. Le remuement cause par la faune facilite I’encracinement
en profondeur et, par consequent, favorise la production d’une biomasse plus elevee.

INTRODUCTION

Every soil provides a habitat suitable for animal life. Because of the presence of large
quantities of living and dead plant and microbial materials which serve as a continuous food
resource, soil animals are generally restricted to the organic or mineral surface horizons.
However, animals may also be active at lower depths.

In his early studies, Kubiena (1938) pointed out that the association, activity and structure
of soil organisms are primarily controlled by the space condition, microclimate (temperature,
I moisture, air pressure, insolation, and air movement), pH and salt concentrations, and food
I conditions in the microhabitat. The heterogeneous nature of soil provides a variety of habitats
I in which animals can survive and reproduce.

596

Mermut

Aquatic forms such as protozoa and nematodes restrict their lives to the moist zone where
free or capillary water is available. Apart from these aquatic forms, soil animals are organized
into two major groups as suggested by Lee and Wood (1971): (i) animals unable to burrow, and
(ii) animals that burrow and reshape the soil. Because of the changes taking place in the soil
year-round such as heating, desiccation, freezing of the surface and/or sheltering from
predation, members of the second group have the ability to accommodate themselves quickly by
moving to a soil zone that meets their requirements. Despite the concentration of biological
activity in surface horizons (Bal, 1970; Whittaker, 1974; Bal, 1982), tracks of small soil
animals were found down to 150 cm, even in soils of arctic and subarctic regions (Federova and
Yarilova, 1972). In semi-arid climatic regions, many animals commonly occur down to 3 m
(Kubiena, 1953; Price and Benham, 1977; Valiakhmedov, 1977a, b).

Improvement in preparation techniques for thin sections of soil has made it possible to
directly observe and study the influence of biological activity and the role of animals in soil
genesis. Much of our present knowledge results from the pioneering studies and efforts of
Zachariae (1965, 1967) who described the development of humus forms by specific soil fauna.
Attempts were also made, especially in semi-arid and arid regions, to establish the relationship
between particular groups of animals and the soil types (Valiakhmedov, 1977a, b; Ghilarov,
1978). Because of the complexities in temperate humid regions, Bal (1982) found great
difficulties in the interpretation of the relationship between the soil and its community. The
major problem today is the lack of study in the interface between biology and pedology.

Several signs of animal activities are found in thin sections of soil. Excrements are one of the
easily recognized features which characterize the nature and the feeding habits of the animals.
Structures within the soil body such as chambers, including pupal chambers of soil dwelling
invertebrates described by Valiakhmedov (1977a, b) and pedotubules described by Brewer
(1976) are other important features that may be used in recognizing and understanding animal
activity in soil. As well, studies indicated that a certain part, or even an entire profile, can be
partly to completely reworked by animal activities (Buntley and Papendick, 1960; Mulders,
1969; de Meester, 1970; Valiakhmedov, 1977a, b; Bal, 1982). Such mixing processes are
referred to as “faunal pedoturbation” (Hole, 1961; Jongerius, 1970).

Despite the efforts made in the past, our present understanding of soil animals and their
effects on soil characteristics seems to be far from complete. This paper is aimed only at
elaborating on the present level of knowledge on micromorphological features that are formed
by animal activity. As an integral part of the interpretation of soil micromorphology, faunal
effects on other soil properties are also included in the present report.

FAUNAL INFLUENCE ON SOIL FABRIC

A wide range of soil microfabrics are directly associated with animal activity in soils.
However, at this stage there is a need for detailed micromorphological studies to fully
understand and recognize the special soil fabrics which are induced by the faunal activities. As
an attempt in this direction, the information noted below was summarized from a few available
studies dealing with the microstructure of termite mounds (Stoops, 1964; Lee and Wood, 1971,
Sleeman and Brewer, 1972 and Mermut et al., 1984). Microfabrics so far recognized in termite
mounds (landscape features that are entirely biologically produced) include organic laminae,
lenticular fabric, lamellar fabric, cross hatching and mesh fabric, which are considered to
reflect the process of construction of the termite nest, and comprise remnants of their activities

Soil microfabrics and other soil properties

597

Fig. 1. Organic laminae (dark areas) and lenticular-fabric. Macrotermes subhyalinus gallery wall from Kenya (plane
light). Fig. 2. Lenticular fabric formed by lenticular units. Macrotermes subhyalinus nursery section from Kenya (crossed
nicols).

598

Mermut

in the soil.

Organic Laminae

These consist of dark reddish brown to black, very weakly and strongly anisotropic material
occurring as bands commonly 20jum wide found adjacent to existing gallery walls (Fig. 1) or
within gallery fills. Their distinctive color allows an easy recognition and delineation of
boundaries with the microscope. Organic laminae may be characteristic for certain termite
species. They are richer in organic matter than the surrounding soil material. According to Lee
and Wood (1971), they are likely made of excrements with a semiliquid consistency. Organic
matter in some of the laminae is more highly humified than in others. More melanization,
considered as an indication of increased humification, suggests that this organic material has
passed through the gut of the termite.

Lenticular Fabric

The elongated lens-shaped units found in termite mounds were described as lenticular by
Stoops (1964). Strong welding of single small lenticular units may develop into a large
lenticular unit exceeding 5 mm in size (Mermut et al., 1984). Low content of skeleton grains
and high amount of inorganic plasma encourages the formation of larger pellets (Fig. 2). The
units consist of dominantly mineral plasma and some pedological features, skeleton grains and
little organic matter. Micromorphological observations indicate that each unit consists of
different proportions of plasma and skeleton grains. The fabric consists of strongly
accommodated lenticular units.

Lamellar Fabric

Fabric consisting of alternate parallel alignments of skeleton grains and plasma in which the
parallel arrangement is sometimes associated with planar voids was termed “lamellar
structure” by Stoops (1964) and “lamellar fabric” by Sleeman and Brewer (1972). This fabric
was found in soils which had high contents of both sand and silt. The skeleton grains are
embedded within the soil matrix; the plasma shows extremely well-developed masepic fabric.
Because of the parallel arrangement of clay domains, Stoops (1964) termed this fluidal
structure. The groundmass of such structures appears to be very dense. Aside from the
above-mentioned planar voids, there are no voids visible in such a fabric. The majority of the
colors of the groundmass in the lamellar fabrics are similar to the original soil plasma from
which the mound was built. However, addition of humic particles may cause the plasma to be
somewhat darker in color.

Cross Hatching

This may be considered a subtype of lamellar fabric in which two sets of parallel
arrangement of plasma and skeleton grains cross each other (sometimes 90°). The resulting
feature is like lattisepic fabric of Brewer (1976), which by definition considered only the
arrangement of the clay domains. This type of fabric is found near the gallery surfaces of
termite mounds (Mermut et al ., 1984) (Fig. 3). Our experience so far indicates that both
lamellar and cross-hatching can be considered to result from activity of burrowing animals.

Soil microfabrics and other soil properties

599

Mesh Fabric

This fabric type results from the specific arrangement of either spheroidal or lenticular
construction units. Each construction unit has a separation zone of plasma that can be
compared with skelsepic fabric (Brewer, 1976). Welding of individual units in a preferred
direction results in a type of plasma orientation resembling a mesh, called “mesh fabric” by
Mermut et al. (1984). If strong welding occurs in each construction layer, the borders of earlier
units appear more diffuse. This type of fabric is attributed to the homogeneity as well as high
content of plasma.

OTHER MICROMORPHOLOGICAL FEATURES FORMED BY ANIMAL ACTIVITY

This group includes fecal pellets, pellets built as construction units, faunal tubules and
chambers, and mammilated metavughs.

Fecal Pellets

Fecal pellets are the excreta that have left an animal’s intestines as shaped,
three-dimensional units (Bal, 1973). Recognizable fecal pellets can be seen in a pedotubule,
inorganic horizons (Fig. 4), or within large interconnected pore spaces. Unfortunately, little is
known about the morphological characteristics of the fecal pellets. Brewer (1976) suggested
that a major subdivision of fecal pellets can be based on the external shape. Bal (1973) was able
to distinguish five main groups: spherical (Fig. 5), ellipsoidal, cylindrical, platy and threadlike.
Bal suggested that one should study and describe the characteristics in the following order of
succession: shape, size, composition, basic distribution.

Easily recognizable fecal pellets are found in burrows (Fig. 6), tunnels or chambers, which
may extend deep into the profile. Therefore, this feature subjected to the proper recognition can
be used as an absolute indication of biological activity. However, as a result of a disturbance of
soil material, for example by pedoturbation, the fecal pellets may become embedded in the soil
material, and thus become difficult to identify and describe.

Pellets Built as Construction Units

Pellets built as construction units are termed “construction elements” by Stoops (1964).
Those units recognized in termite mounds are composed of skeleton grains, plasma, and
pedological features. They are spheroidal, ovoid or lens-shaped and vary in size from 125 /urn to
1000 /urn. Mermut et al. (1984) described two distinct types of pellets; one is highly isotropic,
light yellowish in color with more clay mineral plasma appearing to be oral pellets mixed with
saliva, and the other is brownish, slightly isotropic probably mineral plasma mixed with
excreta. According to Arshad (1981), some Macrotermes species selectively prefer fine soil
separates (particles less than 0.5 mm) to construct their mounds. Lee and Wood (1971)
indicated that in a Podzolic soil, the termites preferred to use clay-rich subsoil to encase their
mound. Stoops (1964) observed that, during restoration of q mound, the termites piled up the
little units of sand and clay, moistened with their saliva. Quick-drying of the outer crust of the
pellets creates a plasma separation around each unit, allowing their recognition, even when they
are extremely welded. This separation of plasma is comparable to the skelsepic plasmic fabric
of Brewer (1976). Except for granular aggregates found in the cracks of the Vertisola which
resemble these units, pellets can also be used as a sign of biological activity in non-selfmulching
soils. However, the random distribution pattern of the granular aggregates in the swelling clay

Quaest. Ent., 1985, 21 (4)

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Mermut

Fig. 3. Cross-hatching observed in a Macrotermes herus mound from Kenya (cross nicols). Fig. 4. Single organic
ellipsoidal fecal pellets from the organic horizon of a Luvisolic forest soil in Saskatchewan (plane light).

Soil microfabrics and other soil properties

601

Fig. 5. Single organic spherical melanized fecal pellets from the organic horizon of a Luvisolic forest soil in Saskatchewan
(plane light). Fig. 6. Fecal pellets around a zoogenic tube (an African termite burrow), as seen with a binocular
microscope.

Quaest. Ent., 1985,21 (4)

602

Mermut

soils may serve to differentiate the pellets even in these soils.

Faunal Tubules and Chambers

These are features which have sharp external boundaries and are generally larger than the
voids in the soils. Faunal tubules have a tubular external form and are made by soil animals
that can burrow and reshape the soil materials. Tubular-shaped voids can also be formed by
growing roots and may be confused with faunal tubules. Characteristics like wall lining (Bal,
1973) and imprints caused by the bodies of animals may be used to identify zoogenic tubes
(Jongerius and Reijmerink, 1963). Excreta in tubules support their origin as faunal burrows.

The burrowing of earthworms is more uniform and generally larger than the tubular-shaped
voids produced by growing roots. The voids created by earthworms are the same diameter as
their body. However, it is not always possible to use size of the burrow to identify the animals
responsible for construction of tubules in the soil. For example, termites and ants make a tubule
much larger (Fig. 6) than their bodies; however, we know now that these animals produce
special soil fabrics around their burrows.

Chambers differ from large voids in soils in that their walls are regular and smoothed (Fig.
7). Their formation has been attributed to faunal activity. In termite nests, chambers have
arched or domed roofs and relatively flat floors. As can be clearly seen in Fig. 6, animals (for
example, termites) sometimes coat the walls of their tunnels with clay, forming a smooth
cutanic feature. Dark staining of gallery walls which is a characteristic found in termite
mounds can also be attributable to faunal activity. Surface smoothing, discoloration of burrows,
cementing and dense packing and excreta in voids are other characteristics that can be
recognized in soil thin sections.

Mammilated Metavughs

As indicated by Brewer (1976), these special vughs have smoothed walls and mammilated
conformation. Mermut et al. (1984) found this special void in almost all the thin sections from
the African termite mounds. Brewer (1976) reports that mammilated vughs occur commonly in
soils with strong evidence of extensive faunal activity, especially earthworms. The voids are
formed by either welding of pellets used for mound construction (Fig. 8) or faunal excreta.
Coalescing fecal pellets and/or pellets used for construction may form empty spaces with sharp
protuberances. For pellets used for construction, the plasma reorientation around each unit is
clearly observed. Therefore, the soil matrix in contact with the vughs invariabily shows a
vosepic plasmic fabric. Mammilated metavughs often bear some additional marks (orderly
arrangement of units, darkening of soil material with increased density) which can be used for
further evaluations. Many mammilated vughs may serve as good indicators of animal activity
in soils.

FABRICS FORMED BY DECOMPOSITION OF ORGANIC MATTER

Soil animals play a very important role in fragmentation of litter and redistribution of
organic materials in soil. Excrements of different shape, size, color, orientation, and
composition indicate consumption of litter by soil fauna. Some of the animal species, as for
example enchytraeids (Zachariae 1964) consume decaying arthropod faeces, pierce and cleave
the compact excrement of big earthworms, and eat humus earth as well. Thus, decomposition is
a continuous process in which the animals work in close association. All plant residues and

Soil microfabrics and other soil properties

603

Fig. 7. Interconnected chambers built by an African termite species, as seen with a binocular microscope. Fig. 8
Mammilated metavughs formed by coalescing of inorganic pellets used for construction from an African termite mound
(crossed nicols).

Quaest. Ent., 1985, 21 (4)

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Mermut

Fig. 9. SEM micrograph of a comb from the nest of an African termite ( Macrotermes michaelseni). The comb appears to
be constructed from plant fragments.

especially excrements are also subject to further microbial decomposition. The effect of specific
soil animals on soil microflora (Arshad et al., 1982) is becoming an important area for
investigation. Fecal pellets are found with fungal hyphae, indicating their close association
during decomposition of organic matter.

According to Geyger (1967), excrement together with organic residues are essential in the
formation of soil structure, expecially in organic horizons depending on the decomposition stage
of organic matter and the admixture of inorganic plasma and skeleton grains. Brewer and
Pawluk (1975) have recognized certain types of granic and related fabrics, especially in the
organic horizons of some Canadian soils. Their concept is strictly based on morphology;
however, it can be related to soil genesis. It seems quite clear that more research is needed to
understand the role of the fauna in the formation of soil fabrics.

Soil animals in organic horizons have a role in chemical decomposition and humification of
plant residues and therefore accumulate nutrients in the biomass. Babel (1978) demonstrated
that rate of fabric differentiation increased from the L to F and decreased in the H horizons,
indicating the close positive relationship between humification and population of soil fauna.
Despite research efforts in this area, details of these processes are still unknown.

In tropical countries, termites are capable of decomposing up to a third of the fresh annual
grass, wood and leaf litter (Collins, 1981). This makes them the most important group of
invertebrates in the decomposition of organic matter in their natural habitat. It is interesting to
note that the fungi of the genus Termitomyces are the predominant microorganisms growing on
the comb in certain termite nests. As can also be seen in Fig. 9, the comb is made up of mostly
uningested plant materials (Rohrmann, 1978) and/or termite fecal pellets (Sand, 1960).
Fungus cultivated on the comb was of nutritional value for the termites. This indicates a
well-balanced cycle of organic matter in a harmony in which the termites have an important
role.

INFLUENCE ON OTHER SOIL PROPERTIES

About 1.5 million kinds of animals are living on more than a million kinds (equivalent to the
soil series, U.S. classification) of soils (Buol, et al. 1980). The action of soil fauna in
combination with plants on initial parent material is prerequisite to soil formation.

Soil microfabrics and other soil properties

605

Turbation of soils by animal activity (faunal pedoturbation) is generally well known to the
pedologist ( e.g ., Hoeksema, 1953; Slager, 1966). Krotovinas (Soil Survey Staff, 1951) are
common features of Chernozem and dark colored soils commonly developed under prairie
vegetation. They are the result of infillings of animal burrows by transportation of soil material
from any direction. This contributes to the process of soil homogenization. Krotovinas appear in
various sizes and are texturally and structurally unlike the surrounding soil materials. In
| semi-arid soils of Turkey, de Meester (1970) found that the contents of organic matter and
nitrogen of krotovinas and the root development in krotovinas were considerably higher than
the soil material of the horizon in which they existed.

Worm-worked soils are typical examples of how an animal group may significantly alter the
soil characteristics. Such unique soil profiles were reported in several places in North America
(e.g., Buntley and Papendick, 1960; Nielsen and Hole, 1964; Wilde, 1971). Striking differences
in macromorphological features and certain chemical characteristics are evident between the
worm-worked profiles and the non-affected adjacent areas. Horizon boundaries within the
turbation zone were obliterated. Buntley and Papendick (1960) clearly demonstrated that the B
horizon which had high clay content originally was reduced in clay with the materials
transported from both A and C horizons. As a result, the clay content of the A and C horizons
were increased, whereas the B horizon was decreased. There was a distinct increase in organic
matter and nitrogen content in the soils perforated with earthworms. Micropedological studies
showed argillams of argillic horizons intensely reworked by faunal activities (Jongerius, 1962).
We have observed similar effects in thin sections of some Kenyan Oxisols perforated by
termites. Preferential selection of clay for mound construction by some termite species (Stoops,
1964; Lee and Wood, 1971) can cause a complete mixing of all horizons in Podzolic soils and
soils with higher clay content in the subsoil than the topsoil.

It has long been recognized that the biological homogenization of soils is also of great
importance in land reclamation from the sea sediments (Hoeksema, 1953; Slager, 1966). In the
sediments of the famous Ijsselmeer polder, Bal (1982) observed that many channels which were
; formed by animals facilitated the deep rooting and consequently higher production of biomass.

Based on apparent lack of the original soil characteristics in worm-worked soils in South
Dakota, Buntley and Papendick (1960) suggested that such soil be named “Vermisol”. These
soils display the features resulting from intensive perforation of worms. The humus type of
forest soil mixed by earthworms is called a “Vermiol” (earthworm mull) by Wilde (1971). The
recognition of such soils resulted in the establishment of the three great groups, namely,
Vermiborolls, Vermudolls and Vermustolls in the Soil Taxonomy (Soil Survey Staff, 1975).
These soils have a mollic epipedon that, below any Ap horizon, has 50% or more by volume of
i wormholes, wormcasts, krotovinas, or filled animal burrows of especially earthworms and their
ji predators.

Because of the close relationship with soil organic matter, animals may play an important
i role in the reduction of water erosion. They create high water infiltration capacity as well as
li absorption due to production of organo-mineral complexes. In a detailed study, Arshad (1982)
found that the soils influenced by termites were high in nutrients. This, together with favorable
water availability and good drainage, resulted in a considerable increase in biomass in dry
tropical parts of Kenya.

One has to be very careful in sampling, measuring and evaluating the effects of soil animals
i on soil properties. Micromorphometric measurements on samples reworked by animal activities
(Mermut et al., 1984) showed that in some parts of the biologically disturbed areas, porosity

1 Quaest. Ent., 1985,21 (4)

606

Mermut

increased; however, compaction caused a considerable decrease in porosity in other parts.
Apparently, improper sampling is one of the causes of controversy among scientists on the
pattern of animal activity.

Among other influences, there is an increase of the availability of mineral nutrients and
their distribution within the rooting zone. For example, earthworms increased the availability
of N, Ca, Mg, K, P, and Mo (Nye, 1955) and Pb, Zn and Ca (Ireland, 1975).

CONCLUSIONS

Advances in soil micromorphology have made it possible to directly observe and study the
influence of soil animals on soil characteristics and the role of the animals in soil genesis.
Despite the studies and efforts of the past, our understanding of soil animals and their influence
on soil microstructure is far from complete. There is need for more detailed studies dealing with
micromorphological features and fabrics that are associated with special soil animals.

Soil fauna together with soil microorganisms have a very important role in accumulation,
decomposition and redistribution of organic matter in the soil. Some animals in the tropics are
capable of decomposing up to a third of the fresh annual grass, wood and leaf litter. Because of
the close relationship with soil organic matter, animals may play an important role in
increasing the aggregate stability and in reducing water erosion. They increase the availability
of mineral nutrients within the rooting zone and they play an important role in land
reclamation from sea sediments by homogenization.

Much knowledge on the effect of soil fauna on soil characteristics can be gained by
experimental studies. Breakdown of litter, formation of biopores, perforation and behavior of
each animal species can be determined in cultures under controlled laboratory conditions. With
the help of micromorphological studies, details of the features produced by animals can be
characterized. It is certain that without such experimental studies, present problems in
micromorphological identification, quantification and description of faunal activity in soils will
remain unsolved.

REFERENCES

Arshad, M.A. 1981. Physical and chemical properties of termite mounds of two species of
Macrotermes (Isoptera Termitidae) and the surrounding soils of the semiarid savanna of
Kenya. Soil Sci. 132: 161-174.

Arshad, M.A. 1982. Influence of the termite Macrotermes michaelseni (Sjost) on soil fertility
and vegetation in a semi-arid savannah ecosystem. Agro-Ecosystems 8: 47-58.

Arshad, M.A., N.K. Mureria, and S.O. Keya. 1982. Effect of termite activities on the soil
microflora. Pedobiologia 24: 161-167.

Babel, U. 1978. Some aspects of fabric dynamics in a modern profile (investigated by the
application of a graphical scheme), p. 169-187. In, M. Delgado (Editor). Proceedings of the
fifth International Working Meeting on Soil Micromorphology, May 24-28, 1977,
Granada, Spain.

Bal, L. 1970. Morphological investigation in two moder-humus profiles and the role of soil
fauna in their genesis. Geoderma 4: 5-36.

Bal, L. 1973. Micromorphological analysis of soils. Lower levels in the organization of organic
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Netherlands. 174 pp.

Bal, L. 1982. Zoological ripening of soils. Agricultural Research Reports 850. PUDIC,
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Brewer, R. and J.R. Sleeman. 1963. Petotubules: their definition, classification and
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Brewer, R. and S. Pawluk. 1975. Investigations of some soils developed on hummocks of the
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Brewer, R. 1976. Fabric and mineral analysis of soils. Robert E. Krieger Publishing Company,
Huntington, New York. 482 pp.

Buntley, G.J. and R.I. Papendick. 1960. Worm-worked soils of eastern South Dakota, their
morphology and classification. Soil Sci. Soc. Am. Proc. 24: 128-132.

Buol, S.W., F.D. Hole, and R.J. McCracken. 1980. Soil genesis and classification. The Iowa
State Univ. Press, Ames. 404 pp.

Collins, N.M. 1981. The role of termites in the decomposition of wood and leaf litter in the
Southern Guinea savanna of Nigeria. Oecologia (Berlin) 51: 389-399.

de Meester, T. 1970. Soils of the Great Konya Basin, Turkey. PUDOC, Research Reports 740,
Wageningen, The Netherlands.

Federova, N.N. and E.A. Yarilova. 1972. Morphology and genesis of prolonged seasonally
frozen soils of western Siberia. Geoderma 7: 1-13.

Franz, H. 1975. Die Bodenfauna der Erde in Biozonotischer Betrachtung. Erdwissenschaftliche
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Geyger, E. 1967. Die Structurbildungen in den Humushorizonten von Boden der hohen Sierra
Nevada (Spanien). Geoderma 1: 229-248.

Ghilarov, M.S. 1978. Bodenwirbellose als Indikatoren des Bodenhaushaltes und von
bodenbildenden Prozessen. Pedobiologia 18: 300-309.

Hoeksema, K.J. 1953. Die natuurlijke homogenisatie van het bodem profiel in Nederland Boor
en spade 6: 24-30.

Hole, F.D. 1961. A classification of pedoturbations and some other processes and factors of soil
formation in relation to isotropism and anisotropism. Soil Sci. 91 : 375-377.

Ireland, M.P. 1975. The effect of the earthworm Dendrobaena rubida on the solubility of lead,
zinc, and calcium in heavy metal contaminated soil in Wales. J. Soil Sci. 26: 313-318.

Jongerius, A. 1962. Recente vorderingen in de micropedoligie en haar mogelijkheden.
Landbouwk. Tijdschr. 74: 973-999.

Jongerius, A. and Reljmerink, A. 1963. Over de betekenis van de zogenaamde perforatiegraad
als onderdeel van het profielonderzoek. Boor Spade. 13: 178-193.

f Jongerius, A. 1970. Some morphological aspects of regrouping phenomena in Dutch soils.
Geoderma 4: 311-331.

Kubiena, W.L. 1938. Micropedology. Collegiate Press, Inc., Ames, Iowa. 243 pp.

I Kubiena, W.L. 1953. Bestimmungsbuch und Systematik der Boden Europas. Enke, Stuttgart.
392 pp.

! Lee, K.E. and T.G. Wood. 1971. Termites and soils. Academic Press, London and New York.
251 pp.

! Mermut, A.R., M.A. Arshad, and R.J. St. Arnaud. 1984. Micromorphological study of termite
mounds of three species of Macrotermes in Kenya. Soil Sci. Soc. Am. J. 48: 613-620.

i Mulders, M.A. 1969. The arid soils of the Balikh Basin (Syria). Ph D. Thesis, Soils

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Department of the State University Utrecht, Utrecht, The Netherlands. 196 pp.

Nielson, G.A. and F.D. Hole. 1964. Earthworms and the development of coprogenous A]
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Nye, P.H. 1955. Some soil-forming process in the humid tropics. 4. The action of the soil fauna.
J. Soil Sci. 6:73-83.

Price, D.W. and G.S. Benham. 1977. Vertical distribution of soil-inhabiting microarthropods in
an agricultural habitat in California. Environmental Entomology 6: 575-580.

Rohrmann, G.F. 1978. The origin, structure and nutritional importance of the comb in two
species of Macrotermitinae (Insecta, Isoptera). Pedobiologia. 18: 89-98.

Sands, W.A. 1960. The initiation of fungus comb construction in laboratory colonies of
Ancistrotermes guieneensis (Silvestri). Insectes Soc. 7: 251-259.

Slager, S. 1966. Morphological studies of some cultivated soils. PUDOC Agricultural Research
Reports 670. Wageningen, The Netherlands. Ill pp.

Sleeman, J.R. and R. Brewer. 1972. Micro-structures of some Australian Termite nests.
Pedobiologia 12: 347-373.

Soil Survey Staff. 1951. Soil survey manual. Agric. Handb. No. 18, USD A. U.S. Government
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Soil Survey Staff. 1975. Soil Taxonomy: a basic system of soil classification for making and
interpreting soil surveys. Agric. Handb. No. 436, USDA. U.S. Government Printing Office,
Washington, D.C. 754 pp.

Stoops, G. 1964. Application of some pedological methods to the analysis of termite mounds,
pp. 379-398. In A. Bouillon (Editor). Etudes sur les Termites Africains. Edit, de l’Univ.
Leopoldville.

Valiakhmedov, B. 1977a. Pupal chambers of soil-dwelling invertebrates in the Sierozems of
Tadzhikistan and their effect on the development of the soil profile. Soviet Soil Sci. 9:
184-191.

Valiakhmedov, B. 1977b. Spuren de Lebenstatigkeit von Bodenwirbellosen als diagnostische
Merkmale fur Subtypen von Grauen Wiistenboden in Tadschikistan. Pedobiologia 17:
60-69.

Whittaker, J.B. 1974. Interaction between fauna and microflora at Tundra sites, pp. 183-196.
In A.J. Holding et al. (Editors). Soil organisms and decomposition in Tundra. Proceedings
of the microbiology decomposition and invertebrate working groups meeting, Fairbanks,
Alaska, U.S.A., August 1973.

Wilde, S.A. 1971. Forest humus: its classification on a genetic basis. Soil Sci. Ill: 1-12.

Zachariae, G. 1964. Welche bedeutung haben Enchytraeen im Waldboden? pp. 57-68. In A.
Jongerius (Editor). Proc. Sec. Int. Working Meeting on Soil Micromorphology, Arnhem,
The Netherlands, September 22-25, 1964. Elsevier, Amsterdam.

Zachariae, G. 1965. Spuren tierischer Tatigkeit im Boden des Buchenwaldes.
Forstwissenschaftliche Forschungen 20. Parey, Hamburg. 68 pp.

