Taxonomy and phylogeny of the Hister servus group (Coleoptera: Histeridae): a Neotropical radiation

Taxonomy and phylogeny of the Hister servus group(Coleoptera: Histeridae): a Neotropical radiation

M I C H A E L S . C A T E R I N ODepartment of Environmental Science, Policy and Management, Division of Insect Biology, University of California, U.S.A.

Abstract. The Hister servus group contains ten species and two species

complexes. Although the group's diversity is centred in Mexico and Central

America, one lineage, comprising H. indistinctus Say, H. defectus LeConte and

H. fungicola Schaeffer, has probably diversi®ed entirely in the south-eastern U.S.

The servus group also contains the following species: H. servus Erichson, H.

nodatus Lewis, H. doyeni sp.n., H. lagoi sp.n., H. bullatus Lewis, H. coronatus

Marseul and H. diadema Marseul and two diverse species complexes, dubbed the

comes complex and the montivagus complex. These two complexes require

additional sampling and study before species within them can be recognized. They

are, however, closely related to each other and appear to be paraphyletic with

respect to a clade comprising H. bullatus, H. coronatus and H. diadema. A prior

hypothesis that the servus group includes H. sallei, H. matador sp.n. and the

species of Hister (Spilodiscus) is not well supported. Several sister groups within

the servus group show geographical disjunction between seasonal tropical habitats

of western North America and wetter montane habitats of the eastern parts of the

continent along the Gulf and Caribbean coasts.

Introduction

The Hister servus group is endemic to the New World. The

species are largely tropical or warm temperate, with only a few

species reaching their distributional limits in the northern

U.S.A. and Canada. As is the case with most of the species of

the genus Hister Linnaeus, the species in this group exhibit

only subtle external differences, and identi®cation is proble-

matic. This study is the ®rst to recognize this lineage as a

putatively monophyletic unit and is the ®rst to treat the species

across their entire known ranges. The purpose of this study is

primarily to provide a means for identi®cation and to

document the distributions and known natural history of the

species of this prominent group. This work also entails the ®rst

examination of phylogenetic relationships among the known

species. Caterino (1998) hypothesized the inclusion of the

species of Spilodiscus Lewis, Hister sallei Marseul and H.

matador sp.n. in the servus group based on genitalic

characters. The phylogenetic analyses presented here serve,

in part, to test this idea. The phylogenetic hypothesis also

provides a framework for examining historical biogeographic

patterns exhibited by the species.

Surveys of variation across the ranges of two of the species

in this study (H. montivagus Lewis and H. comes Lewis)

revealed substantial variation in morphology among popula-

tions. It is clear that genetic isolation varies among popula-

tions, with many populations likely to merit species level

recognition. However, sampling at present has been insuf®-

cient to warrant description of numerous new species. Instead,

the designation `species complex' is applied to each of these

groups. These designations rest entirely on phenetic grounds at

present; phylogenetic analyses suggest that both complexes

may be paraphyletic. More material, and, ultimately, additional

character data will be necessary to assess the status and

relationships of these groups.

Taxonomic history

The close relationships among some of the species of the

servus group were recognized by several previous workers; the

species have been variously assigned to informal groups.

LeConte (1845), following Paykull (1811), split the North

American species of Hister into ®ve groups based on the striae

of the pronotum and the elytra. Most of the members of the

R

Correspondence: Michael S. Caterino, Department of Environmental

Science, Policy and Management, Division of Insect Biology, 201

Wellman Hall, University of California, Berkeley 94720-3112, U.S.A.

E-mail: [email protected]

# 1999 Blackwell Science Ltd 351

Systematic Entomology (1999) 24, 351±376Systematic Entomology (1999) 24, 351±376

servus lineage known at the time were treated in his group 2,

possessing two pronotal striae and lacking elytral striae.

In one of the earliest comprehensive works on the

Histeridae, Marseul (1854) expanded upon LeConte's system,

splitting the genus Hister into eight groups. Most of the species

of the servus lineage described at the time were placed together

in his group 7, characterized by the presence of two pronotal

striae in combination with the absence of subhumeral elytral

striae, although this group included numerous unrelated

species as well. Under this system Spilodiscus biplagiatus

LeConte was also included in group 7 (lacking subhumeral

striae). Marseul placed H. sallei Marseul in his group 8

(lacking subhumeral elytral striae but possessing only one

pronotal stria.)

Horn (1873) broke with the tradition of dividing Hister into

numbered groups and recognized numerous groups named for

`representative' species. Under this system, the species of

Spilodiscus Lewis were all brought together for the ®rst time in

the `group Arcuatus', and the remainder of the servus lineage

was placed in the `group Depurator.'

Lewis never published any formal system of grouping the

species of Hister aside from the proposal of the genus

Spilodiscus (Lewis, 1906) to accomodate the species of Horn's

`group Arcuatus.' Otherwise, beyond mentioning vague

interspeci®c af®nites in his descriptions, Lewis (1888) only

commented that numerous distinctive groupings exist within

Hister.

Casey (1916) ®rst delineated the `group servus', character-

ized by small size, a feebly sinuate mesosternum and bistriate

prosternum. This group contained H. servus Erichson, H.

densicauda Casey, H. cribricauda Casey, H. sternalis Casey,

H. defectus LeConte, H. indistinctus Say, H. fungicola

Schaeffer and H. nanulus Casey. He did not address the

af®nities of species occurring outside of the U.S.A. and

Canada. Nonetheless, his recognition of this group leads

directly to the present treatment.

Materials and methods

Morphological study

Specimens examined during this study were from the

collections of the following institutions and persons (abbrevia-

tions from Arnett et al., 1993): BMNH, The Natural History

Museum, London; CASC, California Academy of Sciences,

San Francisco; CDFA, California Department of Food and

Agriculture, Sacramento; CMNC, Canadian Museum of

Nature, Ottawa; CMNH, Carnegie Musuem of Natural History,

Pittsburg; CNCI, Canadian National Collection of Insects,

Ottawa; CUIC, Cornell University Insect Collection, Ithaca;

DVC, David Verity Collection; EMEC, Essig Museum of

Entomology, University of California, Berkeley; EMUS, Utah

State University Entomology Museum, Logan; FMNH, Field

Museum of Natural History, Chicago; FSCA, Florida State

Collection of Arthropods, Gainesville; INBI, Instituto Nacio-

nal de Biodiversidad, San JoseÂ; LACM, Natural History

Museum of Los Angeles County, Los Angeles; LSUC,

Louisiana State Arthropod Museum, Baton Rouge; MAIC,

Michael A. Ivie Collection, Bozeman; MCZC, Museum of

Comparative Zoology, Harvard University, Cambridge;

MNHN, Musee National D'Histoire Naturelle, Paris; MSCC,

M.S. Caterino Collection; PKC, Peter Kovarik Collection;

PSC, Paul Skelley Collection; SEMC, Snow Entomological

Museum, Lawrence; UMIC, University of Mississippi Collec-

tion, Oxford; USNM, United States National Museum of

Natural History, Washington; WFBM, William F. Barr

Museum, University of Idaho, Moscow; WSUC, James

Entomological Collection, Washington State University, Pull-

man; ZMHB, Zoological Museum of Humboldt University,

Berlin.

Specimens were examined with a Wild M5 binocular

microscope. Measurement conventions follow Wenzel &

Dybas (1941): lengths are measured from the midpoint of the

anterior margin of the pronotum to the apex of the elytral

suture, widths are measured at the widest point of the humeri,

near the anterior corners of the elytra. A summary of other

morphological terminology can be found in Caterino (1998).

For genitalic extractions, dry specimens were relaxed in

boiling water for a few seconds. The pygidium was then pulled

open using a ®nely hooked pin. The genitalic sheath was

grasped and pulled from the body using ®ne forceps and

transferred to a warm 10% KOH solution for clearing. After

about 15 min genitalia were rinsed in weakly acidic (a few

drops of acetic acid in 20 ml water) water and transferred to

glycerin-®lled shell vials which were then pinned beneath the

specimens for later examination.

The scanning electron microscopy was performed in the

Electron Microscopy Laboratory, College of Natural Re-

sources, University of California, Berkeley. The work was

done on an ISI (International Scienti®c Instruments) DS130

scope at 10 kV. Coating of specimens was generally not

required; however, some were coated with 20 nm of Gold-

Palladium using a Polaron sputter coater.