Zachariae, G. 1967. Der Einsatz mikromorphologischer Methoden bie bodenzoologischen
Arbeiten. Geoderma 1: 175-195.

IN SITU LOCALIZATION OF ORGANIC MATTER IN SOILS

R.C. Foster

CSIRO Division of Soils
Glen Osmond

South Australia Quaestiones Entomologicae

AUSTRALIA 21:609-633 1985

ABSTRACT

Three main size classes of organic matter in soils are ingested by soil animals: (1),
multicellular plant and animal remnants (5000 - 50 /um in diameter); (2), microorganisms (50
- 0.3 jum); and (3), subcellular fragments (1 pm - 10 nm). Ultracytological techniques locate
particular reactive sites (polyphenolic, acidic and neutral carbohydrates, enzymes) on soil
organics in situ in natural soil fabrics. This allows investigation of morphological and
biochemical characteristics of organic resources available to soil animals and changes
involved in organic matter transformations as materials pass from class 1 to class 3. This
involves both breakdown of cellular materials and biosynthesis of new organics by
microorganisms. In particular, I address such questions as: where in soils are the different size
and biochemical types of organic matter located with respect to soil minerals and soil
microorganisms ; how are plant and animal tissues reduced to deposits of submicron and even
macromolecular sizes; how do biodegradable materials (e.g., polysaccharides) escape
microbial degradation; how do the physically and chemically protected organic matter
deposits differ in structure, biochemistry and location; how do microbial products bind soil
components into stable aggregates and how do these subsequently break down; how do these
changes affect their availability and nutrient status for soil animals?

RESUME

II existe dans les sols trois categories principales de grosseurs de particules organiques qui sont ingerees par les
animaux endoges: 1) des restes d'animaux et de plantes multicellulaires d’un diamdtre variant de > 5000 h 50 ^m. 2) des
microorganismes d’une taille variant de > 50 h 0.3 pm, et 3) des fragments subcellulaires variant de 1 pm d 10 p m. Des
techniques d’ultracytologie ont ete utilisees pour localiser des sites reactifs particuliers (hydrates de carbone
polyhenoliques, acides et netures, enzymes) sur des particules organiques en place dans les structures naturelles de sols.
Cette approche permet d’etudier les caracteristiques morphologiques et biochimiques des ressources organiques
disponibles pour les animaux endoges, de meme que les changements qui se produisent durant la transformation des
matieres organiques alors que les materiaux passent de la categorie I a la categorie 3. Cette transformation comprend la
decomposition des materiaux cellulaires et la biosynthdse de nouvelles substances organiques par les microorganismes.
Plus particulierement, je pose les questions suivantes: ou dans les sols se situent les differentes grosseurs et les differents
types biochimiques de matiere organique par rapport aux mineraux et aux microorganismes? Comment les tissus
animaux et vegetaux sont-ils decomposes en des depots de taille submicroscopique ou meme macromoleculaire?
Comment des substances biodegradables (comme les polysaccharides ) echappent-elles d la degradation microbienne?
Comment des depots de matiere organique proteges physiquement et chimiquement different-ils dans leur morphologie,
leur biochimie et leur position ? Comment les produits resultant de /’ action microbienne lient-ils les composantes du sol
en des agregats stables, et comment ceux-ci sont-ils par la suite decomposes? Comment ces changements affectent-ils leur
disponibilite et leur qualite nutritive pour la faune du sol?

610

Foster

INTRODUCTION

Little is known about organic matter in its native state in natural soil fabrics. Nearly all
information on soil organics comes from studies of materials which have been chemically or
physically extracted from the soil and introduced into a quite different biophysical and
biochemical milieu for characterisation and quantification. Detailed knowledge of the
structure, biochemistry, microbiology and location of organic materials in situ in soil fabrics is
of great importance to determine where the various phases of organic matter mineralization
occur and where nutrients are available to soil animals.

Ultrastructural studies of soil organics are concerned with particles from c.100 ^m to 10 nm,
so in investigations of the processes of organic matter decay and nutrient recycling,
ultrastructural studies bridge the gap between materials studied by the soil micromorphologist
and those studied by the soil biochemist.

Organic materials in soils are infinitely variable in their structure and biochemistry,
depending on their source and the amount of microbial decay and chemical weathering they
have undergone. They range in size and ultrastructural complexity from histons of plant and
animal tissues which are structurally almost unchanged from their living condition down to
fragments of almost macromolecular size which have undergone profound morphological and
biochemical transformations (Foster and Martin, 1981).

Although modern SEMs will take specimens weighing up to 1 kg, at useful magnifications
(say 10,000x) the amount of material represented in individual electron micrographs is
(<10'6cc [see below]) so that ultrastructural studies are limited to the finer details of effects of
soil animals on soil structure. Similarly the animals observed in electron micrographs of even
moderate magnifications must be restricted to microorganisms - ciliates, flagellates, amoebae
etc. Although these are very numerous in soils (106 /cc Darbyshire and Greaves, 1967), their
effects on soil organic processes were until recently, much neglected. In this paper I examine
the physical and biochemical environments near, and the location and structure of organic
materials available to soil animals, over a wide range of sizes.

METHODS

Electron optical methods

The study of soils by electron optical methods includes the use of conventional- (CTEM)
and scanning-transmission electron microscopy (STEM), scanning electron microscopy (SEM)
using either secondary electron, or back scattered electron detection (BSEI), and electron probe
microanalysis (EPMA) using energy (EDXRA) or wavelength (WDXPA) dispersive X-ray
analysis, (Bisdom, 1983). Unfortunately, biological materials in soils consist of exotic
molecular species made up of a rather limited number of kinds of atom so elemental analysis by
EPMA is not widely applicable to biological problems (Hayes, 1980). Conventional EPMA
instruments do not easily detect light elements such as C, N, and H, of which most organics of
biological importance are composed. EPMA has been used however to investigate the
distribution of P, K, Ca and Mg in roots and rhizospheres (Tan and Nopamornbodi, 1981).
This review is confined to TEM and SEM studies of soil organics in situ in natural soil fabrics:
Smart and Tovey (1981) and Bisdom (1983) provide excellent reviews of the submicroscopy of
the mineral components.

Organic matter in soils

611

Physical dimensions of specimens

Theoretically, the early stages of organic matter broken down to particles of micron size can
be studied by light microscopy using thin sections of soil but practically, section thickness and
the presence of opaque minerals and organic matter limits resolution to about 5-10 microns.
The use of conventional heavy metal staining methods and TEM of ultrathin sections has
allowed the detection of particles down to nanometer sizes in situ in natural soil fabrics (Foster
and Martin 1981). At these sizes, however, except where the organics have a distinctive
structure (membranes, microfibrils etc.) it is difficult to distinguish between organic and
inorganic particles. Bisdom (1983) summarises the application of more sophisticated
techniques for the identification of materials in soil samples, (ion microprobe mass analysis
(IMMA), secondary ion mass spectrometry (SIMS), laser microprobe mass analysis (LAMNA
etc.) which may be useful in distinguishing between organic and inorganic amorphous
materials. Such sophisticated techniques are not generally available to soil scientists, however,
so I have used ultracytochemical techniques to investigate the biochemical properties of the
small fragments of organic matter in situ in soil fabrics.

SEM specimens may be up to 1 kg in size, but the area sampled in an electron micrograph
depends on magnification and is usually quite small. For CTEM and STEM specimen size is
limited by the distance fixatives and embedding media will penetrate. Blocks of soil only 0.5 - 1
mm cubed give the best results. The actual sections are 0.5 x 0.5 mm and 0.1 /urn thick.

Physical and chemical stabilization of the soil fabric

Various components in plant and animal materials are naturally held together to form
tissues. Similarly, interlocking crystals hold sections of rock samples together. Soils, on the
other hand, are composed of randomly disposed and relatively widely spaced minerals, organic
fragments and soil microorganisms lying free or only loosely interconnected. Hence, except for
apical and sub-apical rhizospheres where the soil fabric is embedded in mucigel (Foster, 1981b;
Campbell and Porter, 1983), before ultracytochemical analysis can begin, the soil must be
stabilized both physically and chemically. Physical stabilization prevents relative movement of
soil components during biochemical processes. It is achieved by embedding the soil sample in an
amorphous gel such as gelatine or agar. Chemical stabilization prevents the loss of soluble
components (lipids, low molecular weight gels etc.) during solvent exchange dehydration and is
achieved with cross-linking agents such as aldehydes and/or polyvalent metals such as
lanthanum. For ultramicrotomy the soil must be dehydrated and embedded in plastic (see
Foster and Martin, 1981; Smart and Tovey, 1981 for details of techniques for soil specimens).

Ultracytochemistry

Ultracytochemistry is the detection and/or identification of (usually organic) materials in
biological tissues by electron optical techniques. Here I use the term for any organic deposit
whether part of a cell or free in the soil fabric. Ultracytochemistry has been used in biology for
more than 30 years. At its simplest, it merely consists of adding solutions of heavy metals
(typically Os, Pb, U) to soil samples. These react with, or are absorbed onto organics so that in
ultrathin sections (50 nm - 100 nm thick) where they were previously electron transparent (and
therefore invisible) they become electron opaque and so readily detectable.

Techniques for specific complex molecules.— Ultracytological techniques have the
advantage that specific complex macromolecules with well defined biochemical properties can
be detected and located in situ in a soil fabric section with a resolution measured in

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Foster

nanmometers. For example, histochemical methods have been devised for examples from every
major group of enzymes (Hayat, 1975). All the methods used here are from Sexton and Hall
(1978).

Techniques for particular chemical groups. — Hayes (1980) has suggested that examples of
pure biochemicals such as particular lipids, proteins, polyphenols and carbohydrates which can
be isolated from living cells may be very uncommon in soils. Instead, uncontrolled enzymatic
reactions during cellular autolysis and chemical reactions between the lysates, soil minerals and
pre-existing soil organics, produce random combinations of these materials to form unique
complexes ( e.g ., humates) which no longer closely resemble the biochemicals found in living
tissues.

Fortunately most ultracytological reagents detect particular reactive groups e.g., 0s04 reacts
with phenolic hydroxyls, alkyl groups, sulphydryl groups (for groups derived from plant
residues see Bland et al., 1971) so that conventional aldehyde/0s04 treatment detects plant cell
wall remnants and humic material. The acidic polysaccharides of mucins, bacterial and fungal
slimes and root mucilages are stained with ruthenium red or lanthanum hydroxide (Foster,
1981b).

Where they are not present naturally, specific reactive groups can sometimes be
experimentally generated by chemical pretreatments. Thus neutral carbohydrates are detected
by partial lysis with periodic acid to generate aldehyde groups which are then labeled directly
with silver methenamin (PAMS) (Pickett-Heaps, 1967) or via thiosemicarbazide with silver
proteinate (PATSP-Thiery, 1967). These methods give electron-dense deposits with a
characteristic structure so that carbohydrates can be detected even against a background of soil
mineral fragments, and have been used to demonstrate carbohydrate coatings in clay fabrics
(Foster, 1981a). Similarly, complex epoxides which occur in leaf (Brown and Holloway, 1981)
and root cuticles are detected by iodenation and addition of suitable silver compounds. Hence a
wide range of biologically significant materials can be positively and specifically located in or
on organic particles in soils.

Detection by specific removal. — Other organic materials can be located by noting sites
where staining disappears if the section is treated with a solvent or an enzyme. Thus Heritage
and Foster (in press) identified native S grains in sulphur bacteria in sections of waterlogged
soil by their solubility in CS2. Since specific, highly purified enzymes are available
commercially this has some potential, but does not seem to have been applied to soil
components other than recognisable tissues (e.g., mycelial strands, Foster, 1981c).

SOURCES OF ORGANIC MATTER IN SOILS
Newly deposited materials

The most common material entering soils is carbohydrates derived from leaves, branches,
bark fragments and fragments and floral parts (especially pollen), and from root mucilages,
exudates, and ephemeral root tissues.

Materials from aerial organs. — Depending on the depth from which the sample is taken,
leaf fragments retain much of their characteristic cellular structure even though their tissues
have been invaded by microorganisms (Plate la, b). Autolysis before leaf fall results in the loss
of cytoplasm in many cells, and release of vaculolar polyphenolics causes the cell walls to be
impregnated with materials which make them more electron dense after heavy metal staining
than would occur in the live leaf (Plate lc). Pine needles often contain extensive deposits of
polyphenols which partially occlude cell lumens and stain the cell walls (Plate la, b). Most of

Organic matter in soils

613

the carbohydrates are quickly removed by microorganisms so the cell walls collapse onto the
vacuolar contents (Plate Id). Eventually only much convoluted, lignified cell wall layers remain
and the origin of the material becomes indeterminate (Plate le).

Materials from Roots. — One of the more interesting facts to emerge in the last 10 years is
that roots deposit large amounts of organic matter into the soil whilst they are still functional.
Up to 30% of the photosynthate reaching the root may be released into the rhizosphere (Barber
and Martin, 1976; Martin, 1977; Martin and Puckridge, 1981) as gels, exudates and lysates
(see Rovira et al., 1979, for definitions). Most classes of plant metabolites (sugars, amino acids,
vitamins, proteins, lipids, hormones etc.) have been isolated from root exudates (Rovira, 1965)
but these are not preserved in preparation for electron microscopy. However they support
bacteria and fungi (Plate 2a, b) which colonise the complex carbohydrates secreted by the root
in the form of mucilages and proteins ( e.g ., enzymes). In some cereal crops more carbohydrate
may enter the soil as root mucilage than is stored in the grain as starch, (Samtsevitch, 1965).
For example White (1983) estimates sloughed cells and gel amount to 3.5 tonnes/ha/yr for
wheat.

Direct evidence from electron microscopy (Plate 2a) (Foster, 1981b; Foster et al., 1983;
Campbell and Porter, 1982) and theoretical calculations (Newman and Watson, 1977; Gardner
et al., 1983) suggest that these materials are mainly confined to the immediate vicinity of the
root (0 - 150 fim). Tan and Nopamorabodi (1981) found a sharp break in P distribution
between 200 - 300 /urn from the root surface which may also indicate the outer limits of the
rhizosphere gel. Using quite independant ultrastructural techniques Campbell and Porter
(1982) and Foster (1981b) showed that there was an inner layer of mucilage near the cell
surface (Plate 2a, b) which was much more dense than that 20 - 50 nm away; so, there may be
partition of root products with distance from the root on the basis of molecular weight.
Carbohydrates are neither preserved nor stained by conventional biological preparation
techniques (Foster and Martin, 1981) but the acidic carbohydrates are preserved and stained
by lanthanum hydroxide (Plate 2a) and the neutral carbohydrates by the PATSP (Plate 2b)
and PAMS reactions (Plate 2c). These electron micrographs of known plant materials serve to
calibrate these cytological tests for non-rhizosphere soils to be presented later.

All these root-derived organics are available to those animals such as collembolans,
nematodes and enchytraeid worms which browse along roots (Head, 1967). At first, root
mucilage may be enclosed by a cuticle (Greaves and Darbyshire, 1972), but this is soon
ruptured (Foster, 1981b) allowing the gel to penetrate into the soil fabric. Mucilage appears to
be a true gel, allowing water and ions to diffuse through it in a manner not significantly
different from that in free water (Greenland, 1979). Although most of the root gel is secreted
by the root cap, epidermal cells and root hairs also secrete mucilage. In drying soil, this may
hold soil firmly to form a rhizosheath (Wallstein and Pratt, 1981). At first, colonies of bacteria
develop in the soil surrounding the root in response to exudates penetrating the soil fabric.
Later, bacteria and fungi attack the gel, especially along the grooves between the epidermal
cells, leaving lysis holes (Plate 2a, b, c) in the mucilage.

In some semi-permanent grasslands, 53 - 98% of the standing crop is below the ground, and
some grasses show a 100% root turnover each year (Dickinson, 1982) amounting to 5000 kg dry
matter/ha/y (Whitehead et al., 1980), so root tissues may be a considerable component of the
annual organic matter input into soils. Little wonder then that Curry and Ganley (1977) found
89,000 microarthropods/sq meter, 80% being acarines and collembolans. Recent work has
shown that death of the root cortex is a normal phenomenon unassociated with disease (Henry

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and Deacon, 1981, van Vuurde et al., 1979). Materials released by the death of cells causes a
population explosion amongst the root surface microorganisms. Thus van Vuurde et al. (1979)
found that in root segments 4 - 5 days old, where 35 - 45% of the cells were dead, supported a 1
- 4% microbial cover. When 45 - 75% of the cells were dead, (segments 7-8 days old),
microbial cover increased to 8% of the root surface area.

Animals and their remains. — Most larger animals (here > 1mm!) escape during
preparation, though nematodes may be observed in dead root fragments (Plate 3a), and soil
ingesting animals recognised by clay and organic particles in their gut (Plate 3d), are
occasionally encountered. The most common animal remains in ultrathin sections of soil are
exuviae fragments and cast off appendages (Plate 3b, c, e, f). These are identified by their
characteristic structure, ultrastructure and histochemical reactions (Foster, 1978; Foster and
Martin, 1981). Live animals most commonly observed in soil sections are microorganisms such
as amoebae and cilliates, (Plate 3 g, h) etc.

The abundance of soil animals in the surface layers of soils means that fecal pellets are of
common occurrence, but they are not necessarily all derived from soil inhabiting animals; in
some forests there is an almost continuous rain of pellets released by phytophagous insects in
the canopy. Fecal pellets are recognised in SEM by their characteristic size and shape and some
can be attributed to particular species. In TEM, fecal pellets and their fragments are easily
recognised because their high enzyme and/or mucin contents make them stain strongly with
heavy metals. The chaotic arrangment of their contents is characteristic, and often they contain
cellular materials which are so little altered by digestive processes that cellular organelles (such
as the thylakoids of chloroplasts) can still be recognised (Plate 3i). Many pellets contain
bacteria, or support bacterial colonies in the surrounding soil (Foster et al ., 1983).

Further microbial breakdown. — Once incorporated in the soil, rapid degradation of tissues
occurs through the action of plasmodia, fungi and bacteria. In turn, fungal hyphae are broken
down by bacteria (Kilbertus and Reisinger (1975). Resins and polyphenolics from pines may be
deposited in the soil (Foster and Marks, 1967). Kilbertus and Reisinger (1975) examined the
stages in breakdown of leaf litter at the ultrastructural level. In clay soils most bacteria are
associated with organic matter (Plate 4a - d). The larger bacteria are associated with cell wall
remnants which still contain carbohydrate (electron transparent materials) (Plate 4a, c), but
throughout the humified organic matter and even in the mineral rich parts of the fabric, there
are many small microorganisms, many 3 fim in diameter (Plate 4a, b, d). Gradually organic
and mineral soil components become intimately mixed. Firstly clay platelets become absorbed
onto the gels secreted both by roots (Plate 2c) and bacteria (Plate 5a, b). Secondly fungal
hyphae, root hairs and pieces of plant cell walls tens of microns in length become enclosed in
extensive clay fabrics several microns thick (Plate 5c, f, g). The same is true for amorphous
materials (Plate 5d, e, h, i). These materials will be physically protected from microbial attack
until they are ingested and broken open in the alimentary tract of soil animals. They constitute
part of the physically protected organic pool in soils.

Secondary sources of organic matter in soils

Microbial tissues and their secondary metabolites. — All the materials previously
mentioned are further modified by microorganisms. Bacteria, actinomycetes and fungi are the
most commonly encountered microflora and their lytic activities may be a prerequirement
before tissues become available to soil animals.

Organic matter in soils

Plate 1. Primary sources of soil organic matter leaves.- (a). Although most of the tissue has been replaced by
microorganisms, the thickness of the cell wall indicates that the tissue was a leaf epidermis, (b). Detail of (a). The former
cuticle is occupied by hyphae. The cell lumen is partly occluded by tannins(T). (c). Later stage of decay-only polyphenol
rich cell wall remnants remain, (d). Most of the electron transparent carbohydrates have been removed from the cell walls
so the cells have collapsed onto the vacuolar tannins(T). (e). Highly decomposed leaf tissue from a waterlogged pasture.
Only distorted, humified cell wall layers remain.

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Plate 2. Primary sources of organic matter- roots - (a). Lanthanum hydroxide reacts with substituted carbohydrates in the
root surface mucilage (RM) which has been partially lysed (L) by soil bacteria. The mucilage holds cell wall
remnants(W), clay particles (C) and a colony of bacteria (B) onto the root surface, (b). Neutral carbohydrates in the root
mucilage (RM) are demonstrated by the PATSP technique. The gel and exudates support colonies of bacteria (B) near the
root. Quartz grains (Q) shatter during ultramicrotomy, (c). The cell wall of both root (W) and bacteria (B) are intensely
stained by the PAMS reaction but the root gel is only lightly stained. Clay particles(C) and quartz grains become
embedded in the gel which is locally lysed by bacteria (L). (d). Even where root mucilage (RM) is separated from the root
(W) by clay it can be recognised by its characteristic granularity and reaction with ruthenium red/0s04 complex, (e).
Extensive decay by colonies (B) and individual microorganisms (arrows) leads to the collapse of the root tissue.

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617

Plate 3. Soil animals and their remains - Soil animals which play a major role in organic matter mineralization, (a).
Nematode (?) in dead root fragment, (b). Appendage with live cells, (c). Empty appendages, (d). Soil ingesting nematode
(?) showing several sections of the gut (G) with clay particles and organic matter, (c), and (0- Cast off and partially
broken down insect parts, (g). An amoeba (A) in an organic rich surface soil. (h). A ciliate attached to a rhizomorph. (i).
A fecal pellet (FP) recognised by its chaotic and electron dense contents, supports bacteria (B). Note nearby cell wall
remnants (arrow heads) and membrane systems (arrows).

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Plate 4. Soil fabrics with microorganisms and organic matter remnants - (a). Bacteria (B) are usually associated with
organic matter which still contains carbohydrate (eg. cell walls W). Other highly lignified and convoluted organic matter
(0) does not support bacteria, but there are numerous microorganisms (0.3 diameter) scattered throughout the clay
(*). P is a pore about 1 /im in diameter, (b). Similar to (a). (0) is amorphous organic matter. Again there are numerous
small soil bacteria in the clay (*). (c). Detail of (a) showing bacteria enclosed in capsule material (Ca) which is not stained
by 0s04.(*) indicates a capsule-less bacterium enclosed in humified organic matter, (d). A less consolidated clay fabric
containing remnants of cell wall remnants (0) and an occasional microorganism (*).

Organic matter in soils

619

Skinner et al. (1952) estimated that conventional plating techniques used by soil
microbiologists only detected between 0.1 and 1% of the organisms present. Morever according
to Bae et al. (1972) more than 63% of soil microorganisms are less than 0.3 /urn in diameter and
so not readily seen by light microscopy. Campbell and Rovira (1973) showed that because
bacteria were enclosed in gel they are not readily detected by SEM and TEM of thin sections is
necessary to clearly see all the microorganisms in a soil and to determine their probable
viability (as indicated by their cytoplasmic ultrastructure).

The abundance of mucilages and exudates in young rhizospheres stimulates the growth and
division of bacteria, fungi and actinomycetes both in and around the root. The more readily
available exudates are used by less specialised bacteria such as the fluorescent pseudomonads
which are particularly common in rhizospheres. A host of fungi, epiphytes, symbionts, cortical
and stelar parasites, then colonise the roots and rhizospheres. These are followed by
saprophytes, and together with bacteria, they remove the less resistant polymeric materials.
Using electron microscopy, Foster and Rovira (1976) showed that a consortium (White, 1983)
of different microorganisms were involved in removing different chemical fractions of the cell
wall. Later, actinomyctes and bacilli become more abundant; these are able to attack more
resistant materials such as lignified secondary walls of tracheids, sclerenchyma etc.
Rhizosphere microorganisms reach population of 1.210 E+10/cc of rhizosphere soil at the
rhizoplane (Foster and Marks, 1967; Malajczuk, 1979) and attract not only other
microorganisms (flagellates, amoebae, parasitic bacteria and viruses) but also larger animals
such as mites, collembolans, nematodes etc. As well as feeding on the rhizosphere microflora,
these may also remove the partially decomposed cortical tissues leading to the complete
decortication of the root (Head, 1967).

Many of the exudates and lysates escaping from roots are used by the microorganisms in
respiration and growth, but some microorganisms secrete new organics which act as secondary
sources of energy for other soil inhabitants. With the development of the rhizoflora, microbial
gels become inextricably mixed with the root-derived gels so that they are no longer
morphologically or biochemically distinguishable: this complex colloidal carbohydrate mix was
named mucigel by Jenny and Grossenbacher (1963). Mucigel and microbial polysaccharides
have an important role in stabilizing soils (Martin, 1971; Forster, 1979; Gaspari-Mago et al.,
1979). Silt sized particles are bound to form larger aggregates by fungal hyphae. Clays become
bound onto the surface of small colonies of bacteria both in the rhizosphere (Foster and Rovira,
1978) and bulk soil (Kilbertus and Reisinger, 1978). Some microbial gums are particularly
resistant to breakdown (Greenland and Oades, 1975). Even after the death of
capsule-producing bacteria, the carbohydrate fibrils may persist in the soil, (Plate 6e) binding
the various mineral and organic components of soil crumbs together (Foster, 198 Id; Foster et
al., 1983). Turchenek and Oades (1978) consider that, within aggregates, bacterial gel is the
most important stabilizing agent as it binds clay particles into silt sized aggregates (Plate 6e,
g). The binding action of microbial gels varies with the concentration of uronic acids (Martin
and Aldrich, 1955).

The rhizosphere microflora Some fix nitrogen; others affect total root length, frequency of
laterals, number, length and distribution of root hairs, number of layers in the root cortex and
hence root diameter (see Foster, 1984, for review). As well, rhizosphere products release P, K,
Fe etc. from insoluble minerals (Moghimi et al., 1978).

Except in the surface layers where all the soil may come under the influence of the root,
roots may occupy less than 6% of the soil volume. Elsewhere, organic energy sources are

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Foster

confined to local remnants of organic matter, but despite the oligotrophic conditions in the
non-rhizosphere (bulk) soil (Pointdexter, 1981), there are still large populations of
microorganisms (109E + 9/gm), and Clarholm and Rosswall (1980) show that in forest soils
their numbers are more than sufficient to account for nearly all the organic C accession each
year, and they consider that even under the most favourable conditions, only 1 5 - 30% of the
bacteria were active. Although microbial populations in soil are so large, bacteria occupy less
than 0.01% of the total soil surface (Grey et al. , 1968). They are not randomly distributed in
soils, but are usually associated with substantial organic matter deposits. Thus, Gray et al.
(1968) showed that in a sandy soil, 60% of the bacteria were attached to organic particles even
though these composed only 15% of the soil volume. Similarly in large-area, ultrathin sections
of soils Foster et al. (1983) showed that the larger (.5 - 1 /nm diameter) microbial cells were
associated with cellular debris which still contained carbohydrate (Plate 4a, b). Bacteria were
also associated with highly lignified materials and occurred sparsely scattered in clay fabrics,
but these cells were generally much smaller in diameter (0.3 /im diameter) and contained less
stored food reserves such as polyhydroxybutyrate and polyphosphate than similar cells in the
rhizosphere (Foster, 1978).

In an extensive ultrastructural study of a single aggregate, Kilbertus (1980) showed that
within aggregates bacteria occured within three different types of pore, those with single
openings, those with multiple openings and those which were totally enclosed. He also showed
that a minimal size of pore could be colonised by bacteria which bore a constant ratio to the
diameter of the microorganism. Bacteria within the aggregates greater than 2 mm may
experience anaerobic conditions (Greenwood and Goodman, 1967).

Bacterial gels and microbial slimes are recognised by their fibrous (Plate 6b) or granular
(Plate 6c) texture, but extensive tracts of amorphous non-Os reactive gels are found in some soil
sections (Plate 6a). These may be secreted by earthworms.

Some fungi deposit resistant substances such as melanins in their walls. As well as ribosomes
rich in N and P bacteria may deposit storage materials eg. lipids, polysaccharides and
polyphosphates in their cytoplasm. The rhizosphere microflora becomes a valuable secondary
resource for the soil microfauna. Microfloral use of simple sugars in root exudates causes
immobilization of inorganic nitrogen. It has been shown using “microcosms”, (small plants
growing in a defined medium to which bacteria, amoebae, flagellates, nematodes etc. can be
added singly or in combination), that addition of predators and browsers, especially mites and
nematodes, markedly increases that amount of nitrate nitrogen available to the host plant
(Wood et al., 1982; Elliot et al., 1979). Chakraborty et al. (1983) showed that soil amoebae
attack Gaeumannomyces hyphae so they may be important in the biological control of soil
borne plant pathogens.

ORGANICS OF SUBMICRON SIZE

By the time many organic fragments are reduced to micron and sub-micron size they may
be so modified morphologically and biochemically that their origin may be obscure. They are
then best classified by their ultrastructure and histochemistry. Other particles may retain
sufficient characteristic fine structural features that their origin is more certain.

Organic matter in soils

621

Fibrous or lamellate materials

Many plant cell walls are composed of alternate carbohydrate-rich (electron-transparent)
and lignin-rich (electron-dense) lamellae so wall fragments can be recognised on the basis of
their distinctive multilamellate structure even when the fragments are less than 1 n m wide,
(Plates 5c, g, 6d). Remnants of terminal and middle lamellae are characteristic in their
dimensions, texture and electron density and are relatively resistant to decay (Plate 5f, i). As
carbohydrate is removed from cell wall fragments and phenolic hydroxyl groups are unmasked,
the remnants take up more and more metal ions either from the soil solution ( e.g ., Mn Fe Al) or
from electron dense stains and appear progressively more electron dens. Removal of the
carbohydrates brings the elctron dense lignin rich lamellae closer together. Finally only the
lignin skeleton remains, distinguishable by its osmiophilic, fibrillar structure.

In contrast, fibrillar remnants which do not react with Os but which are demonstrated by
PAMS, PATSP or Au-labeled lectins may be recognised as filamentous polysaccharides such
as cellulose microfibrils from higher plant cell wall remnants, or fibrils from the extracellular
polysaccharide (ECP) layers of bacteria, actinomycetes or fungi. Some root surfaces are
naturally fibrillar (Leppard and Ramomorthy, 1975; Roland, 1971) and Foster (1982) showed
that in later stages of decay fibrils from internal cell wall layers are exposed by microbial or
physical weathering and make contact with nearby soil minerals.

“Amorphous” or granular materials

Many materials which appear amorphous by light microscopy appear to be granular at the
higher resolution of the TEM, especially after suitable histochemical treatment.

Carbohydrates. — Granular deposits in soil fabrics revealed by ruthenium/0s04 or
La(OH)3 are probably microbial gels or remnants of root mucilages. In their freshly-formed,
fluid state, root mucilages flow freely into pores of submicron size in clay fabrics (Plate 2d) and
if these pores are too small to admit bacteria they may be physically protected by the clay from
microbial decay. Some of these deposits enclosed between clay tactoids are less than 0.5 n m
across (Plate 8g, h) (Foster 1981a, Emerson et al., in press), and although the individual
deposits are of small volume, they may be numerous, and thus contain significant energy
resources. Other materials reactive to PATSP or PAMS are associated with bacterial walls
(Plate 7a, c, g) or are the remains of cell-wall materials of higher plants (Plates 6g, d), but
other deposits are not associated with morphologically distinct remnants (Plates 7c, d, 8f).
These probably represent fragments of root or microbial ECP. Finch et al. (1971) and Griffin
(1981) consider that carbohydrate gels can act as water reservoirs. Rovira and Greacen (1957)
and Powlson (1980) showed that physically disturbing soils increase their respiration. They
proposed that grinding brings bacteria into contact with organic matter from which they were
previously physically separated. It is not clear whether mere grinding would be sufficient to
expose sub-micron sized deposits. Conformational changes due to dehydration or heating, (as in
Australian surface soils), may further reduce the susceptibility of these deposits to decay, and
according to Emerson (1977) when polysaccharides are bound to clays they become less
succeptible to periodate oxidation. Both sectioned material and isolated clay particles show a
very patchy reaction to PAMS (Plate 7d - 0 but whether negative reactions are due to absence
of carbohydate or to their stabilization on the clay is not clear at present. The complete
mineralisation of carbohydrates in and on clays probably depends on their release by grinding
in the gizzards of such soil animals as earthworms.

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Foster

Rybicka (1981) states that periodic acid used in the PATSF reaction specifically oxidises
the 1, 2-gycols in polysaccharides to form aldehydes which subsequently react with
thiosemicarbazide. The thiol group then reacts with silver to form an electron dense conjugate.
The reaction has been widely used in the biomedical sciences to locate carbohydrates, but only
rarely on studies of decomposition of organic materials in soils (Foster, 1981a).

The extracellular polysaccharides (ECP) of fungi and bacteria, whether they are granular or
fibrillar, can be distinguished by their different resistance to periodate hydrolysis. By treating
serial sections of the same soil fabric with periodate for different times before applying the
PAMS reagent, Foster (1981a) showed that fungal ECP’s were more resistant than those
produced by bacteria or roots. Similarly, by incorporating 14C-labeled bacteria and fungi into
a grassland soil, Nakas and Klein (1979) showed that bacterial cell walls and polysaccharides
were more quickly mineralised than those of the fungi. This may explain why carbohydrates
under pastures (rich in root mucilages and bacterial ECP) are more susceptible to periodate
degradation than those under forests (Clapp and Emerson, 1972), where most roots are
mycorrhizal. Foster (1981c) found that enzymes that removed the cell wall polysaccharides
from the thick- walled hyphae of mycelial strands from a forest soil failed to remove the ECP
(Plate 8d, e). This may mean that fungal gels are less readily available to soil animals.

Humic substances. — With the fragmentation of cells, polyphenols from cell walls (Harkin
1967) and vacuoles (Zucher, 1983) are released into the soil, so that rhizosphere soils are richer
in polyphenols than the bulk soil (Bokhari et al., 1979). Several studies have shown that
catalytic polymerisation of phenolic materials by clay minerals occurs to form materials with
the properties of humic and fulvic acids (Wang et al ., 1980), and humic materials account fo
60 - 70% of soil C (Griffith and Schnitzer, 1975). The importance of these products lies in the
N they contain. Ladd (1981) labeled medic plants with both 14C and 15N and showed that 15
- 20% of the 14C was still unmineralised after 4 years but nearly 50% of the 15N remained as
stable organic residues. Granular deposits in soils which stain densely with 0s, U or Pb are
probably “humic” materials rich in polyphenol/protein complexes (Plates 7h, 8g). These may
be remnants of vacuolar polyphenolics but they may also represent secondary products of soil
microorganisms (Tan et al ., 1978), or result from chemical weathering of such materials.
Remnants with similar form, internal structure and staining properties as small as tens of nm
have been seen in ultrathin sections of soils. These are probably to be identified with humic
materials. Early work showed that humic materials formed spheres 60 - 100A in diameter (see
van Dijk, 1971, for review). However Chen and Schnitzer (1976), Ghosh and Schnitzer (1982)
and Stevenson and Schnitzer (1982) have shown experimentally that the precise ultrastructure
of humic compounds e.g., fulvic or humic acid) depends on such environmental factors as pH,
water potential and salt concentrations. Thus, the same material may be deposited as sheets,
bands or fine fibrils depending on pH (Schnitzer and Kodama, 1976) and this is probably true
of similar materials in soils. It might be instructive to fix subsamples of the same soil fabric
with aldehydes buffered to different pH’s and to observe any difference in form of humic
materials. Localized drying, secretion of H + ions by roots, or presence of carbonates may cause
different deposits of the same material within a few microns of each other in the same soil
fabric to have different forms! However it is probable that in nature most humic materials are
complexed with clays.

Unfortunately, most of the humic materials in soils cannot be described in precise chemical
terms, either because the existing techniques are inadequate for their separation and
characterization, or because no two humic molecules are exactly alike (Oades and Ladd, 1977).

Organic matter in soils

623

Design of specific ultracytochemical techniques is therefore impossible.

Membrane systems

It is not unusual to come across membranes in soils. These are 7 - 10 nm thick and are of
variable length. Some are rolled into open or closed tubes and vesicles (Plate 3i (arrows)).
These are probably cytoplasmic membranes released from living cells or when bacteria are
lysed by bacteriophage. Most cellular membranes contain enzymes; it is possible that whole
suites of enzymes, necessary for a sequence of biosynthetic steps which give rise to complex
molecules, are preserved in situ in such membrane fragments, especially if the membranes roll
up or form enclosed vesicles. Such membrane systems may be sites of quite complex
biochemical transformations and could give rise to complex nutrients for soil animals.

Enzymes

Martinez and McLaren (1966) remark that, although hundreds of reports dealing with soil
enzymes have been published, the question of the origin and localization of these enzymes is
still as obscure as it was in the first decade of this century. Burns (1982) has proposed 4 main
sites: (1), in the biomass; (2), inside or adsorbed onto cell wall fragments; (3), adsorbed in or on
clay minerals; and (4), adsorbed onto or as co-polymers with humic materials. There has been a
considerable discussion in the literature as to whether enzyme contents of soils correlate closely
with microbial biomass (Nannipieri et al., 1983) or not. Most classes of enzyme have been
isolated from soils (Skujins, 1976) but their precise location in soil fabrics is unknown;
therefore, the ratio of soil transformations associated with the biomass and with “free” enzymes
remains undetermined.

Both roots (Plate 8a) and many soil bacteria excrete enzymes extracellularly where they are
associated with their ECP’s. (Plates 6b, c, 8b, c). It is also known that enzymes are stabilized
by adsorption onto clay surfaces or by forming complex co-polymers with polyphenols. In
experimentally synthesized enzyme/polyphenol co-polymers, the enzymes retain much of their
catalytic activity and it is likely that similar co-polymers are generated during cellular lysis or
after the release of their components into the soil. Enzymes in these complexes may be
protected from the action of proteases released by nearby soil microorganisms. Non-enzyme
proteins will also be preserved in these sites and may play an important role in animal nutrition.

Although in situ ultracytochemical tests for enzymes in soil fabrics have been successfully
used to locate enzymes associated with microorganisms (Plate 6b, alkaline phosphatase: Plate
8b, acid phosphatase, Plate 6c catalase, Plate 8c peroxidase) and cellular debris (Plate 6h, acid
phosphatase)(Foster, 198 Id, 1982), unfortunately, they have so far failed to locate enzymatic
activity in or on soil minerals. Some mineral fragments appear occasionally to have unusual
electron-dense deposits associated with them after ultracytochemical tests, (Plate 8, b lower
arrow). However, because enzyme histochemistry is generally performed on the bulk soil
sample, it is impossible to perform experimental procedures and control procedures on adjacent
serial sections by present techniques. Moreover since the sections are so thin and the deposits so
small (10 - 100 nm) EPMA could not have been used to determine whether the electron dense
deposits were fragments of electron dense minerals, or enzyme-specific heavy metal
precipitates, so the specificity of these deposits was difficult to establish.

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Foster

Plate 5. Physically protected organic matter - (a), and (b). Clay is adsorbed onto the capsule of microorganisms, (c).
Fragments of wall material (0) become enclosed in clay aggregates and hence protected from microbial decay, (d), and
(e). Humified materials enclosed in a pore within an aggregate. The material appears granular at high magnification (e).
(0, and (g). Amorphous (f)and lamellate (g) organic matter (0) enclosed in clay fabrics, (h). Amorphous organic
materials mixed with clay. (i). Middle lamella fragment enclosed in clay.

Organic matter in soils

625

Plate 6. Acidic carbohydrates (lanthanum reactive) in soils.- (a). General view of a fabric containing amorphous (Ca)
acidic carbohydrates and cell wall fragments, (b). Test for alkaline phosphatese locates the enzyme in microbial capsules,
(c). Catalase is also associated with capsule materials, (d). Detail of (a) showing a small colony of bacteria (B) supported
by amorphous and laellate organic matter (0). (Lanthanum stain), (e). Even after a bacterial colony (B) has died the
capsule materials persist linking other soil components such as humic materials (arrow) together. (0. and (g). Fibrous (0
and amorphous (g) lanthanum reactive organic materials in clay fabrics (arrows), (h). After the acid phosphatase reaction
some organic particles (0) appear to have to have enhanced electron density, (i). Membrane (arrows), possibly remains of
I plant cell walls abound in many soils.

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Plate 7. Neutral carbohydrates (PAMS AND PATSP-reactive) in soils - (a). PAMS stains the cell walls of bacteria (B)
but not their capsule carbohydrates or the root mucilage (RM). (b). At higher magnification the granular product of the
PAMS reagents is associated with linear, presumably wall materials, round objects, perhaps bacteria and clay particles
(arrows), (c). Similarly the PATSP reagents stain the cell wall (W) of bacteria (B) as well as amorphous materials (Ca,
arrow) in the soil. (d). Wall (W) and other organic deposits stained by PAMS, (e), and (0- If clay particles are isolated
from soils and tested with the PAMS reagents only certain particles are stained (arrows), as when sections are stained.
This suggests that only a few clay particles are coated with carbohydrate. (0 is the control which has been treated with
silver methenamine but not with the periodate, (g). PAMS treated “humus” from a compost heap. The material varies
widely in its reaction to the stain; presumably the particles which are less intensely stained are devoid of materials that are
readily periodate-reactive. (B) indicates bacteria, (h). At high magnification Os-treated materials contain granules 25 nm
in diameter. These are probably humic materials.

Organic matter in soils

627

Plate 8. Various histochemical reactions.- (a), and (b). In plant cells (a) acid phosphatase is associated with the
plasmalemma and cell wall. In soils (b) acid phosphatase is generally associated with bacteria, though usually only a few
react. Electron density is sometimes also associated with amorphous materials (lower arrow), (c). In waterlogged soils
peroxidase is associated with narrow tubular microorganisms (arrows) though, again, only a few of the cells are reactive,
(d), and (e). The thick walled hyphae of mycelial strands in soil stain intensely throughout (d) but if the cells arc
pretreated with amylase, only the outer layers of the cell walls still stain. (0. (g). and (h). The final product of organic
matter transformations in soils are amorphous materials. These arc usually mixed in composition since they stain with
PATSP (neutral carbohydrates) (f), osmium alone (humic materials) (g) and ruthenium rcd/osmium (acidic
carbohydrates) (h).

628

Foster

SUMMARY AND CONCLUSIONS

There are three reasons why ultracytochemical studies of soil organics are difficult. First,
the hardness of minerals imposes severe technical restraints; second, electron micrographs are
usually monochrome; and third, organic materials in soils are chemically complex and much
modified from their original structural and biochemical properties.

Dommergues et al. (1977) consider that TEM is useless for quantifying materials in soils
because of the sophisticated and lengthy procedures involved in sample preparation, and
difficulties with respect to microscopic field orientation and size. The great resolution of the
TEM has its premium in the extremely small area examined, so that where quantification may
be attempted, sample variability is the dominant consideration. The role of TEM
ultracytochemistry is, rather, investigation of the structure environment of organic matter at
stages of mineralization which are determined by prior quantitative biochemical or biophysical
studies.

Although image processing can be used to produce colored displays of electron images, e.g.,
distinguish between electron density due to background osmium reaction and that due to an
histochemical reaction on the basis of element distribution (Tanaka and Mitsushima 1984),
most electron micrographs contain only black, white and grey areas. In general only those
products which are electron-dense can be detected, i.e., minerals and materials containing
heavy metals. It is, therefore, much more difficult to design ultracytochemical reactions for
electron microscopy than cytochemical reactions for light microscopy, where colored stains
markedly enhance the visibility of the products of histochemical reactions. Ultrathin sections of
most biological tissues have little or no intrinsic electron density, therefore, the results of
histochemical tests are unequivocal, providing adequate controls are employed in which target
groups are masked or destroyed. These ideal conditions do not hold for ultrathin sections of
soils. Here the minerals are electron-dense to varying degrees; organic matter may adsorb
electron-opaque materials from nearby minerals. Moreover, although histochemcial reagents
give uniquivocal results when tested against relatively pure and well characterised materials in
tissues, the same may not be true of the much modified materials which occur in soils.

Until recently, histochemical reagents were not very specific. The use of antibodies and
lectins labeled with heavy metals, (Knox and Clark, 1978) may prove very useful in the study
of mineralization of organic matter though preliminary experiments showed little sign of
specific staining of rhizospheres (Foster, unpublished). Lectins may almost prove to be too
specific in that they may detect only relatively unmodified materials which can be recognised
anyway from their structure or location.

Ultrahistochemical analysis of soil fabrics is important because it provides information not
easily obtained by other electron optical techniques. Thus ultracytochemistry not only tells us
where organics are located in soil fabrics, but also something of their biochemical properties.
EPMA, SIMS, LAMMA etc. are useful in that they tell us what elements, ions or chemical
groups are present in organic deposits; they do not tell us how these parts are put together to
form an organic complex.

ACKNOWLEDGEMENTS

I thank Ms. Y.K. McEwan and Mr. T.W. Cock for excellent technical assistance in
preparing difficult materials for electron microscopy and for help with the bibliography

Organic matter in soils

629

(Y.K.E.) and plates (T.W.C.).

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Quaest. Ent., 1985, 21 (4)

THE IMPORTANCE OF SOIL FAUNA IN REGULATING SOIL MICROSTRUCTURE
AND SOIL MANAGEMENT IN FORESTS

H.-J. Altemiiller

Institut fur Pflanzenerriahrung und Bodenkunde
Bundesalle 50
D-3300 Braunschweig
Federal Republic of Germany

Quaestiones Entomologicae
21:635 1985

ABSTRACT

The effect of faunal activities on structure formation is most impressive in soils containing a
dark humic horizon of the mull type (mollic epipedon). Especially in the transition zone to a
brighter subsoil it can be shown, in which way aggregates of different origin are locally
deposited, changed, and reincorporated in larger units or coherent areas. Shape and size make
evident that various kinds of soil animals contribute to this cycling process, finally resulting in a
complete contacting and mixing of organic and organic components and forming new mull.

In forest soils often the humic material does not reach the status of dark mull. In such cases
it is often unclear how far the soil fauna is involved in the incorporation of organic substance
into the soil. The study of thin sections by means of an incident light fluorescence-microscope
can indicate that the transport and mixing effect in such soils is often underestimated. Small
organic particles, unvisible with other microscopic techniques, can be observed also in subsoil
areas. Their spatial distribution is not to explain merely by root growing and rotting processes.

In forest management two factors which are related to soil structure are of particular
importance.

The use of heavy machinery leads to increasing problems of soil compaction. Depending
from the soil properties and the climatic conditions hydromorphic features may be formed.
Leaching will cause an instability of the binding forces and the regulating potential of the
fauna is often repressed.

The other factor is acidity. The humus horizons of acid soils onto podzols (spodosols) are
well known. It is obvious that the structure forming activities are strongly reduced. We have to
take in account, that possibly the processes leading to acid soil conditions are much faster today
than supposed before. Meliorative measures will be discussed.

SOIL FAUNA AND AGRICULTURE: PAST FINDINGS AND FUTURE PRIORITIES

Stuart B. Hill
Department of Entomology
Macdonald College of McGill University

Ste-Anne de Bellevue, Quebec H9X ICO Quaestiones Entomologicae

CANADA 21:637-644 1985

ABSTRACT

General findings of soil: soil fauna research are given under the headings of soil pests,
effects of beneficial soil animals, and effects of agricultural practices. Arguments are
presented for a sustainable agriculture and for a more rational approach to problem solving
within agroecosystems. The use of indicators of agroecosystem distress is advocated.
Comments are included on research needs and implementation of sustainable systems of soil
management.

RESUME

Les decouvertes generates de la recherche sur les sols et leur faune sont presentees sous les en-tetes d’organismes
nuisibles, d’effets des animaux benefiques aux sols, et d'effets des pratiques agricoles. L’auteur offre des arguments en
faveur d’une agriculture soutenable et d’une approche plus rationnelle pour resoudre les problemes dont souffrent les
agro-ecosystemes. II preconise I’utilisation d’indicateurs de stress dans les agro-ecosystemes et commente sur les besoins
en recherche et sur la mise en oeuvre de systemes soutenables d’amenagement des sols.

INTRODUCTION

Agriculture is defined as the science or practice of cultivating the soil and rearing animals,
and cultivation as the preparation, tillage and use of soil to produce crops. Because definitions
aim to clarify and simplify meaning, they often perpetuate destructive myths that are harder to
. change than the definitions that incorporate them. The above definitions, for example, paint a
picture of a linear agriculture with a black box, the soil, in the middle. The farmer stirs up the
soil with a tool, sows seeds and harvests the plants that mysteriously grow. At first the system
was thought to be limited to people, land, seeds and tools. More recently, synthesized fertilizers
and pesticides have been added to the equation. These and other developments have led to an
agriculture that is characterized by large parcels of land being kept bare for most of the year,
often only one crop species being grown year after year, and production being maintained
through a heavy reliance on imported seeds, energy as fuel for equipment, fertilizers and
( pesticides. The outcome has been increased dependence, environmental stress and a loss of
| capital from the system in terms of crop cultivars, soil and nutrients, water, natural controls of
pests, and other beneficial organisms.

Studies of the relationships between soil fauna and agricultureOhave been conducted within
isuch systems. They comprise three types of studies: (1), of pest species and their control; (2), of
! beneficial species and their effects; and (3), of the effects of agricultural practices on soil
animals. Because of the difficulties of studying organisms in a stratified opaque medium that is
.complex in terms of its physical, chemical and biological parameters, and that varies in time
and space, progress in all of these areas have been limited. Some general statements, however.

638

Hill

can be made. Useful reviews are provided by Kevan (1962), Edwards and Lofty (1969), Mills
and Alley (1973), Wallwork (1976), and in the Proceedings of the Colloquium edited by Dindal
(1980). As most of the following statements are of a general nature or are based on personal
observation and/or on the soil fauna literature in general, they are not supported by specific
references. These are, however, given where useful reviews or landmark papers are known to
exist, or where particular points need to be stressed.

SOIL PESTS

1. Soil pests are at least as significant, in terms of economic damage, as above ground pests.

2. Pesticides, because of difficulties of distribution in soil, adsorption and decomposition,
have provided less effective control in soil than above soil.

3. The biology and ecology of most soil pests is inadequately understood and relatively few
biological controls have been exploited.

4. The use of cultural methods (crop rotation, use of intercrops, timing of operations, soil
and habitat management) and resistant crop varieties, if available, are essential for the control
of soil pests.

EFFECTS OF BENEFICIAL SOIL ANIMALS

1 . All soil animals have beneficial effects on soil structure and fertility.

2. Although their direct effects on processes such as soil formation and organic matter
decomposition are small in comparison with those of microorganisms, their indirect and
catalytic effects are substantial and essential. These include the improvement of food and space
conditions for microorganisms and higher plants, selective cropping and transportation of
microorganisms, aeration, drainage, biological control of pests and soil mixing. Generally, their
role should be seen as one of “regulation” rather than the simple acceleration of soil processes,
which is a common misconception. (Macfadyen, 1961, 1963; Weetman et al., 1972; Hill et al. ,
1973; Behan and Hill, 1978; Lee, 1979; Anderson et al., 1981; Luxton, 1982; Parkinson, 1983;
Seastedt and Crossley, 1984).

3. Most studies of the contribution of soil animals have failed to deal with the soil system as
a functional whole. Rather, they have focused on isolated groups and processes. Consequently
our views of how the soil works is still very fragmentary.

4. Few attempts have been made to introduce and manage beneficial soil fauna (Edwards
1981). Developments in this area will eventually lead, together with parallel developments in
other areas, to the redesign of our food producing systems and to changes in our approach to
soil management.

EFFECTS OF AGRICULTURAL PRACTICES

1 . Dominant agricultural practices (tillage, clean cultivation, monoculture, row crops, use of
pesticides and certain synthetic fertilizers) simplify the soil community and reduce the
beneficial contribution of soil animals (Edwards and Lofty, 1969; Edwards and Thompson,
1973; Andren and Steen, 1978; Edwards, 1983). Manures and most fertilizers generally
increase numbers and species of soil animals (Marshall 1977).

Soil fauna and agriculture

639

2. Although numerous studies have been carried out on these effects they have not led to any
changes in agricultural practices - the beneficial soil fauna remains a largely unknown and
untapped resource within the food system.

3. Preliminary studies have indicated the value of using the presence and population density
of certain soil animals as indicators of soil conditions (Karg, 1968).

4. The growing concern with soil degradation and interest in miminum tillage and ecological
approaches to agriculture are causing some attention to be focussed on the soil fauna (Stinner
and Crossley, 1983). The questions that are being raised provide soil ecologists with an
important opportunity to make practical contributions to the design of sustainable food
systems.

SUSTAINABLE AGRICULTURE

As responsible scientists we have an important role to play in the evolution of a sustainable
lifestyle for our species. Because of its increasing dependence on distant non-renewable and
renewable resources, and its heavy environmental impact, modern agriculture is clearly not
sustainable.

Systems of agriculture that have increased “productivity” (to satisfy markets manipulated
through advertising), “profit” and “power” as their primary goals, are not sustainable and lead
to the degradation of person and planet. This is because these goals know no limits. They are
exhausting of resources and unresponsive to their harmful side-effects. What I am arguing for
is a greater social conscience among scientists and a translation of that conscience into research
that is relevant to food systems that have goals such as nourishment, fulfillment, flexibility, and
sustainability (Hill, 1982; Hill and Ott, 1982; Hill, 1984a). I am also arguing for soil biologists
and ecologists to speak out on these issues, and to broaden their area of interest to include the
food system as a whole and its sustained operation over the long-term.

Let us now consider what a sustainable food system might entail and what contributions soil
biologists can make towards its development and implementation.

In terms of material flows, a sustainable agriculture may be viewed as a production -
consumption - recycle system. Most of the recycle process takes place within the soil in the form
of organic matter decomposition. For sustainability to be achieved, inputs for decomposition
must meet certain quantitative and qualitative criteria, e.g., comprise a diverse range of
substrates containing adequate amounts of major, minor and trace elements that, together with
those from the earth’s crust and the atmosphere, are capable of supporting plant growth.
Substrates must also meet certain time, space and freedom from toxins, criteria. These criteria
are more likely to be met in a multi-story polyculture that includes soil and ecosystem
maintaining, as well as food producing, plants and animals, than in a uni-story row-crop
monoculture (Mollison, 1979; Altieri, 1983; Todd and Todd, 1984). The agricultural task is the
design and management of such systems and the soil zoology task is to describe the animals and
processes that take place in the soil and, with others, to develop methods of soil management
that can enhance the beneficial contributions of the soil fauna.

PROBLEM SOLVING WITHIN SUSTAINABLE AGROECOSYSTEMS

The approach to problems in such systems will probably differ radically from that employed
today. Currently, agricultural problems usually receive attention only when their short-term

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economic consequences justify the required expenditures. Solutions tend to be confined to
disciplines rather than multidisciplinary: entomologists dealing with insects, nematologists with
nematodes, and so on.

An alternative approach, recognizing that the causes of problems often lie outside of the
discipline concerned with their subject, and that prevention is usually less costly than cure,
might channel the efforts and resources that currently are used to directly attack problems to a
less easily defined, maintenance function for multifacetted agroecosystems. Thus, by working
to optimize the functioning of the agroecosystem as a whole, problems within its parts would be
minimized. Those that arise would be taken as indicators of malfunction, and efforts would be
made to correct the malfunction. To be effective with this approach farmers would need to be
more knowledgable and, “closer” to the agroecosystem, and supported more by society.
Sociologically the process may be viewed as one of integration (of our species into the rest of
the biosphere), balance (the maintenance of sustainable relationship with the support
environment) and feedback (paying close attention to the outcomes of our actions, recognizing
their meaning and responding accordingly). Thus, attention is shifted from problem solving to
system maintenance, the incidence of problems declining as systems approach optimal states.
Problems that do arise are solved largely by removing the causes and strengthening the natural
processes that normally prevent such problems from reaching crisis proportions (Hill, 1984b).

INDICATORS OF AGROECOSYSTEM DISTRESS

Recognition of undesirable processes often involves the identification of environmental
stressors and the detection and measurement of their effects.

Because of the widespread and diverse nature of environmental stressors, and because of the
complex nature of their interactions, there is a need to find ways to detect and measure their
combined effects in a general way. Influenced by Selye’s (1946) recognition of a “biological
distress syndrome” in mammals, Rapport (1983) has proposed that we recognize a parallel
“ecosystem distress syndrome” within environments. This concept is based on two important
assumptions: (1), that different stressors give rise to certain similar symptoms (cf. Selye’s
“general adaptation syndrome”); and (2), that there are common indicators of distress that can
be used in widely different ecosystems subject to different stressors.