Phylogeny

OTUs. No character states are consistent throughout the taxa

treated in this study. Delimitation of the ingroup taxa was

based primarily on the presence of articulated dorsal projec-

tions of the aedeagal median lobe (Fig. 1). This is the primary

character justifying the inclusion of Spilodiscus and the sallei/

matador lineage. The remaining OTUs are included with

signi®cantly more support. Most share the following char-

acteristics: prosternal striae present, subhumeral elytral striae

absent, aedeagal gonopore situated more or less apical within

the tegmen, outer mandibular stria present, bursa copulatrix

poorly developed, lateral tegmenal cleft socket-like and

protibia triangular to rounded with a bidenticulate apical

tooth. Whether these represent synapomorphies or symplesio-

morphies will be discussed below.

Most of the OTUs in this analyses represent distinct species.

The comes and montivagus complexes represent polymorphic

lineages distributed throughout the North American tropics.

These are represented by several exemplar OTUs each in order

L

# 1999 Blackwell Science Ltd, Systematic Entomology, 24, 351±376

352 Michael S. Caterino352 Michael S. Caterino

to encompass the morphological (and probably phylogenetic)

diversity (the localities represented by each OTU are listed in

Table 1). Relationships within Spilodiscus have previously

been analysed (Caterino, 1998). One Spilodiscus exemplar is

included here to try to establish the relationships of this lineage

to the servus group.

Outgroups. Designating an outgroup to the servus group was

dif®cult because no hypotheses of phylogeny have been

published for Hister. I have selected outgroups used in this

analysis based primarily on a putative genitalic homology. The

independently articulated aedeagal armature is known in only a

few other Histerini; H. civilis LeConte and H. depurator Say

are included on that basis (Caterino, unpublished data). I have

also included Hister curtatus LeConte because it appears to be

only remotely related (and thus not likely part of the ingroup).

Characters (see Table 2 for scoring)

1. Supraorbital stria: (1) present (e.g. Fig. 7C,F); (2) absent

(Fig. 2A).

2. Frontal stria: (1) smoothly rounded; (2) bisinuate, frons not

depressed (Fig. 7C,F); (3) bisinuate, frons depressed behind.

3. Labrum: (1) rounded apically, as long as wide; (2) more

rectangular, from 1/2 to 2/3 as long as wide; (3) slightly

emarginate apically, 1/2 or less as long as wide.

4. Mandibles: (1) smoothly convex dorsally (Fig. 2A); (2)

¯attened or only shallowly depressed at base of dorsal

surface (Fig. 7C); (3) dorsal surface strongly depressed and

margined laterally (Fig. 7F).

5. Mandibles: (1) outer surface without longitudinal stria; (2)

outer surface with short longitudinal stria along lower

margin of outer face (Fig. 2C).

6. Submentum: (1) evenly convex, not produced anteriorly; (2)

acutely produced anteriorly.

7. Pronotum: (1) inner lateral stria continuous with anterior

marginal stria; (2) marginal stria continuous with anterior

marginal stria, the inner lateral stria ending free anteriorly.

8. Outer lateral pronotal stria: (1) complete; (2) abbreviated

posteriorly; (3) absent.

9. Striae of prosternal keel: (1) present; (2) absent.

10. Protibiae: (1) rounded, multidenticulate, 2 rows denticles

(Fig. 2B); (2) no basal marginal dentation (Fig. 4A); (3) tri-

or quadridentate, apical lateral tooth bidenticulate, promi-

nent apical marginal denticle present medial to apical

tooth, submarginal carina present (Fig. 15A); (4) tri- or

quadridentate, apical lateral tooth bidenticulate, lacking

prominent subapical marginal denticle, submarginal carina

present (Fig. 10A); (5) tri- or quadridentate, apical lateral

tooth bidenticulate, no submarginal carina; (6) tridentate,

apical lateral tooth bidenticulate, submarginal carina

present; (7) bidentate, lateral teeth strong, submarginal

carina present. While each of these states encompasses

several features, they each designate a recognizable

protibial con®guration which is shared by multiple OTUs.

R

Fig. 1. The aedeagus of Hister doyeni, in presumed copulatory

position, showing terminology of the various structures as applied in

this paper.

Table 1. Localities of origin of H. comes and H. montivagus OTUs.