The situation in mammals, however, is much more complicated than Selye has indicated.
Randolph (1976), uses five levels to describe recognizable points along a continuum from
healthy to severe illness within affected humans. One valuable insight from his observations is
that at different times the symptoms present themselves in “up” ( e.g hyperactive) and “down”
( e.g ., depressed) states. While these are both recognized as being undesirable at the developed
end of the spectrum, during the early stages of development the “up” condition (active,
responsive, enthusiastic, ambitious, witty) may easily be regarded as desirable, its connection
with the “down” condition (stuffy nose, occasional coughing and sneezing, skin disorders, gas,
diarrhea, constipation, frequent urination and various eye and ear symptoms) not being
recognized.

There may well be parallels to these observations with respect to the soil ecosystem (Hill,
1980). Thus, certain management practices may at first appear to be beneficial when measured
in terms of their short-term influences on productivity. The negative effects of these practices
are either hidden or not taken seriously until they reach crisis proportions, when it maybe too
late to correct the situation.

Soil fauna and agriculture

641

The following indicators of environmental distress, identified by Rapport (1983) for the
Great Lakes Ecosystem, are equally applicable to soil ecosystems:

1 . Imbalance in nutrient concentrations (loss of some, accumulation of others)

2. Reduced species diversity

3. Replacement of longer lived by shorter lived species (adapted to transitory novel
environments)

4. Replacement of larger by smaller life forms

5. Decline in biomass of macrofauna

6. Increase in amplitude of population fluctuations of key species.

Some of these were recently recognized by Andren and LagerTf (1983) in their study of the
effects of various agricultural practices on soil mesofauna.

One problem with these indicators is that they only provide an after-the-fact indication of
distress. This limitation similarly applies to many specific indicators of environmental
contamination, such as the accumulation of toxins up the food chain, and the incidence of
reproductive failure among top predators (Rapport, 1983).

In addition to these indicators, we urgently need others that are able to provide us with an
early warning of deteriorating conditions. For this, Rapport (1983) has proposed that we
identify “indicator-integrator” organisms, species that are representative of their communities,
are able to survive only in relatively unstressed ecosystems, and that are sensitive to a broad
range of stressors.

Among soil invertebrates, predators within the air spaces and water film and highly mobile
burrowers would seem likely candidates for this role. Karg (1968) has, long ago, stressed the
value of using predatory soil mites as indicators, and Greenslade and Greenslade (1983) make a
similar case for using ants. Predatory nematodes would probably serve a similar function within
the water film. In fact, all soil animals are indicators of soil conditions. The problem is the
interpretation of the information provided. Predators are particularly valued because their
presence, population density, behaviour and body composition can provide, in a sense, a
summation of most of the information provided separately by the organisms lower down in the
food web. Among the non-predators, earthworms are already widely regarded by farmers as
indicators of soil health, and have been successfully used as indicators of soil pollution by
pesticides and industrial chemicals (Edwards, 1979, 1980). Ghilarov (1965) and Krivolutsky
(1975) have proposed using soil fauna as indicators of soil type. The person with the greatest
need for this “indicator information” is the farmer, and researchers should keep this in mind.

While it is essential that more work be done in this area, experience from other fields is not
encouraging with respect to the ability of such studies, on their own, to bring about appropriate
changes in agricultural practices. While most human populations are willing to support studies
of the side-effects of their behaviour, it is rare to Find changes in behaviours as a result of such
studies. I have observed that most people only want to hear truths that validate their present
i lifestyles, that do not cause them to feel guilt, and that do not suggest that they should change
their behaviour. It is often implied that, as scientists, we are more objective and more willing to
be open to truths that disturb, but this has not been my observation. I believe that most of us
conduct our science (and our lives), just as non-scientists conduct their lives, within a territory
determined by our vulnerability to the truths that are likely to distress us. This implies that by
increasing our vulnerability we are likely to improve our science. This involves opening-up more
to our colleagues, to those in other disciplines, to non-scientists and, in a somewhat different
sense, to the subjects of our research. The fact that this meeting has taken place, bringing

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together soil micromorphologists and soil zoologists from around the world, is a positive step in
this direction.

REFERENCES

Altieri, M.A. 1983. Agroecology: The Scientific Basis of Alternative Agriculture. 173 pp. Divn.
Biol. Control, Univ. Calif., Berkeley, CA.

Anderson, R.V., D.C. Coleman and C.V. Cole. 1981. Effects of saprotrophic grazing on net
mineralization, pp. 201-217. In: F.E. Clark and T. Rosswall (Editors). Terrestrial Nitrogen
Cycles. Ecol. Bull. (Stockholm) 33.

Andren, O. and J. Lagerlof. 1983. Soil fauna (microarthropods, enchytraeids, nematodes) in
Swedish agricultural cropping systems. Acta Agr. Scand. 33: 33-52.

Andren, O. and E. Steen. 1978. Effects of pesticides on soil organisma. 1. Soil fauna (In
Swedish, with English summary) SNV Stockholm. PM 1082: 95 pp.

Behan, V.M. and S.B. Hill. 1978. Feeding habits and spore dispersal of oribatid mites in the
North American Arctic. Rev. Ecol. Biol. Sol. 15(4): 497-516.

Blumberg, A.Y. and D.A. Crossley, Jr. 1983. Comparison of soil surface arthropod populations
in conventional tillage, no-tillage and old field systems. Agro-Ecosystems 8: 247-253.

Dindal, D.L. (Editor). 1980. Soil Biology as Related to Land Use Practices. Proc. 7th Int. Soil
Zool. Colloq., Syracuse, N.Y. EPA-560/ 13-80-038. EPA, Wash.

Edwards, C.A. 1979. Tests to assess the effects of pesticides on beneficial soil organisms, pp.
249-253. In: Tests for the Ecological Effects of Chemicals. Pub. Erich. Schmidt, Verlag,
Berlin.

Edwards, C.A. 1980. Interactions between agricultural practice and earthworms, pp. 3-12. In:
D.L. Dindal (Editor). Soil Biology as Related to Land Use Practices. Proc. 7th Int. Soil
Zool. Colloq., Syracuse, N.Y. EPA-560/ 13-80-038. EPA, Wash.

Edwards, C.A. 1981. Earthworms, soil fertility and plant growth, pp. 61-85. In: A. A. Appelhof
(Compiler). Workshop on the Role of Earthworms in the Stabilization of Organic Residues.
Proceedings, Vol. 1. Beech Leaf Pr., Kalamazoo, MI.

Edwards, C.A. and J.R. Lofty. 1969. The influence of agricultural practice on soil
micro-arthropod populations, pp. 237-247. In: J.G. Sheals (Editor). The Soil Ecosystem.
Symp. Publ. 8. Syst. Assoc., London.

Edwards, C.A. and A.R. Thompson. 1973. Pesticides and the soil fauna. Residue Rev. 45:
1-79.

Ghilarov, M.S. 1965. Zoological Methods of Soil Diagnosis. (In Russian, English summary).
278 pp.

Greenslade, P.J.M. and P. Greenslade. 1983. Ecology of soil invertebrates, pp. 645-669. In:
Soils, An Australian Viewpoint. Division of Soils, CSIRO, Melbourne, Academic Pr.,
London.

Hill, S.B. 1980. Observing stressed and unstressed ecosystems and human systems: means for
recovery and value identification, pp. 1 121-1 138. In: Absolute Value and the Search for the
Peace of Mankind. Vol. 2. Int. Cultur. Fdn. Pr., N.Y.

Hill, S.B. 1982. A global food and agriculture policy for western counties: laying the
foundations. Nutr. Health 1(2): 107-117.

Hill, S.B. 1984a. Controlling pests ecologically. Soil Assoc. Quart. Rev., March: 13-15.

Hill, S.B. 1984b. (in press) Implementing a sustainable food system. 14 pp. manuscript. Proc.

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Rights to Food Conference, Concordia Univ., Montreal.

Hill, S.B. and P. Ott (Editors). 1982. Basic Technics in Ecological Farming. 365 pp.
Birkhausser, Basel, Switzerland.

Hill, S.B., L.J. Metz and M.H. Farrier. 1973. Soil mesofauna and silvicultural practices, pp.
119-135. In: Bernier, B. and C.H. Winget (Editors). Forest Soils and Forest Land
Management. 675 pp. Pr. Univ. Laval, Que.

Karg, W. 1968. Bodenbiologische Untersuchungen iiber die Eignung von Milben, insbesondere
von parasitiformen Raubmilben, als Indicatoren. Pedobiologia 8: 30-39.

Kevan, D.K.McE. 1962. Soil Animals. 237 pp. Witherby, London.

Krivolutsky, D.A. 1975. Oribatoid mite complexes as the soil type bioindicator, pp. 217-221.
In: J. Vanek (Editor). Progress in Soil Zoology. 630 pp. Academia, Prague.

Lee, K.E. 1979. The role of invertebrates in nutrient cycling and energy flow in grasslands, pp.
26-29. In: Crosby, T.K. and R.P. Pottinger (Editors). Proc. 2nd Austr. Conf. on Grassland
Invertebrate Ecology. Govt. Print., Wellington.

Luxton, M. 1982. General ecological influence of the soil fauna on decomposition and nutrient
circulation. Oikos 39(3): 355-357.

Macfadyen, A. 1962. The contribution of the microfauna to total soil metabolism, pp. 3-17 In:
van der Drift, J. and J. Doesken (Editors). Soil Organisms. N. Holland Publ., Amsterdam.

Marshall, V.G. 1977. Effects of manures and fertilizers on soil fauna: a review. Commonwealth
Bureau of Soils. Spec. Publ. 3: 79 pp. CAB, Farnham Royal, U.K.

Mills, J.T. and B.P. Alley. 1973. Interactions between biotic components in soils and their
modification by management practices in Canada: review. Can. J. PI. Sci. 53: 425-441.

Mollison, B. 1979. Permaculture Two. 150 pp. Tagari, Stanley, Tasmania.

Parkinson, D. 1983. Functional relationships between soil organisms, pp. 153-165. In: Lebrun,
P., H.M. Andre, A. De Medts, C. Gregoire-Wibo and G. Wauthy (Editors). New Trends in
Soil Biology. 709 pp. Proc. 8th Int. Colloq. Soil Zool. Louvain-La-Neuve, Belgium.

Randolph, T.G. 1976. Adaptation to specific environmental exposures enhanced by individual
susceptability, pp. 46-66. In: Dickey, L.D. (Editor). Clinical Ecology. Charles C. Thomas,
Springfield, 111.

Rapport, R.J. 1983. Indicators of water quality from an ecosystem perspective. 12 pp.
manuscript for “Informal Meeting on Water Use and Quality Statistics” at Conference of
European Statisticians, Geneva, 12-14 Dec. Statistics Canada, Ottawa, Ont.

Seastedt, T.R. and D.A. Crossley, Jr. 1984. The influence of arthropods on ecosystems.
Bio-Science 34(3): 157-161.

Selye, J. 1946. The general adaptation syndrome and the diseases of adaptation. J. Allergy 17:
231-47,289-323,358-398.

Stinner, B.R. and D.A. Crossley, Jr. 1980. Comparison of mineral element cycling under till
and no-till practices: an experimental approach to agroecosystem analysis, pp. 280-288. In:
Dindal, D.L. (Editor). Soil Biology as Related to Land Use Practices. Proc. 7th Int. Soil
Zool. Coloq., Syracuse, N.Y. EPA-560/1 3-80-038, EPA. Wash., D.C.

Stinner, B.R. and D.A. Crossley, Jr. 1983. Nematodes in no-tillage agroecosystems, pp. 14-28.
In: Freckman, D. (Editor). Nematodes in Ecosystems. Univ. Texas Pr., Austin, TX.

Todd, N.J. and J. Todd. 1984. Bioshelters, Ocean Arks, City Farming: Ecology as the Basis of
Design. 210 pp. Sierra Club Books, San Francisco.

Wallwork, J.A. 1976. The Distribution and Diversity of Soil Fauna. 355 pp. Academic Pr.,
London.

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Weetman, G.R., R. Knowles, and S.B. Hill. 1972. Effects of different forms of nitrogen
fertilizer on nutrient uptake by Black spruce and its humus and humus mesofauna.
P.P.R.I.C. Woodl. Rep. 19: 1-20.

SOIL FAUNA AND SOIL STRUCTURE: FEEDBACK BETWEEN SIZE AND

ARCHITECTURE

W.B. McGill

Chairman, Department of Soil Science
University of Alberta
Edmonton, Alberta T6G 2E3
CANADA

J.R. Spence

Department of Entomology
University of Alberta

Edmonton, Alberta T6G 2E3 Quaestiones Entomologicae

CANADA 21:645-654 1 985

ABSTRACT

The relations between soil fauna and soil structure are examined using papers from this
conference as a background. Our synthesis focuses on function of the soil system and
reciprocity between soil animals and other soil components.

Advancement of knowledge at this interface has been impeded by disciplinary
specialization and isolation, and failure to frame hypotheses and research strategies in the
context of the entire soil system. Two major challenges must be met before progress will be
possible. First, philosophical beliefs about soil must be separated from objective science. The
second problem is mainly taxonomic. For soil animals, problems of correlating phylogenetic
and ecological groupings must be resolved. For soil micromorphology, classifications must be
simplified and made more accessible to soil ecologists.

We conclude that soil animals regulate soil function through both trophic interactions and
biophysical mechanisms which influence microhabitat architecture. The mixed culture aspect
of soil communities involves diverse species interactions which regulate the structure of soil
communities. We propose that comminution and disintegration of microstructures be added to
formation of microstructures and comminution of plant debris as a third biophysical
regulatory mechanism. This leads to a dynamic view of micropedology. Establishing links
between groups of soil organisms and specific soil microstructures as seen in thin section will
require substantial collaborative effort. Such efforts will yield basic information necessary for
solving pressing applied problems in management of renewable resources depending upon soil.

RESUME

Nous synthetisons les rapports entre la faune edaphique et la structure des sols to la lumi'ere des articles presentes au
cours de la conference. Cette synthase se concentre sur les fonctions des sols en tant que systimes et sur la reciprocity des
rapports entre les animaux edaphiques et les autres composantes du sol.

Le progres des connaissances ti ce niveau a ete entrave par la specialisation et I'isolement des diver ses disciplines, et
par le manquement h formuler des hypotheses et des strategies de recherche qui considtrent les systtmes edaphiques
dans leur ensemble. Deux defis de taille doivent etre confrontes si I on est pour progresser. D'abord il faut separer les
convictions philosophiques au sujet du sol de I'approche scientifique objective. Deuxitmement. il faut surmonter les
problemes taxonomiques. En ce qui concerne la faune edaphique. il faut reussir d correler les groupes phylogenetiques
avec les groupes ecologiques. En ce qui concerne la micromorphologie des sols, il est necessaire de simplifier les

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McGill and Spence

classifications et de les rendre plus accessibles aux ecologistes etudiant les sols.

Nous concluons que les animaux edaphiques regularised la fonction du sol par des interactions entre les niveaux
trophiques et par des mecanismes biophysiques qui affectent I’architecture des microhabitats. L’apparence de culture
melangee que presented les communites edaphiques met en jeu des interactions diverses entre les especes qui regularised
la structure de ces communautes. Nous proposons que la pulverisation des debris et la disintegration des microstructures
soient considerees comme formant un troisieme mecanisme regulatoire biophysique en plus de ceux de la formation des
microstructures et de la pulverisation des debris vegetaux. De cette facon on obtient une image dynamique de la
micropedologie. L’etablissement de liens entre les groupes d'organismes edaphiques et les microstructures specifiques des
sols requierera des efforts de collaboration substantiels. De tels efforts permettront d'obtenir des informations
fondamentales necessaires pour resoudre les problemes pratiques d’amenagement des ressources renouvelaoles qui
dependent du sol.

INTRODUCTION

As the circle of knowledge increases, so too does the fringe of ignorance. An objective of this
conference was to increase knowledge without expanding the fringe of ignorance by combining
results of analyses from two spheres: soil micromorphology and soil zoology. The mathematical
proof of the above possibility is simple, but the challenge of bringing about constructive
interaction between soil micromorphologists and soil zoologists is not.

Since the pioneering work of Kubiena (1938) we have known that soil structure and
function are intimately related. In this conference, papers by Hill and Parkinson showed that
soil animals regulate other soil biota both directly and by altering their environment.
Altemiiller, Mermut, Pawluk and Rusek showed convincingly that soil animals play a large role
in organizing and maintaining soil fabrics.

Increased understanding of relations between soil fauna and soil structure will have
important practical benefits. For example, Hill remarked that sustained agriculture depends on
understanding the regulation of complex biological processes occurring in soil rather than
indiscriminately accelerating a few. Several authors repeated the theme that soil animals
contribute to soil quality and modify soil profiles and nutrient supply to agricultural crops. In
particular, the paper by Edwards summarizes information now available about the importance
of earthworms, a topic that was first studied experimentally by Charles Darwin (1881). Papers
by Greenslade, Mermut, Pawluk and Rusek showed that animals generate structural units in
soils from the Arctic through temperate regions to the tropics.

Despite immense opportunities for both basic and applied research, soil ecology has
remained a relatively unstudied discipline. The generality of much ecological theory, developed
from studies of freshwater and terrestrial systems, could be tested by work with soil systems.
Also, working out the relationships among biotic and abiotic components of the soil can provide
interesting proximate frameworks for research. Mechanistic questions about relationships
between soil fauna and soil structure have been raised by most speakers. For example, both
Dindal and Norton pointed out the apparent paradox of persistence of faecal pellets associated
with increased rates of decomposition in the presence of soil animals. An important question,
raised by Foster’s presentation, is the extent to which soil animals are involved in disintegration
of fundamental soil structural units. Resolution of such questions will increase understanding of
the important but poorly understood decomposer food web.

In this paper we review some of the past impediments to interaction between soil zoologists
and soil micromorphologists, develop the concept of the soil system as the unifying link between
their disciplines, and present some ideas flowing from such a conceptual approach to studying
relationships among soil animals and soil structure.

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647

IMPEDIMENTS

In North America, soil morphologists and soil zoologists have not communicated in the
recent past, in part due to a tradition of geological affinity of the former group and the
predominant zoological background of those interested in soil animals. For both groups, the
focus of attention frequently was not the soil but some small portion of it. It was therefore
logical to communicate with those having similar interests. A shift of focus to the soil system
would underscore the important point that soil zoologists and pedologists are working on the
opposite side of the same coin. Effective soil ecology will depend upon increased cooperation
between workers in these two areas.

It is true that the animal and its phylogeny or the organic-mineral complexes and their
fabrics are important analytical frameworks in the respective spheres of soil biology and
pedology. However, we suggest that while such perspectives facilitate analysis of parts of the
soil system, exclusive commitment to these points of view has prevented synthesis. In the
broader view, analysis without synthesis is a scientific dead end. Hoffman’s comment that
“myriapods are not just objects to be classified nor are they simply objects to produce faecal
pellets” is appropriate.

Until recently, pedologists and soil zoologists have been necessarily preoccupied with
description of immense natural diversity. The size of various groups of organisms, and the
diversity of soils and fabrics has inevitably promoted disciplinary specialization. Unfortunately,
it appears that with overemphasis on analysis, proximate goals of such specialization have
become ends in themselves. We do not hold that further analytical work is either undesirable or
unimportant. However, we are convinced that a general framework for synthetic work is
available and that we can now proceed without waiting for more perfect descriptions of all
components of the soil system. In fact, it is likely that descriptions will be improved by
experimental studies of interactions among components and by information about emergent
system properties that is generated through synthesis.

From information now at hand, some immediate requirements are obvious. Rusek pointed
out the need to distinguish ecological groups of soil animals. This requires recognition of the
reciprocity between soil animals and other soil components, and realization that soil animals
are part of soil, not mere inhabitants of it. The idea is not new. In his review of the history of
soil zoology, Kevan remarked that in 1757 Adamson recorded the reciprocity between termites
and soil.

Real progress in science is probably often hampered by disciplinary boundaries which have
been created mostly for the convenience of administrators. The willingness of scientists to
adhere strictly to narrow administrative limits appears to be a recent development, even among
workers interested in the soil. For example, Hoffman reported good work was done in the 19th
century by people sharing their efforts among myriapods, echinoderms and mammals. A
growing awareness of the reciprocity between soil animals and other soil components led to this
conference and is reflected in a remark by Parkinson in his presentation: “Kubiena was
remarkably perceptive both as a soil biologist and soil scientist - I suppose they are
synonymous.” Recognition of that unity is growing and is the central thesis of this summary
and synthesis.

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SOIL SCIENCE

SOIL

ZOOLOGY

many species
and

interesting lifestyles

SYSTEM FUNCTION

many structures
and

interesting materials

SOIL

MICROMORPHOLOGY

Fig. 1 . Disciplinary interests showing overlap of soil micromorphology with soil zoology and the concept that the study of
neither is complete without the other.

UNIFYING LINK

Systems consist of several components interacting with each other, and controlled by their
environment. They are characterized by many cause-effect pathways and feedback processes,
which give individuality to each system. Knowledge of that individuality is essential to structure
man’s interaction with ecological systems in a way that permits use of renewable natural
resources that is stable in the long run. With respect to soil, it is clear that soils are being lost
and degraded worldwide much faster than they are being generated and restored (Wolf, 1985).

As pointed out above, the unifying link between soil zoologists and pedologists which
permits advancement of knowledge must be at a broader level of resolution than that required
by either area of study alone. We argue that relationships between system function and system
architecture provide that focus (Fig. 1).

For effective synthesis each part of the soil system merits detailed study and analysis in its
own right. However, there are problems in each area which require information about the
other. For example, while it is generally held that soil animals generate soil microstructures, it
is not often clear which animals are responsible for a specific fabric or structure observed in
thin sections of soil. In fact the relative impacts of soil organisms and abiotic processes are not
well enough known to formulate general hypotheses. Similarly, habitable space and accessible
substrates for various groups of soil animals cannot be evaluated without knowledge of soil pore
size distribution and geometry relative to soil animal sizes and water film thicknesses needed to
permit movement. Predator-prey interactions in soil are also controlled by pore size and
geometry relative to organism sizes. Elliott et al. (1980) presented data consistent with the
hypothesis that soil texture influences habitable pore space and hence trophic interactions in

Soil Fauna and Soil Structure

649

terrestrial ecosystems. The above examples show how system function and architecture unite
the two disciplines. The advancement of knowledge and practical benefits mentioned earlier are
to be attained at this more holistic level.

CHALLENGES

Two challenges must be dealt with before progress may be made. The first is philosophical.
Kevan illustrated how past concepts of soil animals have been shrouded in mythology. Ancient
bestiaries portrayed themes of morality. Also, concepts of soils have varied from the mother of
all life, to masses of ground rock, depending upon perspectives of the writer (Simonson, 1968).
Soils have been associated with immortality and this has been passed to animals associated with
them. Hill pointed out that the above metaphysical themes can be frequently found in
discussions about man’s use of soils or his interactions with it (see also Hyams, 1976).

Such a theme has important cultural consequences which are amenable to investigation
within classics, anthropology, and sociology. However, it may lead to two different outcomes
regarding objective examination of soils and soil animals. On one hand, it may generate a set of
beliefs pertaining to function of soil systems and man’s interaction with them which are not
amenable to scientific scrutiny because they have not been derived from objective data. It may
thereby hinder objective scientific examination of biophysical and biochemical interrelations
between soil animals and the structure or function of the soils of which they are a part. On the
other hand, stressing that roots of agricultural man extend from the soil can lead to a
determined curiosity about how the system functions and how man can appropriately interact
with and even become part of it. The challenge is to assure such objective analysis and
synthesis.

The second challenge is mainly taxonomic. Soil animals are among the most abundant
multicelled animals anywhere on earth (up to 106/ni2) and their rates of reproduction and
turnover can be startling. As pointed out by many authors in this proceedings, identification
and classification of soil animals is both time consuming and difficult because of their small
size, great diversity and relative obscurity among other members of the animal kingdom. For
example, Greenslade estimated that 130,000 species of beetles in 1 1 families occur in soil. As
documented by Fjellberg, Hoffman and Norton, the situation with respect to other groups of
important soil arthropods is equally challenging and much more poorly known. However, few
workers are engaged in soil animal taxonomy and, as Hoffman lamented, there is not much
support for basic taxonomic work. Because research support is society’s way of establishing
value and prestige of workers, few young scholars are being attracted to these vital tasks (see
also Crowson, 1970). As groups of animals are made accessible through production of
| taxonomic monographs, links between species and their environment or interactions within the
I system can be better explored. Edwards’ presentation dealing with the effects of earthworms on
I soil structure and function illustrated what sort of advances are possible through experiments
once a taxon is adequately known for ecological work. However, even with respect to
composition of earthworm assemblages, we are relatively uninformed in North America.
Similarly, soils contain innumerable fabrics with few researchers involved in their
classification.

A proposal by Greenslade may partially resolve the zoological dilemma in the short run. He
, suggests that taxonomists be encouraged to reverse their usual procedures and start analysis by
j separating large groups of important soil animals into genera and species groups. Details of

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species level classification can be worked out after a fauna is packaged for understanding by
non-taxonomists. A first step in this important process in now underway. Dan Dindal is editing
a general guide to soil zoology for North America which has been scheduled for publication by
Wiley. Such treatments will be invaluable to soil biologists and should stimulate ecological
work.

As noted by the Biological Survey of Canada (1982), a major impediment to development of
soil ecology is a lack of taxonomic monographs and keys which are accessible to the
non-specialist. Production of such material should receive high priority. As pointed out by
Hoffman, the production of such basic descriptive taxonomic and faunistic work is often looked
upon with disdain, even though it is most important for stimulating ecological work in the short
run. Both Fjellberg and Rusek recognized need to distinguish ecological groups among taxa
important in soils. Norton pointed out that study of phylogenetic relations is a major stimulus
for classification and that such work has important benefits for synthetic studies. We do not
argue that this approach should be abandoned. However, we submit that ecological
interrelations can provide an alternative stimulus with different but complementary
approaches.

Similarly, complexity of micromorphological classification of soils must be reduced and
useful descriptions of microscale heterogeneity should be made available to non-specialists. The
workshop session organized by McKeague and Fox provides direction for this effort. Again,
synthetic work is appropriately focused by attention to the entire soil system (Fig. 2).
Ultimately, this sort of work will be accomplished best by a new breed of scholar. We hope that
the needs identified by this conference will be addressed by more flexible training of graduate
students in soil ecology in the context of blended research programs that cross traditional
departmental boundaries.

Studies of nutrient or energy flow through the soil system may be taken as an example of the
above approach. Understanding energy flow requires, among other things, knowledge of where
substrates are, where organisms are, and where they can go. A large proportion (40-80%) of
soil pore space and surface area is inaccessible even to organisms of /um size (McGill, in
preparation). Information is therefore required on physical and biological agents which
reorganize soil fabrics to redistribute substrates and organisms. Such needs also link
micromorphology, soil zoology and soil microbiology. The morphologist provides information on
architecture, habitable spaces, and locations of substrates while soil biologists examine feeding
habits and metabolism of various groups of organisms, their abilities to reorganize or produce
specific fabrics, and to ingest mineral or organic material or both.

This conference has underscored the major advantages of joining the disciplines of soil
zoology and pedology to foster growth of knowledge and understanding. Continued detailed
analyses of each component are essential, but interactions among other components of the
system can be an appropriate synthetic focus for study. We argue that the link between soil
morphology and soil biology might best be described as soil biophysics. Thus, it includes but
transcends faecal pellets.

SOME IDEAS

Microhabitats and Microcommunities

Although soils are viewed classically over the landscape at a macro scale of km2 or m2 many
significant processes and mechanisms controlling them occur at a micro scale. Dindal showed
that many distinct microenvironments exist in soil which lead to formation of distinct

Soil Fauna and Soil Structure

651

Fig. 2. Use of the soil system as a central focus for research. Work in the many subdisciplines of soil zoology and pedology
can be synthesized in the dynamic framework of the soil system. The diagram emphasizes that spin-offs from synthesis will
contribute to analysis in each subdiscipline. Spin-offs will also contribute to general theory and find applications in
agriculture and forestry.

microcommunities and add to the spatial complexity of the macroenvironment. The soil system
has tremendous spatial diversity which has been little studied in relation to its biological
communities.

Implications of such microhabitat structure were cited by several authors. Greenslade
estimated that only about 10,000 years are required for an area to be completely reworked by
termites and Mermut showed the unique building block structures of such materials. Therefore,
it is reasonable to deduce that much of the soil in tropical areas is composed of remnants of
reworked termite mounds. Fjellberg mentioned that aggregation pheromones have been
detected for Collembola and the resultant aggregations have obvious but unstudied implications
for generation of microcommunities. Both Hill and Parkinson commented that soil animals are
themselves microhabitats which move, influencing dispersal of smaller animals, bacteria and
fungi. Water retained by surface tension around soil animals or their larval stages can be a
significant proportion of the total water film space available to soil microorganisms (McGill, in
preparation).

The guts of soil animals are also important microhabitats with respect to soil function.
Parkinson mentioned that bacteria are unaffected or increase in numbers upon passage through
the gut while fungi are damaged by passage through small organisms such as Collembola. The
gut of earthworms is a moist microhabitat where substrates are in motion and new surfaces are
acted upon by many smaller organisms. Fungal sporulation and spore movement are affected
by soil pore size distribution.

A recurring theme of the conference has been the importance of faecal pellets as
microhabitats which may dominate the fabric of some soils. Microcommunities and

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McGill and Spence

microenvironments may be characterized as mixed culture systems. Three postulates flow from
this concept: (i) species interactions such as symbiosis and, perhaps, mutualism may be more
characteristic and important to soil communities than are the results of succession, (ii) soil
animals not only alter their own environment, but are microhabitats for smaller organisms, and
(iii) the environment of a soil organism, and hence controls of its activity, are a function of its
size. Investigation of these three postulates could provide an initial framework for a more
synthetic soil ecology.

Fabric Reorganization and Locational Control

Pawluk emphsized that the exact involvement of soil fauna in forming soil microstructures is
inadequately understood for Canadian soils. A further problem, alluded to by Mermut, is the
lack of agreement among micromorphologists about standardized interpretation of soil fine
structure. Because Foster and Mermut, respectively, showed that soil animals can be involved
in both breakdown of structural units and in homogenization of materials, a dynamic picture of
soil micromorphology emerges. It appears that soil fabrics are in a constant state of slow
change; being generated, broken down and reorganized in cycles over long times. Such fabric
reorganization, when combined with the above ideas about microenvironment, lead to a concept
of biotic flux among substrates and environments. Such alterations in environment and
relocation of organisms near fresh substrates, or in barren locations could profoundly influence
how the system functions. It also provides an additional link with soil microbiology, further
emphasising the mixed culture aspect of the soil system.

The role of soil fauna in comminution of plant debris and in formation of the soil matrix is
becoming better understood (Seastedt, 1984). Ideas about communities developed from studies
of nutrient cycling can now be extended to include disintegration or comminution of soil
microstructures. Further research into this aspect of relations between soil animals and soil
structure is needed before the extent and significance of the process is known. Soil
microstructure influences the local environment and probability of substrate-organism contact
at microsites where biological processes occur. As a result, soil organic matter dynamics, and
soil quality, are influenced by fabric reorganization which comprises both formation and
comminution of microstructures. Soil animals may thereby provide an important control on soil
organic matter dynamics and soil quality.

Associated with the above is the effect of location, within or on soil, on the activities and
survival of organisms. For example Fjelberg pointed out the sensitivity of Collembola to water
supply because of the absence of an exoskeleton. One strategy is to live within soil layers where
relative humidity is higher. Other soil animals migrate up and down the profile in response to
soil moisture changes. Altemiiller showed that what an organism does in soil is influenced by its
position, and so behavioural studies of soil fauna must take micromorphological diversity into
account. At an even smaller scale, Foster showed how entrapment of organic molecules or
bacterial cells can result in their persistence through protection from decomposition or lysis.
The above locational control on organism function is fundamental to soil systems and appears
in turn to be modified by fabric reorganization. A type of feedback is thereby generated
because soil animals are among the agents responsible for fabric reorganization.

Soil Fauna and Soil Structure

653

SUMMARY AND CONCLUSIONS

The structure and function of soil systems are interrelated. Feedback between microhabitat
conditions and soil animals is characteristic of terrestrial ecosystems. The above interactions
link soil micromorphology and soil biology. System function and soil biophysics therefore
become the focus which permits advancement of knowledge in soil biology and pedology beyond
the capabilities of either discipline in isolation. Reciprocity between soil fauna and other soil
components must be recognized, however, and studied objectively before progress can be made.

Several ideas which may help guide future research have resulted from this synthesis. It is
postulated that soil fauna regulate soil systems through trophic interactions and biophysical
mechanisms. Trophic interactions which involve soil animals as microhabitats have been
reemphasized. Symbiosis, mutualism, and cohabitation are characteristic of soil communities,
perhaps superceding in importance interactions associated with successional changes.
Biophysical issues relating to size and location appear important. The relevant
microenvironment of an organism is clearly a function of its size. A related concept is that the
location of an organism determines its behaviour and the dynamics of its populations. We
propose that comminution and disintegration of microstructures be added to formation of
I microstructures and comminution of plant debris as a third biophysical mechanism by which
fauna regulate soil systems. Faunal influences on the dynamic relationships between soil
structure and function should receive major emphasis.

An immediate challenge remains to link specific groups of soil organisms to defined soil
microstructures as seen in thin sections. Related to this challenge is our recommendation for a
more ecologically useful approach to classifying both organisms and soil fabrics which is needed
I, to permit such links to be developed.

ACKNOWLEDGEMENTS

We thank the conference participants for a rich potpouri of stimulating ideas only partially
reflected in this summary, and J.S. Scott of the Department of Entomology, for preparing the
1 figures.

REFERENCES

Biological Survey of Canada (Terrestrial Arthropods). 1982. Status and research needs of
Canadian soil arthropods. Bull. Entomol. Soc. Can. 14(1), Suppl.

Crowson, R. A. 1970. Classification and biology. Heinemann Educational Books, Ltd.,
London. 350 pp.

Darwin, C. R. 1881. The formation of vegetable mould through the action of worms. London.
I; 326 pp.

j Elliot, E.T., Andereson, R.V., Coleman, D.C., and Cole, C.V. 1980. Habitable pore space and
microbial trophic interactions. Oikos , 35, 325-335.

: Hyams, E. 1976. Soil and Civilization. Harper and Row, New York. 312 pp.
i Kubiena, W. L. 1938. Micropedology. Collegiate Press, Inc., Ames, Iowa. 243 pp.
i Seastedt, T. R. 1984. The role of microarthropods in decomposition and mineralization
processes. Ann. Rev. Entomol. 29: 25-46.

| Simonson, R.W., (1968) Concept of soil. Adv. Agron ., 20, 1-47.

Quaest. Ent., 1985, 21 (4)
I

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Wolf, E. C. 1985. Erosion of productive soils by wind and water is changing the face of the
earth. Nat. Hist. 94(4): 53-57.

655

ADDENDA: TECHNIQUES, EQUIPMENT, ADDITIONAL REFERENCES, AND
PRIORITIES FOR FUTURE STUDY

Quaest. Ent., 1985, 21 (4)

SOIL MICROMORPHOLOGY

J.A. McKeague
and

C.A. Fox

Land Resources Research Institute
Agriculture Canada
Ottawa, Ontario K1A 0C6
CANADA

Quaestiones Entomologicae
21:657-664 1985

ABSTRACT

Soil micromorphology is a tool for studying a part of the continuum, from landscapes to
microvoids between soil particles. Methods are outlined for sampling soils and preparing thin
sections for study under the microscope. Features such as voids, aggregates, coatings, mineral
and organic particles and their arrangements are shown and described briefly. The potential is
outlined for applications of soil micromorphology to studies of soil genesis and of the
influence of fauna on soil properties. The annotated reference list aids interested readers to
delve farther into the fascinating architecture of soil as viewed in thin sections rather than as
amorphous dirt.

RESUME

La micromorphologie des sols represente un outil permettant d’etudier une partie du continuum de la morphologie
des sols, qui s’etend des pay sages jusqu’aux espaces microscopiques entres les particules. Les auteurs exposent dans leur
grandes lignes des methodes pour echantillonner les sols et pour preparer des coupes fines pour etude au microscope. Ils
montrent et decrivent brievement certaines particularites des sols telles que des vides, des agregats, des pellicules, des
particules minerales et organiques et leurs arrangements. Ils discutent du potentiel qu’ off rent diverses applications de la
micromorphologie des sols dans ietude de la genese des sols et de I'influence de la faune sur les proprietes des sols. Une
liste de references commentees aidera les lecteurs interesses d se familiariser davantage avec le sujet fascinant de
I’architecture des sols tels qu’observes en coupes minces, plutot que comme amas de terre amorphe.

INTRODUCTION

Soil micromorphology is the sub-discipline of soil science that includes studies of the
structure of relatively undisturbed soil samples with the aid of microscopes. It is part of
continuum that begins with observations of the pattern of soils in the landscape, proceeds to
{ studies of pedons (units of soil) representative of segments of that landscape, continues with
description and sampling of horizons within those pedons, of aggregates within the horizons,
and so on at increasing detail, to the study of features within aggregates as seen in thin sections
with the microscope (Fig. 1). Soil features ranging in size from approximately 10 to 10,000
can be studied in this section with the polarizing microscope. Scanning electron microscopy
| (SEM) is applied to the study of smaller features ( Bisdom, 1981).

Micromorphological techniques were applied rarely in the study of soils prior to the
| publication of the book ‘Micropedology’ (Kubiena, 1938). The use of micromorphology

658

McKeague and Fox

Fig. 1. Diagrammatic sketch indicating that soil micromorphology is a part of a continuum that includes macromorphology
of soils in the landscape.

increased slowly until after the publication of Brewer’s (1964) book “Fabric and Mineral
Analysis of Soils”. In it, Brewer defined terms precisely and outlined a system for describing
soil microstructure; the book continues to be a basic reference. Currently, many soil scientists
use micromorphological techniques in studies of soil genesis, physics, chemistry, minerals and
living organisms. Only a few, however, specialize in micromorphology.

In this paper we outline the steps involved in sampling, in preparation of soil samples for
study, and in applying micromorphological techniques to the study of soils. Examples are given
of kinds of problems amenable to study by micromorphology; some involving soil fauna and
related to soil structure are included. Further information on all aspects of soil
micromorphology is included in the references listed.

SAMPLING

Sampling is a crucial step in micromorphology. The first step is to decide the purpose of the
study. Suppose, for example, that the pedologist wants to determine the nature of the material
that cements the sand grains in a cemented horizon of a sandy pedon. A single clod of the
cemented material might be an adequate sample for preparation of a thin section, description
by microscopy of the material that links the grains, and analysis of the material by energy
dispersive X-ray analysis (Bisdom, 1981). If, on the other hand, the purpose of a study is to
determine differences in microstructure of surface horizons associated with differences in land
use, a systematic sampling plan involving replication of samples of similar soils under different
land use would be required.

After establishing the purpose and deciding on the sampling plan, the next step is to collect
the samples. The fundamental requirement is to obtain samples without altering the soil fabric,
the arrangement of particles and voids. For unconsolidated mineral soil materials, this is
usually done by pushing a metal frame or an open metal box, a Kubiena box, into the horizon to

Soil micromorphology

659

be sampled. Boxes of different sizes are used depending on habits, purpose of the study, and
nature of the soil. We use metal frames 8x6x5 cm, 8 x 6 x 2.5 cm and 6 x 4 x 2.5 cm made
from 20 gauge galvanized iron with the largest faces, 6 x 8 or 6 x 4 cm, open. The frame may be
pushed by hand or preferably jacked into a vertical or horizontal exposure of the horizon of
interest. The sample is trimmed flush with the edges of the frame, and its orientation is marked
on the metal frame. The trimmed sample is placed in a plastic bag to avoid loss of water, and
the open faces are covered with 9 x 7 cm pieces of plywood which are taped securely in place.
The site number and sample depth are marked on the sample with waterproof ink. Information
on the site and the sample is recorded in either a notebook, or a form such as a CanSIS file
form for soil data (Day, ed. 1982).

If the horizon to be sampled is strongly coherent, such as a cemented horizon, a clod may be
broken out, placed in a plastic bag, taped and labelled as indicated. Organic soil samples such
as peat may be obtained with a Macaulay sampler, which provides half a cylindrical sample
approximately 3.5 cm in diameter. These samples are placed in half cylinders of 3.75 cm PVC
piping, enclosed in a plastic bag, protected with a wooden cover, taped and labelled. Further
information on sampling is available (Sheldrick, 1984).

PREPARING SAMPLES FOR MICROMORPHOLOGICAL STUDY
Stereomicroscope

For some purposes such as describing the shapes and sizes of aggregates smaller than 2 mm,
it is useful to examine samples under a stereomicroscope at magnifications of 5 to 30x. Such
data complement macromorphological information obtained in the field.

Impregnating Samples for Preparation of Thin Sections

Most soil materials are unconsolidated so it is necessary to consolidate them by filling the
pores with plastic and hardening before preparing thin (20 or 30 ;um) sections. Water must be
removed prior to impregnating the soil with most of the plastics that are used. Three methods
have been used for removing water (Murphy, 1983): (1) oven drying - this results in shrinkage
of many soil materials and hence alteration of pore sizes, (2) freeze drying - this results in ice
crystal formation within the sample and, hence, some disruption of the fabric, (3) exchange of
water by acetone - this results in some dissolution of organic components but it is the best of the
three methods for preserving the soil fabric. Details of the drying procedues are given by
Fitzpatrick (1984), Sheldrick (1984) and references cited therein.

After water is removed by oven - or freeze-drying, the sample is put under vacuum so as to
remove air from voids, and the plastic mixture is added under vacuum. For acetone-exchanged
samples, voids are filled with acetone so vacuum is not necessary during the addition of plastic.
A variety of polyester resins diluted with thinners such as acetone, or epoxy resins, are used (see
Jongerius and& Heintzberger, 1975; Sheldrick, 1984; Fitzpatrick, 1984). Catalysts may be
added to increase the rate of polymerization and fluorescent dyes may be added to facilitate
study of pores. We use a polyester resin - acetone mixture; Uvitex OB (Ciba-Geigy), a
compound that fluoresces in ultraviolet light, is added in some studies (Sheldrick, 1984).
Usually in our laboratory 2 weeks to 2 months are required from the time the resin is added
until the impregnated block is hard. For some purposes, it is appropriate to use a resin mixture
that hardens in a few hours. Final curing of the plastic is done by heating the block to 60°C.

Quaest. Ent., 1985, 21 (4)

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McKeague and Fox

Fig. 2. Photomicrographs of features in soil thin sections. A. Partly-fused, rounded aggregate ‘a’ due to earthworms, a
large void ‘b’ and organic fragment ‘c’ in the Ah horizon of a clayey Hunic Gleysol near Ottawa. The width of the field is 8
mm. B. Fecal pellets ‘d’ of mites in decaying wood tissue ‘e’ from an Om horizon of an organic soil in Ontario. The width of
the field is 1.3 mm. C. Dark reddish-brown, amorphous coatings ‘f on sand grains ‘g\ and packing voids ‘IT in the Bhc
horizon of an Ortstein Humic Podzol from New Brunswick. The width of the field is 1.3 mm. D. Clay coating ‘F on
surfaces of dense clayey peds with embedded silt and sand grains ‘j’ and a planar void ‘k’ in the Bt horizon of an Orthic
Gray Luvisol from Ontario. The width of the field is 1.3 mm. E. Reddish-brown and black nodules enriched in Fe or Mn,
of Fe and Mn in the Bg horizon of an Orthic Humic Gleysol from the Fraser Valley, British Columbia. The field is 3.9 mm
wide. F. Dark reddish-brown aggregates ‘1’ in spaces between sand grains ‘m’ in a porous ‘n‘ Bf horizon on an Orthic
Humo-Ferric Podzol from Quebec. The width of the field is 1.3 mm.

Soil micromorphology

661

Preparing Thin Sections

The hardened sample is cut with a diamond saw to obtain a horizontally or vertically
oriented slab depending on requirements, approximately 1 cm thick. For samples containing
Uvitex OB, the cut face of the block may be photographed under ultraviolet light to show the
pore pattern. Pore configuration may be characterized quantitatively using an image analyzer
(Murphy, 1982; Bullock and Murphy, 1983). The sample orientation and number is marked on
the slab and the area to be used for thin sections is selected. The dimensions of the thin section
may be as large as the block or approximately 2x3 cm depending on the equipment available
and the purpose of the work. A chip of the appropriate size is cut, its orientation is marked, one
side is ground smooth on a diamond lap and cleaned. The chip is warmed on a hot plate, epoxy
cement is applied and a glass slide is fixed to the chip. The mounted chip is cut to a thickness of
approximately 0.5 mm on a diamond saw. It is ground on diamond laps with progressively finer
grit to a thickness of 20 to 30 iim. The thickness is checked by observing the section under
crossed polarizers with a polarizing microscope. Quartz grains appear white to grey if the
thickness is correct. The section is cleaned, a cover glass is applied with epoxy, and the sample
orientation and number are marked on the microscope slide.

Describing Thin Sections

Systems for describing thin sections are outlined in several publications (Brewer, 1964;
FitzPatrick, 1984; Bullock et al., 1985). We refer to the last system as it was developed by an
international committee. Sections are described under the following headings:

Microstructure. — The size, shape and arrangement of particles and voids. For example,
note the rounded aggregates, large voids and organic fragments in an Ah horizon (Fig. 2A).

Mineral and Organic components. — Mineral grains larger than approximately 20 /urn can
be identified by skilled microscopists. The nature and degree of decomposition of organic
components may be identified. For example Fig. 2B shows a decaying woody root fragment
with a cluster of mite pellets.

Groundmass. — The proportions and arrangements of coarse and fine components. In some
samples, the fine material occurs as coatings on coarse grains (Fig. 2C); in others coarse grains
are imbedded in a fine matrix (Fig. 2D).

Pedofeatures. — Features of the fabric due to soil genesis. For example, the coating of clay
on the heterogeneous matrix material adjacent to the planar void (Fig. 2D) is a pedofeature due
to deposition of clay from suspension. The dark brown and black nodules (Fig. 2E) are
pedofeatures due to segregation of Fe and Mn oxides in a soil that is saturated and under
reducing conditions periodically. The microaggregates in the B horizon of a Podzolic soil may
be due to physical processes or to soil fauna (Fig. 2F).

APPLICATIONS OF SOIL MICROMORPHOLOGY

Soil micromorphology has been applied principally to studies of soil genesis. In recent years,
however, applications to other areas of soil science, including soil zoology, have increased
markedly. Some examples of these applications are discussed briefly; others are found in
Bullock and Murphy (1983) and in proceedings of previous meetings of the International
Working Meeting on Soil Micromorphology.

Among the major applications of soil micromorphology in studies of soil genesis is seeking
direct evidence of translocation of fine particules from near-surface to subsurface horizons (see

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McKeague and Fox

papers in Bullock and Murphy, 1983; and Douglas, 1985). Surfaces of peds in horizons from
which clay has been removed are commonly uncoated or they may have coatings of coarser
grains due to loss of clay. Horizons in which clay has been deposited commonly have surfaces of
peds coated with oriented clay which appears finer and more uniform than the matrix material
(Fig. 2D). Micromorphology has been applied also in many studies of podzols, especially their
B horizons in which amorphous organic Fe, A1 materials accumulate as coatings (Fig. 1C), as
aggregates between grains (Fig. 2F), or both. A controversy continues over the origin of the
aggregates (Fig. 2F). Some believe that they are fecal pellets; others believe that they are the
result of physical processes, especially shrinkage on drying of the gel-like amorphous materials.

Micromorphology may be applied to soils studies other than those focused on genesis; a few
examples are listed. Micromorphological and associated sub-microscopic techniques are
powerful tools for studying the weathering of minerals in soils (Bisdom, 1981). Attempts have
been made to relate the sizes and shapes of voids seen in thin section to water flow in soils
(Bouma et al., 1979), Babel (1975) has shown the potential of micromorphological techniques
in studying, at high magnification, the decomposition of organic materials. Fox (1984) outlined
a system for describing the complexity of organic materials at a wide range of magnifications.
Other examples are given in proceedings of this symposium.

Kubiena (1938) was ahead of his time in recognizing the influence of soil fauna on structure
and he observed soil fauna directly in the field. Bal (1982) reviewed the literature on the
subject and reported results of his experiments showing faunal effects on soil structure. The
growing awareness in North America of the role of soil fauna will be accelerated by this
symposium. Many questions regarding the origin of aggregates and tubules in soils remain to be
resolved and caution will be required to avoid overstating the roles of soil fauna. Hypotheses
that could account for the common presence of rounded aggregates ranging in size from
approximately 20 /um to several mm must be tested objectively for different soil horizons.
Micromorphology will be a useful tool in such studies.

CONCLUSION

Examination of thin sections with the microscope complemented by submicroscopic
techniques leaves the observer with an expanded appreciation of the organized heterogeneity of
soil horizons. Soil samples prepared for chemical analysis appear to be amorphous dust. Thin
sections show the complex architecture of a host of different mineral crystals, aggregates of fine
particles, amorphous components and voids of differing sizes and shapes, some of them made by
soil fauna. Having viewed soil in thin section, the observer incorporates into his model of soil
the concept of its complex architecture and appreciates the influence of biological forces on that
architecture.

REFERENCES

Key references in soil micromorphology are listed and for some brief comments are added.

These reference and the papers cited therein will lead the reader into specific topics.

Babel, U. 1975. Micromorphology of soil organic matter, pp. 369-473. In: Gieseking, J.E.
(Editor). Soil components, Vol. 1, Organic components. Springer-Verlag. [Microscopy of
organic materials in soils is discussed; photomicrographs show plant tissues at various stages
of decomposition].

Soil micromorphology

663

Bal, L. 1973. Micromorphological analysis of soils. Soil survey papers No. 6. Soil Survey
Institute, Wageningen, The Netherlands. [A system for describing organic materials in soils
is presented].

Bal, L. 1982. Zoological ripening in soils. Agricultural Research Reports 850. Pudoc,
Wageningen, The Netherlands. [This book links soil zoology and soil micromorphology].

Bouma, J., A. Jongerius, and D. Schoonderbeek. 1979. Calculation of hydraulic conductivity of
some saturated clays using micromorphometric data. Soil Sci. Soc. Am. J. 43:261-265.

Brewer, R. 1976. Fabric and mineral analysis of soils. Robert E. Drieger Publ. Co.,
Huntington, New York. [This or the first edition (Wiley 1964) is considered by many soil
micromorphologists as the best comprehensive reference on the subject. It provided the first
systematic framework for describing soil microstructure. It is much stronger on mineral
than on organic soil features. See also a more recent account of some features of the system
by: Brewer, R. and S. Pawluk. 1975. Can. J. Soil Sci. 55:301-319].

Bisdom, E.B.A. (Editor). 1981. Submicroscopy of soils and weathered rocks. Pudoc,
Wageningen, The Netherlands. [This compilation of papers shows how conventional soil
micromorphology can be extended by the use of “submicroscopic” techniques including
scanning electron microscopy, image analysis (Quantimet), energy dispersive X-ray
analysis, and other techniques] .

Bullock, P., N. Fedoroff, A. Jongerius, G. Stoops, and T.Tursina. 1985. Handbook for thin
section description. Waine Research Publ. Wolverhampton. [This product of an
international committee should provide the framework for a standard system used
internationally].

Bullock, P. and C.P. Murphy (Editors). 1983. Soil micromorphology. Vols. 1 and 2. AB
Academic Publ., Berkhamsted, Herts. [These volumes, the proceedings of the International
Working Meeting on Soil Micromorphology, London, 1981, include papers on a wide range
of topics and show the current state of soil micromorphology].

Day, J.H. (Editor). 1982. Manual for describing soils in the field. LRRI No. 82-52. Agr. Can.
Ottawa.

Douglas, L.A. et al. (Editors). 1985. Micromorphology and Soil Classification. Soil Sci. Soc.
Am. Spec. Publ. (in press).

FitzPatrick, E.A. 1984. Micromorphology of Soils. Chapman and Hall. [This book by a teacher
of the subject explains how to prepare, describe and interpret thin sections of soil].

Fox, C.A. 1984. A morphometric system for describing the micromorphology of organic soils
and organic layers. Can. J. Soil Sci. 64:495-503.

Jongerius, A. and G. Heintzberger. 1975. Methods in soil micromorphology. A technique for
the preparation of large thin section. Soil Survey Paper No. 10. Soil Survey Institute,
Wageningen, The Netherlands.

Jongerius, A. and G.K. Rutherford. 1979. Glossary of soil micromorphology, Pudoc,
Wageningen, The Netherlands. [This glossary provides definitions in English of many of the
terms used in soil micromorphology. The terms are given in English, French, German,
Spanish and Russian].

Kubiena, W.L. 1938. Micropedology. Collegiate Press Ames Iowa. [The enthusiam and
preceptiveness of the author, who may be considered as the founder of soil
micromorphology, are clearly evident to current readers, half a century after the book was
written].

Murphy, C.P. 1982. A comparative study of three methods of water removal prior to resin

Quaest. Ent., 1985, 21 (4)

664

McKeague and Fox

impregnation of soils. J. Soil Sci. 33:719-735.

Sheldrick, B.H. (Editor). 1984. Analytical methods manual 1984. Land Resource Research
Institute No. 84-30. Agr. Can., Ottawa.

PRIORITIES FOR THE INTEGRATED DEVELOPMENT OF SOIL
MICROMORPHOLOGY AND SOIL ZOOLOGY: RESULTS OF A BRAINSTORMING

SESSION

S.B. Hill

Department of Entomology
Macdonald College

Ste. Anne de Bellevue, Quebec H9Z ICO
CANADA

V.M. Behan-Pelletier
Biosystematics Research Institute
Agriculture Canada
Ottawa, Ontario K1A ICO
CANADA

Quaestiones Entomologicae
21:665-668 1985

During a two-hour workshop, 50 soil micromorphologists and soil zoologists participated in
an exercise designed to identify priorities for the integrated development of their disciplines.

Several months before this workshop, a list of 20 needs of soil zoologists, identified by us,
was circulated to other speakers as background materials to the workshop. This list was also
included in the registration package given to all symposium participants. These topics have
been integrated into the full list of identified needs given below (those not mentioned during the
workshop have an asterisk).

The participants were divided into 10 groups of five, each group including representatives of
both disciplines. Two five minute exercises were then conducted during which participants
brainstormed (listed, uncritically, as many things as they could think of in the time available)
on the questions “in what ways can members of the other discipline benefit your discipline” and
“in what ways can members of your discipline help the other discipline”. One member from
each group recorded responses, and all responses were then listed on flip charts and posted in
view of all participants.

Groups were then encouraged to expand their initial lists by imagining that there were no
restraints on their proposals and that ideal conditions prevailed. Finally, each group was asked
to identify, taking into account their previous suggestions, the three most important needs to
achieve the integrated development of the disciplines. These priority needs were combined and
then arranged under the following six headings. The more extensive list of suggestions, referred
to earlier, is given in an Appendix using the same headings.

666

Hill and Behan-Pelletier

PRIORITY NEEDS

1. Policy

Identify changes in government policies that will facilitate the integrated
development of soil micromorphology and soil zoology.

Identify ways to translate research findings into improved soil management
practices.

Improve public education (use of media, etc.) concerning the importance of these
disciplines.

Increase the number of university positions in these disciplines.

Establish multidisciplinary “Soil Institutes” (building, for example, on the
experiences of Dr. Josef Rusek, Director of the Laboratory of Soil Biology,
Institute of Landscape Ecology, Czechoslovakia).

2. Research

Facilitate and support multidisciplinary research (from planning to publication).
Improve international planning and cooperation of research programs (building
on International Biological Program experience).

3. Education

Establish educational programs that integrate these disciplines, at least at the
University level.

Provide general and specialized short courses (and field trips), covering the
various aspects of these disciplines.

4. Networking

Produce a directory of specialists noting their fields of interest and current
projects.

Continue to hold joint symposia for these (and other related) disciplines.

5. Literature

Produce low cost, high quality textbooks providing syntheses of what is known
and unknown in these disciplines.

Prepare illustrated, comprehensive, easy-to-use keys and atlases of soils and soil
organisms.

6. Techniques

Develop reliable, standardized, inexpensive and easy-to-use techniques for
conducting research in these disciplines.