OTU Included localities

H. comes 1 Mexico: Oaxaca (nr Mitla)

H. comes 2 Arizona: Chiricahua Mts; Texas: Big Bend N.P.; Mexico: Durango; Chihuahua.

H. comes 3 Arizona: Pajarita Mts

H. comes 4 Mexico: Chihuahua (30 mi NW Cd Chihuahua)

H. comes 5 Arizona: Chiricahua Mts; Huachuaca Mts; Mexico: Zacatecas

H. montivagus 1 Mexico: Oaxaca, south of Juchatengo

H. montivagus 2 Guatemala: Baja Verapaz

H. montivagus 3 Mexico: Veracruz (Sierra las Tuxtlas nr FortõÂn de las Flores)

H. montivagus 4 Mexico: Chiapas (Sumidero Canyon)

H. montivagus 5 Mexico: Tamaulipas; San LuõÂs PotosõÂ; Hidalgo; Veracruz (FortõÂn de las Flores)

H. montivagus 6 Panama: ChiriquõÂ; Costa Rica: Puntarenas (Monteverde)

H. montivagus 7 Costa Rica: Guanacaste; Alajuela; Puntarenas (P.N. Amistad)


The Hister servus group 353The Hister servus group 353

L

Tab

le2.

Ch

arac

ter

stat

esan

din

div

idu

alch

arac

ter

consi

sten

cyin

dic

es.

à'

=1/2

;`b

'=

1/3

.

12

34

56

78

910

11

12

13

14

15

16

17

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

Ch

arac

ter

CI

0.6

0.4

0.7

0.6

0.8

0.5

0.5

0.5

0.5

0.7

0.3

10.3

11

0.3

0.3

0.4

0.7

10.3

0.4

0.8

11

0.3

10.4

0.5

10.8

10.5

0.5

H.

civi

lis

21

21

11

12

21

22

21

22

22

21

11

11

11

13

12

11

11

H.

dep

ura

tor

21

21

11

12

26

22

11

a2

12

11

21

11

11

13

12

31

11

Sp

.b

ipla

gia

tus

21

21

11

11

27

12

21

21

?2

11

21

11

12

13

32

11

22

H.

curt

atu

s2

22

11

21

22

62

12

11

11

21

11

11

11

11

11

11

1?

?

H.

fun

gic

ola

21

31

21

12

15

22

21

12

11

21

21

11

11

22

24

41

11

H.

serv

us

12

33

21

11

15

22

21

11

11

11

22

11

13

21

15

11

11

H.

def

ectu

s2

12

12

11

21

51

22

11

11

12

12

11

11

22

22

44

11

1

H.

lag

oi

22

21

a1

12

14

22

21

11

21

11

21

11

11

21

24

22

??

H.

ind

isti

nct

us

21

21

21

12

15

12

21

11

11

21

21

11

11

22

24

41

11

H.

dia

dem

aa

32

22

11

21

52

22

11

11

a1

12

22

21

22

11

31

21

2

H.

bu

lla

tus

23

22

21

12

15

22

21

12

11

11

22

22

11

22

13

12

??

H.

coro

na

tus

13

22

21

22

25

22

21

12

22

11

22

22

12

22

13

12

??

H.

no

da

tus

13

23

21

11

15

22

21

11

12

11

22

11

13

21

15

11

11

H.

do

yen

i2

22

22

21

11

51

21

11

22

21

11

11

11

12

12

44

11

1

H.

sall

ei1

21

11

12

31

22

22

11

11

11

21

21

11

21

33

21

12

1

H.

ma

tad

or

12

11

11

23

11

22

21

11

11

12

1a

11

11

13

32

31

21

H.

mo

nti

vagus

12

22

21

11

21

42

21

11

21

11

12

11

1?

??

??

??

??

?

H.

mo

nti

vagus

2a

22

22

11

21

42

22

11

11

11

12

12

1?

??

??

??

??

?

H.

mo

nti

vagus

32

22

22

?1

21

42

22

11

11

11

12

12

1?

??

??

??

??

?

H.

mo

nti

vagus

42

22

22

11

21

42

22

11

11

11

12

22

12

12

21

31

2?

?