APPENDIX: FULL LIST OF NEEDS IDENTIFIED DURING WORKSHOP
1. Policy

Dissolve disciplines (at least at the edges).

Shift emphasis to long-term multidisciplinary studies.

Identify potential sources of institutional support.

Soil Micromorphology and Soil Zoology

667

Promote institutional support.

Improve public education (better use of media etc.).

Establish chairs in soil biology /soil micromorphology.

Establish “Soil Institutes” integrating these and other related disciplines.
Facilitate interaction between members of these disciplines in universities,
institutes, etc..

4. Networking

Establish a common journal.

Produce a directory of specialists noting their fields of interest and current
projects (including willingness to identify and describe soils and soil fauna).
Establish data banks with minimum access costs.

Continue to hold joint symposia for these (and othe related) disciplines, including
workshops and think-tanks e.g., to continue the initiative described in this paper.

* Prepare directories of special facilities and equipment that are not widely
available.

2. Research

A. Requests from soil micromorphologists to soil zoologists:

Clarify relationships between soil community and soil type.

Identify and provide information concerning the distribution (horizontal and
vertical) of soil animals, exuviae and faeces.

Clarify the ecological importance and influence on physical factors of different
species/genera/orders of soil animals, e.g., their role in decomposition of organic
matter.

Provide qualitative and quantitative data concerning the feeding habits of
different soil animals.

Describe the niche characteristics of different soil animals.

Collaborate in research projects (from planning to publication) and establish
multidisciplinary research teams.

Consider the needs of soil micromorphologists when selecting soil zoology
research topics.

Provide physico-chemico-biological descriptions of faeces of different soil
animals.

Describe the major ‘types’ of soil communities.

B. Requests from soil zoologists to soil micromorphologists:

Describe the major chemical transformations in soil.

Describe the micromorphology of different soils, including identification of
potential food sources, mineral and humus composition, pore spaces, etc.

Describe the micromorphology of the stages in humification.

Collaborate in research on “problem soil profiles”.

Collaborate in research on the acceleration of soil-forming processes.

Describe the impact of salinization on soil structure and function.

Quaest. Ent., 1985,21 (4)

668

Hill and Behan-Pelletier

6. Techniques

Develop non-destructive research techniques ( e.g ., sampling).

Develop reliable, standardized, inexpensive and easy-to-use research techniques.
Develop improved methods for embedding, staining, making thin sections and
analyzing soil (e.g., discover a water miscible, non-toxic embedding resin).

* Prepare “Cookbooks” of techniques (including hints not usually given in
textbooks).

* Prepare a “Consumer guide” to equipment (giving advantages and
disadvantages).

* Prepare a “Cookbook” of statistical techniques and a list of computer and micro
computer program packages that are especially useful for soil fauna and soil
micromorphology research.

3. Education

Emphasize to soil scientists the living reality of soil.

Share information “at the microscope” (soil fauna/faeces identification, ped
description, etc.)

Organize joint field trips.

Provide general and specialized short courses (including laboratory experience
and field trips) covering the various aspects of these disciplines (for professionals
and non-professionals).

Guide students enrolled in each of the disciplines to attend one or more courses in
the other discipline.

Produce learning packages, tapes, films, slide sets, video-tapes, modules etc., on
all aspects of these and related disciplines, and prepare guides to existing
materials.

5. Literature

Prepare guides (annotated bibliographies, etc.) to the basic literature.

Prepare illustrated, comprehensive, easy-to-use keys to adult and immature soil
animals (by habitat, region, feeding group, etc.).

Produce low-cost, high quality textbooks.

Prepare basic comprehensive atlases of soil micromorphology (including three
dimensional views of pore spaces).

Prepare thesauri of soil zoology and soil micromorphology.

Prepare comprehensive dictionaries of soil zoology and soil micromorphology.
Produce directories of translations of relevant books and papers.

Produce directories of grants, with tips on grantsmanship.

A VARIATION OF THE MERCHANT-CROSSLEY SOIL MICROARTHROPOD

EXTRACTOR

Roy A. Norton

Department of Environmental and Forest Biology

State University of New York

College of Environmental Science and Forestry

Syracuse, New York 13210

U.S.A.

Quaestiones Entomologicae
21:669-671 1985

The Merchant-Crossley extractor is an inexpensive apparatus which rivals more elaborate
constructions in efficiency (Merchant and Crossley, 1970; Seastedt and Crossley, 1978). The
design suggested here involved modifications to improve ease of operation and to minimize
lateral contaminations. As in the original, the 2 inch diameter soil corer used is commercially
available from “Art’s Machine Shop” (Harrison and Oregon Trail, American Falls, Idaho
83211) and costs slightly more than $100 (U.S.). The steel sampling cup can be ordered to any
length, as can the aluminum retaining cylinders.

Soil cores taken in the field are trimmed at the bottom end, placed in individual plastic bags,
left open at the top, and kept in an ice-chest until extraction. The top of each retaining cylinder,
with the core intact, is covered with a small individual fiberglass screen. It is cut slightly larger
than the cylinder and held in place by a plastic retaining-cylinder cover (also available from
“Art”), from which most of the center has been cut out so that little more than a “lip” is left to
hold the screen in place. This soil-cylinder-screen unit is then inverted, placed into the extractor
hole from below, and held up by two heavy rubber bands, wrapped around the upper part of the
cylinder.

The extractor itself is constructed from varnished 1/2” plywood. I have found the most
convenient design to be that shown in the accompanying figure. Heat is provided by 7 watt
“nite-lite” bulbs, either used in a Christmas tree string, or preferably individually wired “cleat
receptacles” attached to the extractor cover. Individual tin or aluminum (12 oz.) cans (with
both ends cut out) between the top and core provide reflection and maintain heat. Due to the
individual nature of each unit, there is no “edge-effect” in terms of extraction efficiency, as
there often is with extractors using a common heat source. Because of the low wattage of each
20-unit extractor, an inexpensive household dimmer-switch can be wired and attached to the
outside of the cover for the control of light intensity.

The extractor is very adaptable in terms of collection method. As in the original version,
collection into alcohol can be accomplished with the use of powder funnels forced through holes
cut into caps of collecting vials. Vials are about 1/3 filled, so that room is left for the funnel
which is rinsed with alcohol after extraction is complete.

If extraction over water is preferred, appropriate size plastic jars are used instead of the
powder funnel-vial unit. An ideal, inexpensive screw-cap plastic jar is a 4 1/2 oz. wide-mouth
specimen-container used in hospitals (Superior Plastic Products Corp., P.O. Box 2128,
Providence, RI 02905). The cap (with the center cut out to the diameter of the cylinder cap) is
I attached to the underside of the middle layer of the extractor, so that the jar (1/3 filled with
| water) can be screwed in from below. An effective, and very inexpensive canister extractor is

670

Norton

the result. It prevents drying of the core from below, and eliminates the use of funnels, which
provide condensation surfaces and allow potential escape. Seastedt and Crossley (1978) had
somewhat poorer results with a canister-style apparatus, but this was probably due to their use
of alcohol, instead of water, as a collecting fluid.

The compact 20-unit extractors can be removably wall-mounted, shelf-mounted, or placed
on small tables; they can even be stacked if side ventilation holes are provided. They can be
used in refrigerators or both small and large environmental chambers and are extremely
portable.

REFERENCES

Merchant, V.A. and D.A. Crossley, Jr. 1970. An inexpensive high-efficiency Tullgren extractor
for soil microarthropods. J. Georgia Entomol. Soc. 5: 83-87.

Seastedt, T.R. and D.A. Crossley, Jr. 1978. Further investigations of microarthropod
populations using the Merchant-Crossley high-gradient extractor. J. Georgia Entomol. Soc.
13: 338-344.

Merchant-Crossley Soil Microarthropod Extractor

671

Fig. 1 . Diagram of a Modified Merchant-Crossley Extractor for Soil Microarthropods.

SUPERIOR MICRO-NEEDLES FOR MANIPULATING AND DISSECTING SOIL

INVERTEBRATES

R. Norton

State University of New York

College of Environmental Science and Forestry

Syracuse, New York 13210

U.S.A.

F. Sanders

Wayne State University
Detroit, Michigan
U.S.A.

Quaestiones Entomologicae
21:673-674 1985

Sturdy, yet very sharp, needles for dissecting microarthropods or manipulating small objects
cannot be purchased, but can be easily made with little equipment. The standard use of insect
minuten-pins is often not satisfactory due to poor quality control, improper taper, flaking and
corrosion and other problems which can be avoided by electrolytically produced needles.

A suggested apparatus is shown in the accompanying figure. A deep well slide is attached to
the bottom of a standard petri-dish with a strong epoxy cement (the top covers the apparatus
when not in use). Then an old microscope stage slip (or a similarly shaped piece of thin
aluminum sheeeting) is epoxyed to the slide, with the bent tip pointing into the well (touching
the bottom, if possible) and the other end bent upwards about 1 /2 inch. Cover the clip with
epoxy, except at the bent ends.

The needle is cut from 10 mil (0.25 mm) tungsten wire, and held in a zero-closure
pin-holder. Wire can be obtained, for example, from Alfa Products, Thiokol/Ventron Division,
152 Andover Street, Danvers, MA 01923 (catalogue #0371) and costs about $20 (U.S.) for a
20 m roll (a life-time supply). The pin-holders can be obtained from Fine Science Tools Inc.,
321 -B Mountain Highway, North Vancouver, B.C. V7J 2K7 and cost $8-10 (U.S.) each
depending on style and length. Good holders are recommended, rather than cheaper varieties
which are not zero-closure.

The only other necessary equipment is a D.C. power supply, about 5-10 volts and 0.5- 1.0
! amp. Some calculator-style supplies are satisfactory, but a transformer from an old
| stereo-microscope or compound microscope illuminator would be ideal. Also, the output leads
from the transformer need to be supplied with electrical attachment clips for good contact.

In operation, one fills the well with 10% KOH, attaches one lead to the clip of the well slide
! and the other to the end of the pin-holder (with the appropriate length of wire in place).

| Correct polarity is essential, but usually has to be determined the first time by trial and error (if
I the wire does not sharpen after a half minute or so, switch leads). Once the transformer is
j activated and the needle is placed in the KOH, rapid bubbling should surround the needle. Rate
of electrolysis is controlled by the transformer setting and the distance the needle is held from
I the well-slide clip. A good simple needle can be made in less than one minute.

The desired taper of the needle is a function of its projected use and individual preference.

I Taper can be controlled by the angle at which the needle is immersed in the KOH bath.

674

Norton and Sanders

Fig. 1. Diagram of Equipment and Procedure for Manufacture of Micro-Needles used in Morphological Dissection of Soil
Invertebrates.

Near-vertical orientation produces a short, thick taper; near horizontal orientation produces a
long, fine taper. The tip will be destroyed if accidentally touched to the well-slide clip while
current is on.

Bending the needle with forceps prior to electrolysis is usually desirable for working
comfort. Micro-hooks can be first bent, then electrolyzed; another way is to first taper a needle
in the usual way and then press it against a hard object to curl it, with finishing touches put on
after that.

Electrolysis causes a mist of KOH to be produced, so a good place to work is in a fume-hood.
Have a nearby stereo-microscope set up to intermittently check progress, but do not operate too
close to the microscope. Once formed, needles can be redressed in a few seconds and the
tungsten wire need be replaced (or re-bent) only after repeated quick sharpenings. Since the
tips are very fine, cover the end of the pin-holder when not in use. The cut-off tips of soft plastic
disposable eye-droppers serve nicely, as do some of the stiffer rubber bulbs, or tubings.

SOIL INVERTEBRATES: MAJOR REFERENCE TEXTS

Compiled by:

V.M. Behan-Pelletier'

S.B. Hill1 2
A. Fjellberg3
R.A. Norton4 and
A. Tomlin5

Quaestiones Entomologicae
21:675-687 1985

This bibliography is incomplete but should serve as an introduction to the literature on the
various groups of soil invertebrates. (Research papers are almost all omitted. Works in
languages other than English (some of which fill important gaps) are almost all omitted;
references to most of them will be found in the books listed.

GENERAL

D’Aguilar, J., C. Athias Henriot, A. Bessard, M.-B. Bouche and M. Pussard. (Editors). 1971.
Organismes du sol et production primaire. IV Colloquium Pedobiologiae. Institut National
de la Recherche Agronomique, Paris. 590 pp.

Important research papers on the soil ecosystem, including one on benefits of seeding a
wormless soil with earthworms. (35 Engl., 1 1 Ger., 9 Fr.). Refs, after each paper.

Anderson, J.M. and A. MacFadyen. (Editors). 1976. The Role of Terrestrial and Aquatic
Organisms in Decomposition Processes. Blackwell, Oxford, England. 474 pp.

Proceedings of British Ecological Society Symposium. Covers physico-chemical aspects of
the environment, interrelationships of organisms involved and their role in soil and fresh
water ecosystems and modelling of decomposer systems. Refs, after each paper.

Bornebusch, C.H. 1930. The Fauna of Forest Soil. Nielsen & Lydiche, Copenhagen. ( From
Forst. Forsogsv. i Danmark, II: English and Danish). 224 pp.

A classic.

Brauns, S. 1968. Praktische Bodenbiologie. Gustav Fisher Verlag, Stuttgart, xviii + 470 pp.
Classic text on Soil Biology with a practical emphasis. Not as yet translated into English.
Refs, after each chapter.

Burges, A. and F. Raw. (Editors). 1967. Soil Biology. Academic Press, New York. 532 pp.

An anthology of technical papers discussing biology of different classes of soil life.
Cloudsley-Thompson, J.L. 1958. Spiders, Scorpions, Centipedes and Mites. Pergamon Press,
Oxford. 278 pp.

(Paperback edition 1968). Includes most groups other than hexapods besides those in title.
Illustrates their diversity and versatility. Refs, after each chapter.

Cloudsley-Thompson, J.L. 1967. Microecology. (Institute of Biology’s Studies in Biology, 6).
Edward Arnold Ltd., London. 48 pp.

1 Biosystematics Research Institute, Agriculture Canada, Ottawa, Ont. K 1 A 0C6, CANADA

department of Entomology, Macdonald College, Stc. Anne de Bellevue, Quc. H9X ICO, CANADA

3Zoologisk Avdeling, Tromso Museum, Tromse, Norway.

4R.A. Norton, Department of Forest Biology, SUNY, CESF, Syracuse, NY 13210, U.S.A

5Research Centre, Agriculture Canada, London, Ont., N6A 5B7

676

Behan-Pelletier et al.

Simple introduction to predominantly soil and related fauna. 23 refs.

Cloudsley-Thompson, J.L. & J. Sankey. 1961. Land Invertebrates. Methuen, London. 156 pp.

Introductory, relates to various groups of soil animals.

Coineau, Y. 1974. Introduction a l’etude des Microarthropodes du sol et de ses annexes.
Documents pour l’enseignement pratique de l’ecologie. Doin, Paris, 188 pp.

Excellent textbook, with chapters on abiotic and biotic factors in the soil, techniques,
taxonomy and biology and a comparison of the microfauna of different soil types.

Danks, H.V. 1979. Canada and its insect fauna. Mem. Ent. Soc. Can. 108: 573 pp.

Synopsis of information available on insects and related arthropods in Canada.

Dickinson, C.H. and G.J.F. Pugh. (Editors). 1974. Biology of Plant Litter Decomposition. Vol.
1 & 2. Academic Press, New York. 241 & 775 pp.

The most comprehensive work on the subject. Refs, after each chapter.

Dindal, D.L. (Editor). 1980. Soil Biology as Related to Land Use Practices. Proc. VII Int. Soil
Zoology Colloquium. Office of Pesticide and Toxic Substances, EPA, Washington, DC. 880

pp.

Current research on the effects of human ameliorations and perturbations on soil organisms.
Doeksen, J. and J. van der Drift. (Editors). 1963. Soil Organisms. North Holland Publ. Co.,
Amsterdam. 453 pp.

Research on the biology and ecology of soil organisms. Refs, after each chapter.

Drift, J. van der. 1951. Analysis of the Animal Community in a Beech Forest Floor. Institut v.
Toegepast Biologisch Onderzoek in de Natuur, Mariedaal, Oosterbeek, Netherlands. [From
Tijdscjr. v. Ent. 94 (1)]. 168 pp.

Oligochaetes and molluscs receive brief notice.

Graff, O. and J.E. Satchell. 1967. Progress in Soil Biology. North Holland Publ. Co.,
Amsterdam. 656 pp.

Anthology of technical and specialized papers from German colloquium. Half of papers are
in German. Refs, after each presentation.

Haarl^v, N. 1960. Microarthropods from Danish Soils: Ecology, Phenology.

Andelsbogtrykkeriet, Odense, Denmark. (From Oikos, Suppl. 3). 176 pp.

Jackson, R.M. and F. Raw. 1966. Life in the Soil. Edward Arnold Ltd., London. 60 pp.

Authors from Rothamsted discuss soil ecology and ways of studying it. 21 refs.

Kaestner, A. 1967. Invertebrate Zoology. Vol. I. John Wiley and Sons Inc. 597 pp.
(Translation of earlier German work).

Includes information on biology, ecology and general classification of platyhelminths,
rotifers, nematodes, molluscs and annelids.

Kaestner, A. 1968. Invertebrate Zoology. Vol. II. John Wiley and Sons Inc. 472 pp.
(Translation of earlier German work).

Includes information on biology, ecology and general classification of tardigrades,
arachnids, centipedes, millipedes, pauropods and symphylans.

Kaestner, A. 1970. Invertebrate Zoology. Vol. III. John Wiley and Sons Inc. 597 pp.
(Translation of earlier German work).

Includes information on biology, ecology and general classification of isopods and
amphipods.

Kevan, D.K.McE. (Editor). 1955. Soil Zoology. Butterworths Scientific Publications, London
& Academic Press, New York.512 pp.

First integrated study of soil fauna on an international footing. It is still a basic reference

Soil Invertebrates-Reference Texts

677

work. Refs, after each paper.

Kevan* D.K.McE. (Editor). 1968. Soil Animals. H., F. & G. Witherby, London, &
Philosophical Library Inc., New York. 244 pp.

Corrected and slightly augmented reprinting of 1962 edition which covers all groups of soil
animals.

Kuhnelt, W. 1976. Soil Biology: with special reference to the Animal Kingdon. 3rd ed. Faber &
Faber, London. 397 pp.

Draws mostly on work from European soils, but is still very good for general description of
soils and the animals in them. 1700 refs.

Lawrence, R.F. 1953. The Biology of the Cryptic Fauna of Forests. A. A. Belkema, Cape
Town. 408 pp.

Emphasis on fauna of indigenous forests of South Africa. 13 pp. of refs.

Lebrun, Ph., H.M. Andre, A. de Medts, C. Gregoire-Wibo and G. Wauthy. (Editors). 1983.
New Trends in Soil Biology, Proc. VIII. Inti. Colloquium of Soil Zoology,
Louvain-la-Neuve, Belgium. 1982. 700 pp.

Most recent text on research in soil biology. Topics are covered in four major areas: the role
played by soil fauna in mineral cycling; functional relationships between soil organisms;
ecophysiology of soil animals and restoration capacity of soil communities.

MacFadyen, A. 1963. Animal Ecology: Aims and Methods. 2nd ed. Sir Isaac Pitman & Sons,
London. 344 pp.

A good ecology text that emphasises the soil ecosystem. Approx. 1000 refs.

Murphy, P.W. (Editor). 1962. Progress in Soil Zoology. Butterworths, London. 398 pp.

Technical papers about methods of extracting soil animals. Refs, after each presentation.
Pesson, P. (Editor). 1971. Le vie dans les sols. Aspects Nouveaux. Etudies experimentales.
Gautier-Villars, Paris, x + 472 pp.

Comprehensive review of ecology of soil organisms with emphasis on current research. Refs,
after each chapter.

Phillipson, J. (Editor). 1971. Methods of study in quantitative soil ecology: Population,
production and energy flow. I.B.P. Handbook No. 18. 297 pp. Blackwell, Oxford.

Good overview of current techniques. Refs, after each presentation.

Pimental, R.A. 1967. Invertebrate Identification Manual. Van Nostrand Reinhold Co., New
York. 151 pp.

Excellent illustrations of the major orders and families.

Richards, B.N. 1974. Introduction to the Soil Ecosystem. Longman Inc., N ew York. 266 pp.

A good modern approach to the subject, from Australia. Refs, after each chapter.

Savory, T. 1971. Biology of the Cryptozoa. Merrow Publishing Co., Watford, England. 56 pp.
Introductory textbook.

Schaller, F. 1968. Soil Animals. Univ. Mich. Press, Ann Arbor. 145 pp.

Introduction to soil ecosystems, emphasizing larger soil animals. No references.

Sheals, J.G. (Editor). 1969. The Soil Ecosystem. Systematics Assoc., London. 247 pp.

Proceedings of conference emphasising problems of classification of soils and their
components. Includes review on impacts of agricultural practices. Refs, after each
presentation.

U.N.E.S.C.O. 1969. Soil Biology: Review of Research. U.N.E.S.C.O., Paris. 244 pp.

Summarizes knowledge of soil biological processes. Refs, after each chapter.

Vanek, J. (Editor). 1975. Progress in Soil Zoology. Proceedings of the 5th International

Quaest. Ent., 1985,21 (4)

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Behan-Pelletier et al.

Colloquium on Soil Zoology held in Prague, September 17-22, 1973. Academia Publishing
House, Prague. 630 pp.

Most recent text on research in soil zoology as of 1975. Section on influence of human
activities on soil organisms. Refs, after each presentation.

Wallwork, J.A. 1970. Ecology of Soil Animals. McGraw-Hill, New York. 283 pp.

Basic text updating some of the material in Kevan (1962). Refs, after each chapter.
Wallwork, J.A. 1976. The Distribution and Diversity of Soil Fauna. Academic Press, London.
355 pp.

A continuation of his Ecology of Soil Animals (1970). Refs, after each chapter.

Webb, J.E., J.A. Wallwork and J.H. Elgood. 1978. Guide to Invertebrate Animals. 2nd ed.
Macmillan Press Ltd., London. 305 pp.

Guide for undergraduates with up-to-date classification scheme. Limited illustrations.
PROTOZOA

MacKinnon, D.L. and R.S.T. Hawes. 1961. An Introduction to the Study of Protozoa.
University Press, Oxford.

Good section about methods.

Stout, J.D. and O.W. Heal. 1967. Protozoa, pp. 149-195. In Burges, N.A. and F. Raw.
(Editors). Soil Biology. Academic Press, New York.

Excellent reference text on soil protozoa.

PLA TYHELMINTHES

Schmidt, G.D. 1982. Platyhelminthes. pp. 727-823. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 1. McGraw-Hill Book Co.

Primarily parasitic, some platyhelminthes are free-living in highly organic moist soils.
GASTROTRICHA

Hummon, W.D. 1982. Gastrotricha. pp. 857-863. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 1. McGraw-Hill Book Co.

ROTIFERA

Donner, J. 1966. Rotifers. Frederick Warne & Co., Ltd., London. 80 pp.

Simple, yet thorough introduction to “wheel animals”. Mainly aquatic. 29 refs.

Nogrady, T. 1982. Rotifera. pp. 865-872. In Parker, S.P. (Editor). Synopsis and classification
of living organisms. Vol. 1. McGraw-Hill Book Co.

Up-to-date classification of this group.

NEMATOMORPHA

Maggenti, A.R. 1981. General Nematology. Springer Verlag, New York. 372 pp.

Contains a short (pp. 27-32) section on Nematomorpha or gordian worms.

NEMATODA (= NEMATA)

Bird, A.F. 1971. The Structure of Nematodes. Academic Press, New York. 318 pp.

General textbook on the morphology and physiology of nematodes.

Chitwood, B.G. and M.B. Chitwood. (Editors). 1950. Introduction to Nematology. University
Park Press, London. 334 pp.

Papers on morphology, physiology and life histories.

Croll, N.A. 1970. The Behaviour of Nematodes, their activity, senses and responses. Edward
Arnold Ltd., London. 117 pp.

Croll, N.A. and B.E. Matthews. 1977. Biology of Nematodes. John Wiley and Sons, New
York, Toronto. 201 pp.

Introductory textbook presenting a unified view of the whole range of nematode types.

Soil Invertebrates-Reference Texts

679

Goodey, J.B. 1963. Laboratory Methods for Work with Plant and Soil Nematodes. 3rd ed.

Ministry of Agriculture, Fisheries & Food (Tech. Bull. 2), London. 47 pp.

Maggenti, A.R. 1981. General Nematology. Springer Verlag, New York. 372 pp.

Excellent general textbook on nematode morphology, physiology and classification. Third of
book deals with parasitic types, no special section on free-living soil nematodes.

Maggenti, A.R. 1981. Nemata. pp. 879-929. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 1. McGraw-Hill Book Co.

Up-to-date classification of group.

Nicholas, W.L. 1975. The Biology of Free-Living Nematodes. Clarendon Press, Oxford. 219

pp.

“This is a book written by an enthusiast for the unconvinced” and covers the morphology,
physiology, biochemistry, ecology, culturing, techniques and introductory taxonomy of these
invertebrates.

Poinar, G.O., Jr. 1983. The Natural History of Nematodes. Prentice-Hall Inc., New Jersey.
323 pp.

Introductory textbook to biology and ecology of nematodes. Includes section on
classification.

Sasser, J.N. and W.R. Jenkins. (Editors). 1960. Nematology: Fundamentals and recent
advances with emphasis on plant parasitic and soil forms. University of North Carolina
Press, Chapel Hill, N.C. 480 pp.

Contains an excellent section on methodology.

Southey, J.F. (Editor). 1959. Plant Nematology. Ministry of Agriculture, Fisheries & Food
(Tech. Bull. 7), London. 175 pp.

Lecture course, which reviews the subject and emphasizes problems in British agriculture.
Refs, after each chapter.

Southey, J.F. (Editor). 1970. Laboratory Methods for Work with Plant and Soil Nematodes.
Ministry of Agriculture, Fisheries & Food (Tech. Bull. 2), London. 148 pp.

A great “how-to” book on methods of studying nematodes.

Thorne, G. 1961. Principles of Nematology. McGraw-Hill Book Co., New York, Toronto,
London. 553 pp. 31 pp. refs.

ANNELIDA (OLIGOCHAETA)

Anonymous. 1982. Earthworms: Raising, uses, beneficial aspects 1978-1981, 97 citations.
Produced from AGRICOLA database. Available free from U.S.D.A. Library, Beltsville,
MD.

Appelhof, M. (Editor). 1981. Workshop on the Role of Earthworms in the Stabilization of
Organic Residues. Vol. 1. Kalamazoo, Michigan, April 9-12, 1980. Beech Leaf Press,
Kalamazoo, Michigan. 315 pp.

Has no bearing on identification but, besides being interesting, this book has an extensive
list of research needs in Appendix.

Bal, L. 1982. Zoological Ripening of Soils. Centre for Agricultural Research and
Documentation, Wageningen, Netherlands. Agricultural Research Reports 850.

A monograph on the contribution of soil animals to soil structure, in which earthworms have
an important role. The stilted text is redeemed by a number of excellent photographs, a
glossary and a large bibliography.

Bouche, M.B. 1972. Lombriciens de France: Ecologie et systematique. Ann. Zool. Ecol. Anim.
72(2): 214-472.

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Darwin, C. 1897. The formation of vegetable mould, through the action of worms with
observations on their habits. John Murray, London, vii + 328 pp.

As relevant today as in 1897. There have been several reprintings. Darwin was one of the
first to realise (and put his thoughts and painstaking observations into print) how
earthworms affect the landscape. His estimates of earthworm ( L . terrestris) density are
probably low. Refs, throughout text.

Edwards, C.A. and J.R. Lofty. 1972. Biology of Earthworms. Chapman & Hall, London. 283

pp.

Comprehensive review of all aspects of earthworm biology and ecology. 32 pp. of refs. 2nd
edition in 1977 with a more complete bibliography.

Gates, G.E. 1972. Burmese Earthworms. An introduction to the systematics of megadrile
oligochaetes with special references to Southeast Asia. Trans. Amer. Philos. Soc. 62(7):
1-326.

Lee, K.E. 1959. The Earthworm Fauna of New Zealand.. N.Z. Dept. Scientific & Industrial
Research, Auckland. 486 pp.

Has general application.

Nielsen, C.O. and B. Christensen. 1959, 1961, 1963. The Enchytraeidae: Critical Revision and
Taxonomy of European Species. Natura Jutlandica. Vols. 8, 9, 10.

Essential for workers on Enchytraeidae.

Reynolds, J.W. 1977. The Earthworms (Lumbricidae and Sparganophilidae) of Ontario. Royal
Ontario Museum, Life Sciences, Misc. Publications. 141 pp.

A comprehensive review of the Ontario earthworm fauna. Very well illustrated by Dan
Dindal. Indispensable for Ontario workers. Good bibliography with the bonus that Reynolds
cites his and Gates’ papers in the Bull. Tall Timbers Res. Stn. and Megadrilogica up to
1977. Reynolds has published extensively on the earthworm fauna of several American
states (particularly in the Northeast) and several Canadian provinces (Ontario, Quebec,
Nova Scotia, New Brunswick, and P.E.I.). This book also briefly summarizes Reynolds’
considerable experience in sampling methods and preservation of earthworm specimens.
Satchell, J.E. (Editor). 1983. Earthworm Ecology. Chapman & Hall Ltd., London. 495 pp.
Papers cover topics under the general headings of: earthworms and organic matter;
earthworm ecology in grassland soils, in cultivated soils, in forest soils, in tropical and arid
soils; earthworms and land reclamation, and soil pollution; utilization of wastes by
earthworm culture; earthworms and microflora; earthworms in food chains; earthworm
evolution and distribution patterns; taxonomy and nomenclature. Refs, after each chapter.
Sims, R.W. In press. A Classification and the Distribution of Earthworms, Suborder
Lumbricina (Haplotaxida: Oligochaeta). Bull. Brit. Mus. Nat. Hist. (Zool.).

Worden, D.D. (Editor). 1981. Workshop on the Role of Earthworms in the Stabilization of
Organic Residues. Vol. II. Bibliography. Beech Leaf Press, Kalamazoo, Michigan.

3036 citations, cumulative author and subject indices. Nematodes and microorganisms that
impact on earthworms are also cited.

Zajonc, I. and J. Cepelak. 1968. Colloquium - Questions on Ecology and Taxonomy of
Earthworms. 149 pp. Institute of Zoology - Institute for Biology and Ecology of Cultural
Plants, Agricultural University, Nitra, Czechoslovakia. 149 pp.

Papers in English, French, German, Russian. Refs, after each presentation.

MOLLUSCA

Cameron, R.A.D. and M. Redfern. 1976. British Land Snails. Synopses of the British Fauna

Soil Invertebrates-Reference Texts

681

(New Series) No. 6. Publ. for Linnean Soc. of London by Academic Press, London. 64 pp.
Keys and notes to the identification of species. Contains sections on morphology, collecting
and preservation.

Leonard, A.B. 1959. Handbook of Gastropods of Kansas. University of Kansas Natural History
Museum, Topeka, Kansas.

Only partially applicable to Canada.

Runham, N.W. and P.J. Hunter. 1970. Terrestrial Slugs. Hutchinson University Library,
London. 185 pp.

Review of biology, ecology and economic importance of slugs. 21 pp. of refs.

CRUSTACEA

Bousfield, E.L. 1982. Amphipoda. pp. 254-293. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 2. McGraw-Hill Book Co.

Crustaceans common in tropical soils.

Richardson, H. 1905. A Monograph on the Isopods of North America. Bull. U.S.N.M. 54:
Reprinted by Antiquariaat Junk, Netherlands 1972.

A classic, and still useful.

Schultz, G.A. 1982. Isopoda. pp. 249-254. In Parker, S.P. (Editor). Synopsis and classification
of living organisms. Vol. 2. McGraw-Hill Book Co.

Up-to-date classification of Woodlice.

Sutton, S.L. 1972, 1980. Woodlice. Ginn & Co., Ltd., London. 144 pp.

An introduction to the biology, ethology, genetics, ecology, and identification of woodlice.
Includes a section on techniques.

Walker, E.M. 1927. The Woodlice or Oniscoidea of Canada (Crustacea, Isopoda). Can. Field -
Nat. 41: 173-179.

Van Name, W.G. 1936. The American Land and Freshwater Isopod Crustacea. Bull. Am.
Mus. Nat. Hist. 71: 1-535.

Useful for identifying Canadian species.

TARDIGRADA

Morgan, C.I. 1982. Tardigrada. pp. 731-739. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. McGraw-Hill Book Co.

Up-to-date classification of group.

MYRIAPODA

Blower, J.G. 1958. British Millipedes (Diplopoda). Linnean Soc. Synopses of the British
Fauna, 1 1: 74 pp.

Mainly taxonomic; useful in E. Canada where most species are European introductions.
Blower, J.G. (Editor). 1974. Myriapoda. Symposia of the Zoological Society of London, 32.
Academic Press, London & New York. 712 pp.

General work, including some taxonomy.

Broleman, H.W. 1932. Chilopodes. Faune France 25. 405 pp.

Useful for the numerous introduced species.

Demange, J.-M. 1981. Les Mille-Pattes, Myriapodes. Societe Nouvelle des Editions Boubee,
Paris. 284 pp.

Morphology, ecology and ethology of class with key to determination to species of myriapods
in France. Excellent figures plus 4 plates in colour and 9 colour photographs.

Eason, E.H. 1964. Centipedes of the British Isles. Frederick Warnc & Co., Ltd., London. 294

pp.

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Has general application; useful in E. Canada where most species are the same or similar.
Edwards, C.A. 1952. A Revision of the British Symphyla. Proc. Zool. Soc. London. 132:
403-439.

Edwards, C.A. 1952. Keys to the Genera of the Symphyla. J. Linn. Soc. Zool. 44: 164-169.
Most useful text on Symphyla.

Hoffman, R.L. 1979. Classification of the Diplopoda. Museum d’Histoire Naturelle, Geneve.
237 pp.

Hoffman, R.L. 1982. Chilopoda. pp. 681-688 In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 2. McGraw-Hill Book Co.

Most recent classification of centipedes.

Hoffman, R.L. 1982. Diplopoda. pp. 689-724. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 2. McGraw-Hill Book Co.

Most recent classification of millipedes.

Kevan, D.K.McE. 1983a. A Preliminary Survey of Known and Potentially Canadian and
Alaskan Centipedes (Chilopoda). Can. J. Zool. 61: 2938-2955.

Present state of knowledge of centipedes, in Canada and Alaska, including economic and
biological aspects. Excellent reference section.

Kevan, D.K.McE. 1983b. A Preliminary Survey of Known and Potentially Canadian and
Alaskan Millipedes (Diplopoda). Can. J. Zool. 61: 2956-2975.

Diplopod fauna of Canada, including a checklist of species known to, and likely to occur, in
Canada. Excellent reference section.

Lewis, J.G.E. 1981. The Biology of Centipedes. Oxford University Press. 476 pp.

Most recent textbook on this subject.

Remy, P.A. 1958. Pauropodes des Etats-Unis d’Amerique et de la Jamaique. Mem Soc. Natn.
Sci. nat. math. Cherbourg 48. 77 pp.

Probably will be useful for some Canadian species.

Scheller, U. 1982. Pauropoda. pp. 724-726. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 2. McGraw-Hill Book Co.

Up-to-date classification of pauropods.

Scheller, U. 1984. Pauropoda (Myriapoda) from Canada. Can. J. Zool. In press.
COLLEMBOLA

Christiansen, K. and P. Bellinger. 1980-1981. The Collembola of North America North of Rio
Grande. Part 1 (1980 Poduridae and Hypogastruridae. Part 2 (1980) Onychiuridae and
Isotomidae. Part 3 (1980) Entomobryidae. Part 4 (1981) Neelidae and Sminthuridae. 1322
pp. Grinnell College, Grinnell.

The current standard work on North American Collembola. Illustrations are numerous, but
finer details are obscured by bad printing. Part 1 contains a good general description of
Collembola morphology and describes several slide preparation techniques. Part 4 contains a
useful morphological glossary and an exhaustive author index.

DeHarveng, L. 1982. Cle de determination des genres de Neanurinae (Collemboles) d’Europe
et de la region Mediterraneene avec description de deux nouveaux genres. Trav. Lab.
Ecobiol. Arthr. Edaph., Toulouse 3(4): 7-13.

Identification keys to the many new Neanurinae genera described from Europe since Gisin
(1960).

Ellis, W.N. & P.F. Bellinger. 1973. An annotated list of generic names of Collembola (Insecta)
and their type species. Mon. Ned. Ent. Ver. 7: 1-74.

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683

A necessary publication for taxonomists in particular.

Fjellberg, A. 1980. Identification keys to Norwegian Collembola. Norsk Entomologisk
Forening, As. 152 pp.

Fairly up to date, illustrated keys covering most of the Nordic species.

Fjellberg, A. 1984. Arctic Collembola 1. Alaskan Collembola of the families Poduridae,
Hypogastruridae, Odontellidae, Brachystomellidae and Neanuridae. Ent. Scand. Suppl. (In
press).

Gisin, H. 1960. Collembolenfauna Europas. Museum d’Histoire Naturelle, Geneve. 312 pp.
Well illustrated identification keys and short description of species. Contains a general
introduction to collembole morphology and identification technique. Although largely out of
date, the book is still the standard work among European collembologists.

Massoud, A. 1967. Monographies des Neanuridae, Collemboles Poduromorphes a pieces
buccales modifiees. Biol. Amer. Austr. CNRS 3: 1-399.

A standard work on the family Neanuridae, but now largely out of date, at least concerning
the European fauna.

Maynard, E.M. 1951. A Monograph of the Collembola or Springtail Insects of New York
State. Comstock Publishing Co. Inc., Ithaca, NY. 339 pp. + 29 pi.

A few biological notes included; nomenclature not up to date.

Rusek, J. 1977. Protura, Collembola, Diplura, Thysanura. Enumeration Insectorum
Bohemoslovakiae. Acta Faun. Ent. Mus. Nat. Pragae 15(4): 9-21.

A check list to the apterygotan species recorded from Czechoslovakia.

Salmon, J.T. 1964. An Index to the Collembola. Bull. Roy. Soc. New Zealand 7(1-2). 644 pp.
Wellington.

An index to world literature on Collembola up to 1962 arranged (1) by author and (2) by
species. Contains also a key to world genera. A very useful book despite numerous minor
errors.

Yosii, R. 1977. Critical check list of the Japanese species of Collembola. Contr. Biol. Lab.
Kyoto University 25(2): 141-170.

In addition to the species list, the paper also provides identification keys to the Japanese
genera.

INSECTS ( OTHER THAN COLLEMBOLA)

Arnett, R.H. Jr., N.M. Downie and H.E. Jaques. 1980. How to Know the Beetles. Brown
Publishing Co., Dubuque. 416 pp.

Chandler, L. 1957. The orders Protura and Diplura in Indiana. Proc. Indiana Acad. Sci. 66:
112-114.

Of very limited use.

Chu, H.F. 1949. How to know the immature insects. Brown Publishing Co., Dubuque. 234 pp.
Lee, K.G. & T.G. Wood. 1971. Termites and Soils. Academic Press, London and New York.
Morgan, C.I. and P.E. King. 1976. British Tardigrades, Tardigrada: Keys and notes for the
identification of the species. Synopsis of the British Fauna No. 9. Academic Press, London.
133 pp.

Important reference source.

Nosek, J. 1973. The European Protura. Museum d’Histoire Naturelle, Geneve. 345 pp.

The most up-to-date work on taxonomy, ecology and distribution. Includes keys for
identification.

Ramazzotti, G. 1972. II Phylum Tardigrada. Mem. 1st Ital. Idrobiol. 28: 1 732.

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An introduction to the world literature on the group.

Smith, L.M. 1960. The family Projapygidae and Anahapygidae (Diplura) in North America.
Ann. Ent. Soc. Am. 53: 575-583.

Sudd, J.M. 1967. An introduction to the behaviour of ants. Edward Arnold Ltd., London.

A useful introduction to ants.

Tuxen, S.L. 1964. The Protura. A revision of the species of the world with keys for
determination. Hermann, Paris. 360 pp.

A major work on Protura.

ARACHNID A (EXCEPT ACARI)

Comstock, J.H. 1940. The Spider Book. Cornell University Press, Ithaca, NY. 727 pp.

A classic spider work, though outdated taxonomically.

Gertsch, W.J. 1978. American Spiders. 2nd ed., Van Nostrand-Reinhold, New York. (1st ed.
1949).

A readable summary for general readers.

Hoff, C.C. 1949. The Pseudoscorpions of Illinois. Illinois Nat. Hist. Surv. Bull. 24: 412-498.

A somewhat dated, but still very useful, introduction to morphology and key.

Hoff, C.C. 1958. List of the Pseudoscorpions of North America North of Mexico. Amer. Mus.
Nov. No. 1875: 1-50.

With a key to genera.

Hoff, C.C. 1959. The Ecology and Distribution of the Pseudoscorpions of North-Central New
Mexico. University of New Mexico Publications in Biology, No. 8. 68 pp.

Contains much general information on biology.

Kaston, B.J. 1948. Spiders of Connecticut. Bull. Conn. Geol. Nat. Hist. Surv. 70: 1-874.

A classic study, very useful for northeastern U.S. and southeastern Canada. Supplement
published in 1977 (Jour. Arachnol. 4: 1-72) updates nomenclature and selected keys.

Kaston, B.J. 1972. How to Know the Spiders. 3rd ed., W. Brown, Dubuque. 272 pp.

Collection techniques, picture keys to orders and families and keys to most common genera;
a good place to start.

Levi, H.W., L.R. Levi and H.S. Zim. 1968. A Guide to Spiders and Their Kin. Golden Press,
New York. 160 pp.

A non-technical, “look-see” guide.

Muchmore, W.B. 1982. Pseudoscorpionida. pp. 96-102. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 2. McGraw-Hill Book Co.

Nelson, S. Jr. 1975. A Systematic Study of Michigan Pseudoscorpionida (Arachnida). Amer.
Midi. Nat. 93:257-301.

Savory, T. 1977. Arachnida. 2nd ed. Academic Press, London, New York. 340 pp.

General introduction to the morphology, physiology, ecology, and taxonomy of the class.
Weygoldt, P. 1969. The Biology of Pseudoscorpions. Harvard University Press, Cambridge,
MA. 145 pp.

Translation from German of 1966. Covers anatomy, physiology, ecology, taxonomy and
techniques. An excellent, readable summary.

ACARI

Baker, E.W. and G.W. Wharton. 1952. An Introduction to Acarology. The Macmillan Co.,
New York. 465 pp.

Largely but not exclusively taxonomic.

Balogh, J. 1972. The Oribatid Genera of the World. Akad. Kiado, Budapest. 188 pp. + 71 pis.

Soil Invertebrates-Reference Texts

685

Balogh, J. and S. Mahunka. 1983. The Soil Mites of the World. 1. Primitive Oribatids of the
Palaearctic Region. Elsevier, Amsterdam. 372 pp.

Keys to Palaearctic species.

Evans, G.O., J.G. Sheals, & D. MacFarlane. 1961. The Terrestrial Acari of the British Isles:
An Introduction to their Morphology, Biology and Classification. British Museum, London.
219 pp.

Evans, G.O. and W.M. Till. 1979. Mesostigmatic Mites of Britain and Ireland (Chelicerata:
Acari-Parasitiformes). Trans. Zool. Soc. Lond. 35: 139-270.

An introduction to their external morphology and classification.

Gilyarov, M.S. (Editor). 1975. A Key to the Soil-inhabiting Mites, Sarcoptiformes. (In
Russian). Nauka, Moscow. 491 pp.

Translation (on fiche) available from Canadian Index of Scientific Translations, Canada
Institute for Scientific and Technical Information, National Research Council of Canada,
Ottawa K1A 0S2. (Translation #4328).

Gilyarov, M.S. 1978. A Key to the Soil-inhabiting Mites, Trombidiformes. (In Russian).
Nauka, Moscow.

Relevant to North American fauna. Translation available as per Gilyarov (1975).
(Translation #4569).

Gilyarov, M.S. and N.G. Bregetova. (Editors). 1977. A Key to the Soil-inhabiting Mites,
Mesostigmata. (In Russian). Nauka, Leningrad. 718 pp.

Very relevant to North American fauna. Translation available as per Gilyarov (1975).
(Translation #4371).

Hughes, T.E. 1959. Mites, or the Acari. University of London Athlone Press, London. 225 pp.
Kethley, J. 1982. Acariformes - Prostigmata, pp. 117-145. In Parker, S.P. (Editor). Synopsis
and classification of living organisms. Vol. 2. McGraw-Hill Book Co.

Krantz, S.W. 1978. A Manual of Acarology. Oregon State University, Corvallis, OR. 509 pp.
2nd ed.

Largely, but not exclusively, taxonomic. The most widely used introductory text on
acarology.

O’Connor, B.M. 1982. Astigmata. pp. 146-169. In Parker, S.P. (Editor). Synopsis and
classification of living organisms. Vol. 2. McGraw-Hill Book Co.

NOTE: A manual dealing with soil organisms in North America is presently being edited by
D.L. Dindal (to be published by J. Wiley & Sons).

STATISTICS, EXPERIMENTAL DESIGN & SAMPLING TECHNIQUES FOR SOIL

ZOOLOGY

Unlike the epigeaic fauna, there are few “models” which have been explicitly established for
soil animal populations. Many epigeaic models have been adapted for soil animal models with
varying degrees of success. The euedaphic fauna is rarely homogeneous even throughout the
small volumes of sampling cores; the fauna varies dramatically with depth and is highly
dependent upon climate factors, soil type, vegetation cover and, structures as roots or
earthworm tunnels, for example. Soil animals often exhibit aggregative behaviour, which,
presumably, is their response to exploitation of food resources (which again are often
aggregated and not evenly or even randomly dispersed in a plot) or reproductive requirements.
There are a large number of statistical texts available for consultation for the more difficult

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statistical analyses, and the wide availability of computers often makes it tempting to get
heavily (perhaps even unnecessarily) involved in this end of the work. An appraisal of statistical
texts is not provided, but listed are a few texts and papers which have an ecological bent, and
that are useful in accessing and comprehending this literature. The assistance of a sympathetic
biometrician for assistance in field experiments should not be underrated.

Marked-capture-recapture techniques, which should have some utility for estimating the
abundance of soil animal populations, have not been widely used in the past for this purpose,
but these techniques might be particularly applicable to earthworms, for example. Some of the
listings given provide numbers and analyses of soil faunal data on which to hang your hat or
against which you may compare your own data.

Gauch, H.G., Jr. ca. 1976-present. The Cornell Ecology Programs Series. Available from
Cornell University, Dept, of Ecology & Systematics, 224 Langmuir Laboratory, Ithaca, NY
14850.

A series of main-frame computer programs (with considerable documentation) for analysing
ecological data. The catalog has been revised and up-dated several times. Program &
documentation are available at nominal cost. Several Canadian universities and government
institutions now provide and support these programs. The programs deal mainly with
ordination and classification of data (particularly useful for large sets of data).

Jeffers, J.N.R. Statistical Checklists. Nos. 1,2, & 3 (Design of Experiments, Sampling &
Modelling, respectively). Institute of Terristrial Ecology, Cambridge, UK.

Lists of questions to ask of yourself and your experiments. Thought-provoking and helpful.
Jeffers, J.N.R. 1978. An Introduction to Systems Analysis with Ecological Applications.
University Park Press, Baltimore.

For those of you into ecological modelling - this is a very readable account with lots of
worked examples.

Lewis, T. and L.R. Taylor. 1967. Introduction to Experimental Ecology. Academic Press.

A relatively basic approach to quantitative ecology, but a treasury of techniques, graphs,
lists and analytical methodology for ecologists.

Macfadyen, A. 1963. Animal Ecology. 2nd Ed. Pitman & Sons.

Less mathematical approach than Southwood or Taylor & Lewis, but Macfadyen’s
credentials as a soil zoologist mean that there are plenty of illustrative examples from soil
ecology.

Petersen, H. (Editor). 1982. Quantitative ecology of microfungi and animals in soil and litter.
Oikos 39: 388-422.

Extensive tabulations and comparisons of various components of the soil fauna for various
global biomes and their impact on decomposition and soil processes. An invaluable aid for
comparison purposes and highlighting the many deficiencies.

Phillipson, J. (Editor). 1971. Methods of Study in Quantitative Soil Ecology. Blackwell
Scientific Publications. IBP Handbook No. 18.

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A comprehensive account of various soil animal sampling techniques. An indispensable
handbook for this type of work. Phillipson’s final chapter on “Other Arthropods” is brief but
helpful entree to many of these neglected taxa ( e.g ., Tardigrada).

Pielou, E.C. 1976. Mathematical Ecology. John Wiley & Sons.

A very mathematical (not statistical) approach to ecology. Heavy going in many places. A
source book for the mathematically-minded ecologist.

Seber, G.A.F. 1973. The Estimation of Animal Abundance and Related Parameters. Hafner
Press, New York.

Very heavy going in places, but a comprehensive review of various sampling and
marked-capture-recapture techniques. The final chapter is an excellent summary of
methods discussed in the book.

Southwood, T.R.E. 1978. Ecological Methods. 2nd Ed. Chapman & Hall.

A somewhat more sophisticated approach than Lewis & Taylor, but otherwise comparable.
Excellent bibliographies.

Wallwork, J.A. 1970. Ecology of Soil Animals. McGraw-Hill.

The best single “read” on the details of soil ecology from concepts to specimen preservation.
Good bibliographies. There is a brief section on extraction techniques for soil animals.
References are now somewhat dated.

Wallwork, J.A. 1976. The Distribution and Diversity of Soil Fauna. Academic Press.

Lots of relevant goodies here. Written by a soil ecologist specializing in soil mites. Chapter 2
is a “quickie” review of statistical and measuring techniques and ecological concepts. Highly
recommended.

Quaest. Ent., 1985, 21 (4)

688

Index

INDEX

Abies alba , 508

Acacia aneura, 579

Acanthoceridae, 576

Acanthodrilidae, 518

Acarina see mites, 506

Acer saccharum Marsh, 588

Aclopinae, 576

Adela spp., 502, 510

Aenigmopus alatus , 551

agricultural practices, effects on soil, 638

agriculture, definition, 637

agriculture as multi-story polyculture, 639

agriculture, sustainable, 639

agroecosystems, indicators of distress, 640

agroecosystems, sustainable, 639

Ah horizons, 475

amoebae, 619

ant-lions, 573

Anthribidae, 574

ants, 506, 572, 577

ants as indicators of soil conditions, 641
Anurophorus , 563
Aphodiinae, 576

Aporrectodea tuberculata (Eisen), 591
Arachnida, 524, 582
arachnids, soil, phylogeny, 534
supraspecific classification, 534
systematics, 525
Artemetopoidea, 574
Arthronota, 534
Arthroptyctima, 534
Astigmata, 525
Attini, 579
bacterial gel, 619
Ballophilidae, 552
bark beetles, 502
Berlese Oribatei, 500
Bibionidae, 510

Bilobella aurantiaca Caroli, 565
Blattodea, 573
booklice, 573
Bostrychoidea, 574
Bourletiella lutea , 508
Bourletiella spp., 507
Brachycybe, 551

BSEI, 610
bugs, 573

Buprestidae, 502, 574
Buprestoidea, 574
butterflies, 573
Byrrhoidea, 574
caddisflies, 572
Camponotus intrepidus, 577
Cantharidae, 574
Cantharoidea, 574
Carabidae, 573-574, 576
Caraboidea, 574
centipedes, 524
Cephalotoma grandiceps, 507
Cerambycidae, 502, 574, 576
Cetoniidae, 502
Cetoniinae, 576-577
Chelodesmidae, 547
Chernozem, 605
Chilopoda, 544, 548-549
checklists, 548
regional surveys, 548
Chironomidae, 510
Chordeumatida, 547
Chrysomelidae, 574, 576
Chrysomeloidea, 574
Cleidogonidae, 547
Cleridae, 574
Cleroidea, 574
Coccinellidae, 574
cockroaches, 573

Coleoptera, 502, 511, 572-573, 575, 582
larvae, 502

Collembola, 500, 502-50}, 507-509, 524,
559

aggregations on snow, 561
anhydrobiosis, 561
behaviour, 561
chaetotaxy, 564
cyclomorphosis, 564
cytogenetics and physiology, 564
dependence on moisture, 561
ecomorphosis, 563
effects of pollution and human
activities, 562

Index

689

epitoky, 563
fecal pellets, 479
feeding and nutrition, 559
reproductive biology, 560
response to water supply, 652
Conotylidae, 547
coprophagy, 589
Coprosphere, 593
crickets, 506, 573
Cryptocephalinae, 576
Cryptostigmata, 524
CTEM, 610
Ctenacaridae, 528
Cucujoidea, 574-575
Cupedoidea, 574
Curculionidae, 502, 574, 576
Curculionoidea, 574
Dascilloidea, 574

Dendrobaena octaedra (Savigny), 591
Dendrobaena rubida, 503, 505, 513
Dendrodrilus rubidus (Savigny), 591
Dermaptera see earwigs, 573
Dermestoidea, 574
Diplocardia, 5 1 8

Diplopoda, 500, 502, 544-545, 547-549
checklists, 548
regional surveys, 548
Diptera, 573

larvae, 477, 499, 502-503
Dragonflies, 572
Dryopidae, 511
Dryopoidea, 574
Dry ops rudolfi, 499, 510-512
dung beetles, 506
Dynastidae, 502
Dynastinae, 576
earthworm casts, 478
earthworm fecal pellets, 478
earthworms, 477, 500, 502-503, 517, 588
mineralization of carbohydrates, 621
earthworms and soil development, 474
earthworms, anecic, 500, 503, 506
earthworms, endogeic, 500, 503, 505
earthworms, epigeic, 500, 502, 505
earwigs, 573

Edaphophyllosphere, 593

EDXRA, 610
Eisenia foetida, 503, 513
Eisenia lucens, 513
Elateridae, 503, 574, 576
Elateroidea, 574
Embioptera, 572
Enchytraeidae, 477, 507, 510
Endeostigmata, 528
Entomobryidae, 565
Enzymes, 623
EPMA, 610
Euarthronota, 534
Eucinetoidea, 574
Eumolpinae, 576
Eupelops, 526
Eupodidae, 527, 530, 533
fecal pellets, 479, 588, 591
construction units, 599
soil microstructure, 599
fecal rain, 589
Fensterfrass, 500
fertilizers and pestcides, 637
flagellates, 619
fleas, 572
flies, 573

Folsomia spp., 503, 507
food web, decomposer, 646
Formica cinerea, 580
Formicidae see ants, 572
Fraxinus spp., 588
Friesea spp., 507
fungi and bacteria, 622
Gaeumannomyces, 620
Geolycosa, 590
geophagy, 589
Geophilomorpha, 548-549
Geotrupidae, 576
Glomeris spp., 500
Glyceria maxima , 5 1 2
Gomphodesmidae, 547
Grandjeanicus, 528
grasshoppers, 573
Grylloblattodea, 572
Haeckel’s biogenetical law, 502
Harpagophoridae, 547
Harpaphe haydeniana , 552

Quaest. Ent., 1985, 21 (4)

690

Index

harvestmen, 524
Hemiptera see bugs, 573
Heterostigmata, 530
Hexapoda, 582
Histeridae, 574
Histeroidea, 574
history of soil zoology, 369-472
Humic fecal pellets, 478
humon, 475
Hybosorinae, 576
Hydrophilidae, 574
Hydrophiloidea, 574
Hymenoptera, 506, 572
Hypochtonius sp., 508
Hypogastrura, pheromones, 561
phototaxis, 561

Hypogastrura socialis (Uzel), 560
Hypogastrura spp., 507
Hypogastrura tullbergi (Schaffer), 564
Hypogastruridae, 563
IMMA, 61 1

Iridomyme x purpureus, 579
Isopoda, 500, 502
isopods, 588
Isoptera, 572, 578, 582
Isotoma , 560

Isotoma nivea Schaffer, 564
Isotoma spp., 507

Isotoma tigrina ( olivacea auct.), 563

Isotomidae, 563

Julida, 548

Julus, 544

Julus spp., 500

kingfisher, belted, 590

Krotovinas, 605

lace wings, 573

LAMNA, 61 1

leaf litter, decomposition of, 614
Lepidoptera, 573
lice, 572

Lithobiomorpha, 548
Lithobius , 544

Litter disintegration in a forest soil, 503,
588

Lochfrass, 500
Lucanidae, 502, 576

Lumbricidae, 503, 513
Lumbricus rubellus Hoffmeister, 591
Lumbricus terrestris L., 588, 591
Lycidae, 574
Lycoridae, 500, 502, 510
Lymexyloidea, 574
macroarthropod droppings, 503
macroarthropods, 503
Macrotermes species, 599
Mantodea, 572