H.

mo

nti

vagus

52

22

a2

11

21

42

22

11

11

11

12

2a

11

22

21

31

21

2

H.

mo

nti

vagus

62

22

11

11

21

42

22

11

11

21

12

2a

11

12

21

31

21

2

H.

mo

nti

vagus

72

22

22

11

21

42

22

11

11

11

12

11

12

12

21

31

2?

?

H.

com

es1

22

22

a1

12

14

22

21

11

1a

11

21

11

12

22

14

12

11

H.

com

es2

22

22

a1

12

14

22

21

11

11

11

21

11

12

22

14

12

11

H.

com

es3

22

22

a1

12

14

22

21

11

11

11

21

11

12

21

14

12

11

H.

com

es4

22

22

a1

12

14

22

21

a2

22

b1

21

11

12

22

14

12

11

H.

com

es5

23

22

a1

12

14

22

21

1a

aa

11

21

11

12

22

14

12

11



11. Anterior face of protibia: (1) distinct row of setae arising

near base of lateral margin; (2) row of setae indistinct, at

most a few setae subserially arranged.

Note on characters 12±17. Elytral striae show substantial

intraspeci®c variation, though some aspects of their distribu-

tion are informative. The stria is scored as present if at all

detectable. These characters have been collectively weighted

as equivalent to one character in all analyses.

12. Outer subhumeral elytral stria: (1) present; (2) absent.

13. Inner subhumeral elytral stria: (1) present; (2) absent.

14. Third discal elytral stria: (1) present; (2) absent.

15. Fourth discal elytral stria: (1) present; (2) absent.

16. Fifth discal elytral stria: (1) present; (2) absent.

17. Sutural elytral stria: (1) present; (2) absent.

18. Anterior portion of metasternal stria: (1) anteriorly

complete; (2) convex posteriorly along mesometasternal

suture.

19. Posterolateral portion of metasternal stria: (1) extending

R

Fig. 2. A, Head of Hister defectus. B, Left protibia of H. civilis, posterior view. C, Lateral view of the left mandible of H. servus showing the

stria along the lower outer margin. D, Metatarsus of H. sallei, showing four spines of basal tarsomere. E, Metatarsus of H. servus, showing

bispinose basal tarsomere.



laterally to metepisternum; (2) curved outward posteriorly

but not reaching metepisternum; (3) ending posteriorly

without curving outward.

20. Metatibia: (1) 2 parallel, proximate rows of marginal

spinules; (2) inner row of marginal spinules displaced onto

outer face of tibia, especially basally, rows not parallel

(Fig. 4C).

21. Meso- and metatarsus: (1) basal tarsomere with > 2 ventral

spinules (Fig. 2D); (2) basal tarsomere with 2 spinules on

ventral apical margin (Fig. 2E).

22. Propygidium: (1) ¯at or evenly convex; (2) with lateral

depressions.

23. Pygidial punctures: (1) shallow, with centres slightly

raised (Fig. 7B); (2) deep, especially at pygidial apex

(Fig. 7E).

24. Pygidial punctures: (1) evenly and usually densely spaced

(Fig. 7B); (2) irregularly spaced, with areas devoid of

punctation (Fig. 15C).

25. Basal piece: (1) length one-third or less of total aedeagal

length (Fig. 8A); (2) half of aedeagal length (i.e. equal in

length to the tegmen; seen only in a few populations of H.

montivagus).

26. Dorsal lobes of tegmen: (1) broad, generally straight and

not widely separated dorsally (Fig. 13C); (2) curved,

tapering, and separated dorsally (Fig. 13D); (3) narrow,

straight, not widely separated dorsally (Fig. 8A).

The tegmenal lobes, when broad (state 1) cover the

tegmenal opening dorsally. In state 2 the lobes are

emarginate on their inner edges, thus appearing curved

and exposing part of the median lobe; in this state the

armature usually remains covered from above by the

tegmenal lobes. In state 3 the tegmenal lobes are reduced

laterally, abutting along the dorsal midline and exposing the

armature at the sides.

27. Lateral tegmenal cleft: (1) narrow, slit-like (Fig. 3C); (2)

rounded basally, socket-like (Fig. 9E±G).

28. Lateral surface of tegmen: (1) longitudinally convex; (2)

longitudinally concave; (3) longitudinally furrowed, with

ventral lamellae.

29. Dorsal median rods of median lobe: (1) absent; (2)

present, short (Fig. 9E±G); (3) present, elongate, slender

(Fig. 3B,C).