Mecoptera see scorpion flies, 572
Megaceryle a. alycon, 590
Megalothorax minimus , 507
Megascolecidae, 518
Meloidae, 574
Melolontha spp., 510
Melolonthinae, 576-577
Melyridae, 574
Membrane systems, 623
Meranoplus , 581

Merchant-Crossley extractor, 669
Mesaphorura spp., 502, 507
Mesostigmata, 524

edaphic members, 533
metavughs, mammilated, 599, 602
Micranurida spp., 507
microbial gums, 619
microbial polysaccharides, 619
microbial slimes, 620
microcosms, 620
microfabrics, spongy, 474
microstructure formation, processes, 499
mites, development, 529
fecalpellets, 506
fungivorous, 524
fungivorous oribatid, 524
fungivorous prostigmatid, 524
gut enzymes, 526
gut microflora, 526
mycophagy, 524, 527
nutrition, 525

population dynamics, 525, 529
predation, 524
saprophagy, 524, 527
sarcoptiform, 528

mites, mycophagous soil, taxonomy and

Index

691

monographs, 533
mites, oribatid, cannibalism, 525
nutrition, 525
mites, phthiracarid, 500
mites, saprophagous, comminution, 530
mites, saprophagous soil, taxonomy and
monographs, 533
modexi, 475
Monera, 526
mor humus, 474
moths, 573
mucigel, 619
Mull, 500
mull humus, 474
Mycetophilidae, 500, 502, 510
Mycoridae, 510
Myremecia pilosula, 577
Myremeleontidae see ant-lions, 573
Myriapoda, 582
diversity, 545
knowledge, 544
morphology, 549
systematics, 544
Myriapodology, history, 544
Neanura spp., 507
Neanuridae, 565
Necrosphere, 593
Nematocera, 500
larvae, 502

nematodes, bacteriophagous, 524
nematodes, predatory, as soil indicators,
641

Neuroptera, 573
Nidusphere, 593
Nitidulidae, 574

nova (Oudemans), Oppiella, 527
Octolasion tyrtaeum (Savigny), 591
Odonata, 572
Odontopygidae, 547
Ommatoiulus more let i, 550
Oniscus asellus L., 588
Onychiuridae, 503

Onychiurus armatus , contamination with
lead and copper, 562
Onychiurus spp., 503, 507
Opisthogoneata, 544

Oppiella nova (Oudemans), 527
Orchesella cincta , 509
Orchesella spp., 507
Organics of submicron size, 620
organo-clay complexes, 475
Oribatei, 500, 506-508
Oribatida, 524
Oribatidae, K-selection, 530
Orthoptera, 573
owl, burrowing, 590
Oxydesmidae, 547
Paradoxosomatidae, 547, 549
Parajulidae, 548, 551
parasitic bacteria, 619
Paratullbergia callipygos, 507
Passalidae, 502, 576
Passalus sp., 502
Pauropoda, 544-545, 549
pedology, interface with biology, 596
pedoturbation, faunal, 605
Phasmatodea, 572
Pheretima, 518
Phgmephoridae, 527
Phthiracaridae, 507
xylophagy, 526
Phthiracarus , 526
Phthiraptera, 572
pitchmoder rendzina, 507
plant tissues in soil, bacterial
decomposition, 614
Platynothrus poltifer (Koch), 529
poltifer (Koch), Platynothrus, 529
Polydesmida, 547
polysaccharides, 622
polysaccharides, extracellular, 622
predators in soil, 641
preying mantises, 572
Proctostephanus , 563
Progoneata, 544
Prostigmata, 524, 527
Protacarus , 528
Protacarus crani , 528
Pselaphidae, 574
pseudoscorpions, 524
Pseudosinella spp., 507
Pterygota, 572-573

Quaest. Ent., 1985, 21 (4)

692

Index

Pygmephoridae, 533
rabbit, cottontail, 591
rendzina soils, 506
Rhiscosomididae, 547
Rhisotritia minima, 500
rhizoflora, 619

rhizosphere microflora, functions of, 619

Riparia r. riparia, 590

root exudates, 612

root mucilages, 6 1 2

root tissues, 612

Russian Chernozems, 474

Rutelinae, 576-577

Scarabaeidae, 574, 576-577

Scarabaeinae, 576

Scarabaeoidea, 574, 576

Scolopendromorpha, 548

Scolytidae, 502

scorpion flies, 572

Scutacaridae, 527, 533

Scutigeromorpha, 548

SEM, 610

Sigmoria, 547

SIMS, 611

Siphonaptera, 572

Sminthurus spp., 507

S mint hums viridis, 507

Soil, drainage, 518

soil aeration, 5 1 8

soil animals, 587

Beneficial effects, 638
microcommunicies, 587
microcommunities, 650
taxonomic challenge, 649
soil arthropod densities, predation in
regulating, 524

soil fabrics, decomposition of organic
matter, 602

faunal influence, 587, 596
organic luminae, 598
transformation, 652
ultrahistochemical analysis, 628
soil fabrics, decomposition of, 589
organic matter, 590
soil fabrics, enaulic, 475
soil fabrics, granic, 476

soil fabrics, granoidic, 476
soil fabrics, granoidic porphyric, 476
soil fabrics, monic, 475
soil fabrics, porphyric, 476
soil fauna, comminution of plant debris,
652

regulation of soil systems, 653
relationship with structure, 646
soil fauna as a resource, 639
soil fauna coenoses, 499
soil homogenization, land reclamation, 605
soil invertebrates, dissection, 673
Soil litter, 587
soil litter components, 588
soil materials, f-matrix, 475
f-member, 475
soil mesofauna, 503
soil microfabrics, 474
soil micromorphology, 498
application, 661
collecting samples, 658
definition, 657
preparation of samples, 659
sampling, 658
techniques for study, 657
soil microstructure formation, 473, 589
processes, 500, 507
soil nematode densities, predation in
regulating, 524
soil pests, 638

soil systems, structure and function, 653
soil texture, trophic intercations, 648
Soil zoology, history of, 369-472
soils, amorphous and granular materials,
submicron size, 621
arthropod moder, 500
carbohydrates, 621
cross hatching, 598
decomposers, 499, 506
development of humus forms, 502
fibrous or lamellate materials,
submicron size, 621
humic substances, 622
lamellar fabrics, 598
lenticular fabric, 598
materials in animals and their remains.

Index

693

614

mesh fabrics, 599
micro fabrics, 596
microarthropod droppings, 503
microarthropod moder, 500
microhabitats, 650
physical dimensions, 61 1
Prairie Parkland Region, 479
predator-prey interactions, 648
secondary sources of organic matter,
614

sources of organic matter, 612
sources of organic matter in soils, 623
thin sections, 498, 661
ultracytochemistry, 611, 628
soils fabrics, physical and chemical
stabilization, 611

soils, biochemical properties of, 61 1
soils, materials in aerial organs, 612
soils, pedzolic, termites, 599
soils, regional, Boreal Forest Region, 478
Forest-Tundra Transition Region, 477
North American Prairies, 474
Northern Tundra Region, 476
Southern Tundra Region, 476
soils, study of, back scattered electron
detection, 610

conventional-transmission electron
microscopy, 610

electron probe microanalysis, 610
energy dispersive X-ray analysis, 610
ion microprobe mass analysis, 611
laser microprobe mass analysis, 61 1
philosophical challenges, 649
physical properties, 610-61 1
scanning electron microscopy, 610
scanning-transmission electron
microscopy, 610

secondary ion mass spectrometry, 61 1
wavelength dispersive X-ray analysis,
610

Speotyto cunicularia, 590
Sphaerioidea, 574
spider predation, 524
spiders, 524
Spirobolida, 548

Spirobolidae, 548
Spirobolus, 544
Spirostreptida, 547
Spirostreptidae, 547
Staphylinidae, 574
Staphylinoidea, 574
Steganacarus magnus, 506
STEM, 610
Stemmiulida, 550
stick insects, 572
Strepsiptera, 572
succession of humus forms, 500
swallows, bank, 590
cliff, 590

Symphyla, 544-545, 549
Tarsonemidae, 527, 533
Tectocepheus velatus (Michael), 527
Tenebrionidae, 574-575
termite mounds, microstructure, 596
termites, 506, 572, 578, 582
plant decomposition, 604
Terpnacarus, 528
thrips, 573
Tingupidae, 547
Tipulidae, 502, 510
Tomocerus minor , 509
Tomocerus spp., 507
Trichoptera, 572
Trogidae, 576
Tullbergiinae, 503
Tydeidae, 524, 527, 530, 533
Typhloblaniulus lorifer , 551
Tyrophagous putrescentiae , 530
Uropodina, 527
Valginae, 576

velatus (Michael), Tectocepheus , 527

Vermiborolls, 605

Vermisol, 605

Vermudolls, 605

Vermustolls, 605

Vertagopus sp., sun orientation in, 561

Vertagopus westerlundi , 561

viruses, 619

WDXPA, 610

wireworms, 503

Xenylla maritima , 561

Quaest. Ent., 1985,21 (4)

694

Index

xeric protorendzina, 507
Xystodesmidae, 547
zooedaphon, 498
Zoraptera, 572

Index to Volume 21

695

INDEX TO VOLUME 21

( Progateritina ) bicolor Drury, Galerita ,
351

Acacia greggii, 236
acanthobia delicatula vittata Forel,
Pseudomyrma, 238
Achworth, A.C.,

see Morgan, A.V., 334
Allen, R.T., 352-353, 364
Ambrosia artemisiifolia , 235
Anacardium, 239
Ancystroglossus Chaudoir, 360
Ancystroglossus dimidiaticornis
Chaudoir, 351

Ancystroglossus new species, 351
Ancystroglossus ovalipennis Reichardt,
351

Andropogon , 235, 237-238
Anthrax spp., 321
apache Creighton, Pseudomyrmex ,
215-216, 218, 229-230, 234-235
arcticum Malloch, Simulium, 176-177,
180-182, 189-190, 192, 196-197,
204-206

Arctostaphylos manzanita , 230
Ardisia revoluta, 239
Arnason, A.P.,

see Rempel, J.G., 176
attelaboides Fabricius, Galerita , 352
auduboni LeConte, Cicindela, 328
Avicennia germinans, 228, 239

Baccharis , 225, 236
Baccharis halimifolia, 228, 239
Bacillus thuringiensis , 206
Bailey, I.W.,

56* Wheeler, W.M., 225-226, 235
Ball, G.E., ,351-354, 364-365

see also Allen, R.T., 352-353, 364
balli Reichardt, Galerita , 352
Basilewsky, P, 351

beetles, carrion, 247-317
Bidens, 235

bimaculatus MacLeay, Planetes , 351
Bombiliidae, 321

boucardi Chaudoir, Galerita , 352

brunnea F. Smith, Pseudomyrma ,
231-232

brunneus F. Smith, Pseudomyrmex ,
215-217, 231-232, 240, 242
Buren, W.F.,

see Whitcomb, W.H., 225
Callicarba, 235

capperi Forel, Pseudomyrmex, 215
carrion beetles, 247-317
Carroll, J.F.,

see Whitcomb, W.H., 225
Carya, 232
Carya floridana, 239
Cicindela auduboni LeConte, 328
Cicindela cinctipennis LeConte, 332
Cicindela cuprescens LeConte, 332-333
Cicindela decemnotata Klug, 323, 329,
335

Cicindela duodecimguttata Dejean,
320-321,324-325,335
Cicindela formosa formosa Say, 320
Cicindela formosa gibsoni Brown, 320,
328

Cicindela formosa Say, 320-321, 323,
328, 330, 335

Cicindela fulgida fulgida Say, 330
Cicindela fulgida Say, 324, 330, 335
Cicindela guttifera LeConte, 325
Cicindela hirticollis ponderosa Thoms.,
326

Cicindela hirticollis Say, 320, 323-326,
335

Cicindela hyperborea LeConte, 326
Cicindela imperfecta LeConte, 332
Cicindela kirbyi LeConte, 331
Cicindela knausi Leng, 332
Cicindela lengi Horn, 320-321, 323, 330
Cicindela lengi lengi Horn, 330
Cicindela lengi versuta Casey, 330, 335
Cicindela lepida Dejean, 319-323, 333,
335

Cicindela limbalis Klug, 329
Cicindela limbata hyperborea LeConte,
323, 326-327, 334
Cicindela limbata limbata Say, 326

Quaest. Ent., 1985,21 (3)

696

Index to Volume 21

Cicindela limbata nympha Casey,
320-323, 326, 330
Cicindela limbata Say, 324, 326
Cicindela longilabris, 327
Cicindela longilabris longilabris Say, 327
Cicindela longilabris Say, 320, 323,
327-328

Cicindela montana LeConte, 327
Cicindela nebraskana LeConte, 320, 322,

327- 328

Cicindela nebraskana nebraskana
LeConte, 328

Cicindela nevadica knausi Leng, 332-333
Cicindela nevadica LeConte, 321, 323,

332.335

Cicindela nympha Casey, 326
Cicindela oregona guttifera LeConte,
324-326, 334

Cicindela oregona LeConte, 320,

324-325, 332, 334

Cicindela oregona maricopa Leng, 334
Cicindela oregona navajoensis Van Dyke,
334

Cicindela oregona oregona LeConte,
324-325, 334-335
Cicindela puarpurea LeConte, 328
Cicindela punctulata Oliver, 321-322, 331
Cicindela puntulata punctulata Oliver,

331

Cicindela purpurea auduboni LeConte,

329

Cicindela purpurea LeConte, 323,

328- 329

Cicindela purpurea purpurea LeConte,
320, 329

Cicindela repanda Dejean, 320-321,
323-326, 335

Cicindela scutellaris Say, 320-321, 324,

330

Cicindela splendida Hentz, 329
Cicindela splendida limbalis Klug, 324,
328-329, 335

Cicindela terricola cinctipennis LeConte,
322, 332, 335

Cicindela terricola imperfecta LeConte,

322.332.335

Cicindela terricola Say, 321-322, 332
Cicindela togata La Ferte, 333
Cicindela tranquebarica borealis
Harrington, 331

Cicindela tranquebarica Herbst, 324, 331,
333, 335

Cicindela tranquebarica kirbyi LeConte,
331

Cicindela versuta Casey, 330
cinctipennis LeConte, Cicindela , 332
Cladium, 225, 228, 232, 240
Cladium jamaicense, 235, 239
Conocarpus erectus, 239
Conostegia , 232

Creighton, W. S., 209, 225-227, 229, 232,
235,239

Criddle, N., 320, 324-326, 333
cubaensis Forel, Pseudomyrmex,

214- 215, 217-218, 226-228, 241
cuprescens LeConte, Cicindela , 332-333
decemnotata Klug, Cicindela , 323, 329,

335

delicatula capperi Forel, Pseudomyrma ,
238

delicatula Forel, Pseudomyrma , 238
delicatula panamensis Forel,
Pseudomyrma , 238

delicatulus Forel, Pseudomyrmex , 215
Denmark, H.A.,

see Whitcomb, W.H., 225
dimidiaticornis Chaudoir,
Ancystroglossus , 351
duodecimguttata Dejean, Cicindela ,
320-321,324-325,335
ejecta F. Smith, Pseudomyrma , 23 1
ejectus F. Smith, Pseudomyrmex ,

215- 217, 219, 231-234, 240, 242
elongata cubaensis Forel, Pseudomyrma ,

226

elongata Mayr, Pseudomyrma , 226-227
elongatus Mayr, Pseudomyrmex ,

214-215, 217, 219, 226-228, 241-242
Environment Canada, ,181
Enzmann, E.V., 231
Erwin, T.L., 364

Eunostus Castlenau, 359, 363, 365

Index to Volume 21

697

Eunostus herrarensis Alluaud, 351
Eunostps new species, 351
Eunostus vuilloti Alluaud, 351

Faunal Influences on Soil Structure
(Symposium), 371.1

flavidula delicatula Forel, Pseudomyrma,
235, 238-239
Flint, B., 334
Forel, A., 226-227, 238
formosa formosa Say, Cicindela, 320
formosa gibsoni Brown, Cicindela , 320,
328

formosa Say, Cicindela , 320-321, 323,
328,330,335
Fraxinus gall, 230

Fredeen, F.J.H., 176, 180, 184, 189-190,
205-206

Freitag, R., 322, 324-325, 334
Frenzel, B., 334
Frey, D.G.,

see Wright, H.E., 334
fulgida fulgida Say, Cicindela , 330
fulgida Say, Cicindela , 324, 330, 335
Galerita (Progaleritina) bicolor Drury,

351

Galerita ( sensu lato ), 351, 354
Galerita attelaboides Fabricius, 352
Galerita balli Reichardt, 352
Galerita boucardi Chaudoir, 352
Galerita Fabricius, 354, 365
Galerita mexicana Chaudoir, 352
Galerita perrieri Fairmaire, 351-352
Galerita procera Gerstaecker, 352
Galerita ruficollis Dejean, 352
Galerita sulcipennis Reichardt, 351-352
Galeritin’a, 354, 359-360, 363
Galeritini, 351-352, 354, 360, 364
Galeritiola Jeannel, 354
Gibbs, D.F.,

see Hinton, H.E., 364
Gliricidia sepium , 228, 239
Gossypium thurberi , 236
Goulet, H., 364

gracilis mexicana Roger, Pseudomyrma ,
225

Graves, R.C., 320

guttifera LeConte, Cicindela , 325
Hamilton, C.C., 320, 324, 326, 332-333
Harris, R.A., 21 1, 354
Hatch, M.H., 327
Helicteres , 228

herrarensis Alluaud, Eunostus , 351
Heterotheca , 238, 240
Heterotheca subaxillaris, 236, 238
Hibiscus tiliaceus, 239
Hilchie, G.J.,

see Ryan, J.K., 192
Hinton, H.E., 352, 364
hirticollis ponder os a Thoms., Cicindela ,
326

hirticollis Say, Cicindela , 320, 323-326,
335

hyperborea LeConte, Cicindela, 326

Hyptis emoryi, 236

imperfecta LeConte, Cicindela, 332

Inga, 228

Iva ciliata, 236

Jeannel, R., 352

Kavanaugh, D.H., 334

Kempf, W.W., 214

kirbyi LeConte, Cicindela, 331

knausi Leng, Cicindela, 332

Laguncularia racemosa, 228, 239

Lavingne, R.J., 320

lengi Horn, Cicindela, 320-321, 323, 330
lengi lengi Horn, Cicindela, 330
lengi versuta Casey, Cicindela, 330, 335
lepida Dejean, Cicindela, 319-323, 333,
335

leptosus sp. nov., Pseudomyrmex, 215,
218-219, 233-235,242
limbalis Klug, Cicindela, 329
limbata hyperborea LeConte, Cicindela,
323, 326-327, 334
limbata limbata Say, Cicindela, 326
limbata nympha Casey, Cicindela ,
320-323, 326, 330
limbata Say, Cicindela, 324, 326
Lindroth,C.H., 351-353, 364
longilabris longilabris Say, Cicindela , 327
longilabris Say, Cicindela, 320, 323,
327-328

Quaest. Ent., 1985, 21 (3)

698

Index to Volume 21

luggeri Nicholson and Mickel, Simulium,
176-177, 180-182, 184-185, 187,
189-193, 196-197, 199, 203-206
Mann, W.M., , 226, 228, 235

see Wheeler, W.M., 226, 228, 239
marginipennis Latreille, Trichognathus,
351

Martin, P.S., 334
Matthews, J.V. jr.,

see also Morgan, A.V., 334
Matthews,, J.V. jr., 334
Mayr, G., 227
Melia azedarach , 231, 236
meridionale Riley, Simulium , 177, 182
Methocha Latreille, 321
Met opium toxiferum, 239
mexicana Chaudoir, Galerita, 352
mexicanus Roger, Pseudomyrmex ,
214-216,218, 225-226
Mickel, C.E.,

see Nicholson, H.P., 187
Mitchell, J.D., 225, 232, 235
montana LeConte, Cicindela, 327
Morgan, A.V., 334
Moss, E.H., 335

nebraskana LeConte, Cicindela , 320, 322,
327-328

nebraskana nebraskana LeConte,
Cicindela , 328
Nectandra coriacea, 239
nevadica knausi Leng, Cicindela , 332-333
nevadica LeConte, Cicindela , 321, 323,
332,335

new species, Ancystroglossus, 351
new species, Eunostus, 351
Nicholson, H.P., 187
nigritus Enzmann, Pseudomyrmex , 215
Nimmo, A.P.,

see Ball, G.E., 351
Nursall, J.R.,

see Paterson, C.G., 177
nympha Casey, Cicindela , 326
oregona guttifera LeConte, Cicindela ,
324-326, 334

oregona LeConte, Cicindela , 320,

324-325, 332, 334

oregona maricopa Leng, Cicindela , 334
oregona navajoensis Van Dyke, Cicindela ,
334

oregona oregona LeConte, Cicindela ,
324-325, 334-335

ovalipennis Reichardt, Ancystroglossus ,
351

pallida F. Smith, Pseudomyrma, 229,
234-235, 237, 239

pallidus F. Smith, Pseudomyrmex , 209,
215-216, 218-219, 228-229, 234-236,
238-240, 242

panamensis Forel, Pseudomyrmex, 215

Paterson, C.G., 177

pendleburyi Andrewes, Planetes, 351

Peperomia , 225

perrieri Fairmaire, Galerita , 351-352
peruvianus Wheeler, Pseudomyrmex, 232
Pierce, W.D.,

see Mitchell, J.D., 225, 232, 235
Pinus attenuata cone, 230
Planetes bimaculatus MacLeay, 351
Planetes MacLeay, 356, 360, 363, 365
Planetes pendleburyi Andrewes, 351
Planetes ruficollis Nietner, 351
Planetina, 354, 356, 360
Populus sp., 230

procera Gerstaecker, Galerita, 352
Progaleritina Jeannel, 351, 360
Prosopis, 225, 228, 232
Prosopis glandulosa, 230
Prosopis sp., 230
Prunus, 236

Pseudomyrma acanthobia delicatula
vittata Forel, 238

Pseudomyrma brunnea F. Smith, 231-232
Pseudomyrma delicatula capperi Forel,
238

Pseudomyrma delicatula Forel, 238
Pseudomyrma delicatula panamensis
Forel, 238

Pseudomyrma ejecta F. Smith, 231
Pseudomyrma elongata cubaensis Forel,
226

Pseudomyrma elongata Mayr, 226-227

Index to Volume 21

699

Pseudomyrma flavidula delicatula Forel,
235, 238-239

Pseudomyrma gracilis mexicana Roger,
225

Pseudomyrma pallida F. Smith, 229,
234-235, 237, 239

Pseudomyrma simplex F. Smith, 238
Pseudomyrmex apache Creighton,
215-216, 218, 229-230, 234-235
Pseudomyrmex brunneus F. Smith,
215-217,231-232, 240, 242
Pseudomyrmex capperi Forel, 215
Pseudomyrmex cubaensis Forel, 214-215,

217- 218,226-228,241
Pseudomyrmex delicatulus Forel, 215
Pseudomyrmex ejectus F. Smith,

215-217, 219, 231-234, 240, 242
Pseudomyrmex elongatus Mayr, 214-215,
217,219, 226-228, 241-242
Pseudomyrmex leptosus sp. nov., 215,

218- 219, 233-235,242
Pseudomyrmex Lund, 209-210, 212, 214,

225, 235, 239-242
Pseudomyrmex mexicanus Roger,

214- 216,218,225-226
Pseudomyrmex nigritus Enzmann, 215
Pseudomyrmex pallidus F. Smith, 209,

215- 216, 218-219, 228-229, 234-236,
238-240, 242

Pseudomyrmex panamensis Forel, 215
Pseudomyrmex peruvianus Wheeler, 232
Pseudomyrmex seminole sp. nov.,
215-216, 218-219, 229, 234-235,
238-240, 242

Pseudomyrmex simplex F. Smith,
215-217, 219, 234-235, 239-240
Pseudomyrmex subater Wheeler &

Mann, 227-228

Pseudomyrmex tandem Forel, 214, 228
Pseudomyrmex vittatus Forel, 215
PseudomyrmexbrunneusF. Smith, 218
Psorealea lanceolata Pursh, 330
Ptelea trifoliata, 236
puarpurea LeConte, Cicindela, 328
punctulata Oliver, Cicindela, 321-322,

331

puntulata punctulata Oliver, Cicindela ,
331

purpurea auduboni LeConte, Cicindela,

329

purpurea LeConte, Cicindela, 323,
328-329

purpurea purpurea LeConte, Cicindela,
320, 329
Quercus, 232
Quercus arizonica, 230
Quercus chrysolepis, 230
Quercus emoryi, 230, 236
Quercus fusiformis, 228, 230
Quercus grisea, 230
Quercus oblongifolia, 229-230, 236
Quercus santaclarensis, 230
Quercus turvinella, 230
Quercus virginiana, 228
Quercus wislizeni, 230
Reichardt, H., 351,365
Rempel, J.G., 176

repanda Dejean, Cicindela, 320-321,
323-326, 335
Rhizophora, 225
Rhizophora mangle, 228
Rhus, 232
Roger, J., 225
Ross, H.E., 334

ruficollis Dejean, Galerita, 352
ruficollis Nietner, Planetes, 351
Ryan, J.K., 192
Sale, P.F.,241
Salix, 225

Saskatchewan Agriculture,,, , 196-197
scutellaris Say, Cicindela, 320-321, 324,

330

seminole sp. nov., Pseudomyrmex,
215-216, 218-219, 229, 234-235,
238-240, 242

Shelford, V.E., 320, 332-333
Shewed, G.E., 180
Shpeley, D.,

Ball, G.E., 364

simplex F. Smith, Pseudomyrma, 238
simplex F. Smith, Pseudomyrmex,
215-217, 219, 234-235, 239-240

Quaest. Ent., 1985, 21 (4)

700

Index to Volume 21

Simulium arcticum Malloch, 176-177,

180- 182, 189-190, 192, 196-197,
204-206

Simulium luggeri Nicholson and Mickel,
176-177, 180-182, 184-185, 187,
189-193, 196-197, 199, 203-206
Simulium meridionale Riley, 177, 182
Simulium vittatum Zetterstedt, 177,

181- 182

Smith, D.R., 209

Smith, F., 231,234, 238

Soil Structure, Faunal Influences on

(Symposium), 371.1

Spilanthes , 232

splendida Hentz, Cicindela , 329
splendida limbalis Klug, Cicindela, 324,
328-329, 335
Spurr, D.T.,

see Fredeen, F.J.H., 180
Statistics Canada,, , 197
subater Wheeler & Mann,

Pseudomyrmex , 227-228
sulcipennis Reichardt, Galerita, 351-352
Swan, L.A., 321
Swietenia mahagoni, 239
tandem Forel, Pseudomyrmex, 214, 228
Terminalia catappa, 239
terricola cinctipennis LeConte, Cicindela ,
322, 332, 335

terricola imperfecta LeConte, Cicindela,
322, 332, 335

terricola Say, Cicindela, 321-322, 332
t ogata La Ferte, Cicindela, 333
tranquebarica borealis Harrington,
Cicindela, 331

tranquebarica Herbst, Cicindela, 324,
331,333,335

tranquebarica kirbyi LeConte, Cicindela,
331

Trichognathus Latreille, 360
Trichognathus marginipennis Latreille,

351

Umbellularia californica, 230
Uniola, 238, 240
Uniola paniculata, 235-236, 238
Vernonia, 232

versuta Casey, Cicindela, 330
Vitis, 232

vittatum Zetterstedt, Simulium, 177,
181-182

vittatus Forel, Pseudomyrmex, 215
vuilloti Alluaud, Eunostus, 351
Wallis, J.B., 319, 322, 325, 329-330, 332,
334

Wheeler, G.C., 226, 229, 232, 235
Wheeler, J.,

see Wheeler, G.C., 226, 229, 232, 235
Wheeler, W.M., 225-229, 232, 234-235,
237-239

Whitcomb, W.H., 225
Willis, H.L., 320-322, 333-334
Wilson, E.O., 227, 235, 239
Wright, H.E., 334

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