30. Armature of median lobe: (1) absent; (2) long, sinuate

(Fig. 3B); (3) short, divergent (Fig. 16C); (4) longer,

divergent and bearing oblique carina separating basal

and apical parts (Fig. 9E±G). (5) longer, divergent and

lacking carina (Fig. 8A).

31. Proximal apodemes of median lobe: (1) thin and well

separated (Fig. 8B); (2) robust and well separated

(Fig. 13C); (3) robust and basally convergent (Fig. 9E);

(4) robust and abutting or nearly abutting along midline

(Fig. 9F).

32. Length of gonopore bearing tube (apical portion of

median lobe): (1) short, with gonopore situated near bases

of armature (Fig. 9E); (2) long, with gonopore situated

near apices of armature (Fig. 16A).

33. Bursa copulatrix: (1) weakly sclerotized and atrophied; (2)

well sclerotized and paddlelike.

The bursa is reduced and more or less sclerotized in many

Histerini. It is not known how this condition affects its

function.

34. Spermathecae: (1) rounded, not tapered apically; (2)

tapered, apically acute.

Phylogenetic analyses

All analyses were performed using PAUP 3.1.1 (Swofford,

1993). For parsimony analysis the following search conditions

were implemented: heuristic search, ten replicates of random

taxon addition sequence, nearest neighbour (NNI) branch-

swapping, all characters unordered, elytral striae weighted as

explained above. Branch support was assessed using boot-

strapping (100 replicates, heuristic search, NNI branch-

swapping). Calculation of decay indices (Bremer, 1994) was

also attempted. However, using the full data set, a large

number of similarly parsimonious resolutions among comes

and montivagus complex populations precluded a thorough

decay analysis. In order to get some idea of decay support for

groups outside of the comes/montivagus populations, indices

were calculated using two OTUs from each complex (comes 2

and 5; montivagus 4 and 6). Branch-swapping was performed

using MacClade 3.0.6 (Maddison & Maddison, 1992) to

examine the cladogram lengths of alternative resolutions.

A key to the species (adults)

There is no single feature with which to diagnose the servus

group species. Most of the species lack subhumeral elytral

striae and have a striate prosternum (with the exception of H.

coronatus Marseul). However, prosternal striae are also seen in

other New World Hister which are not closely related (H.

criticus Marseul, H. latimargo Schmidt and others). The

subhumeral striae are likewise absent in a few unrelated

species. Thus, the following key allows separation of servus

group species from all New World Histerini, including all

North and South American and Caribbean species. Taxa in

parentheses are not members of the servus group. The elytral

striae in histerids are numbered from the outside inward (see

Fig. 3D).

1. Hypomeron setose.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

± Hypomeron not setose... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2(1). Protibiae strongly bidentate; elytra of most individuals

with red maculations; meso- and metatibiae apically

expanded in many individuals; aedeagus character-

istic, with elongate, sinuate armature and elongate

dorsal projections.. . Spilodiscus (see Caterino, 1998).

± Protibiae with three or more teeth; elytra of few

individuals with red maculations; aedeagus of most

individuals with armature short or absent . .. . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (not servus group)

L



3(1). Each elytron with 3 longitudinal costae; most external

striae costate; myrmecophilous .. . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Epiglyptus costatus)

± Elytra not costate.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4(3). External subhumeral stria complete in most indivi-

duals; median lobe of aedeagus with apical

gonopore ¯anked by paired armature hinged to

median lobe at their bases (at meeting point of

median lobe and proximal apodemes), proximal

apodemes short (Fig. 3A; also see Wenzel,

1944) . . . . . . . . . . . . . . . . . . . . . . . . . . (Margarinotus)

± External subhumeral stria abbreviated; median lobe of

aedeagus with gonopore distinctly subapical in most

individuals; armature, if present, consisting of more

than one pair of hinged arms; proximal apodemes

longer (e.g. Figs 8A, 9E) .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5(4). Meso- and metatibiae ¯at, expanded, nearly rectan-

gular; body surfaces, especially frons, strongly

punctate, rufescent; principally occurring in ant

nests .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Psiloscelis)

± Meso- and metatibiae not rectangular; frons not

strongly punctate .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6(5). Mesosternum truncate or only very slightly emargi-

nate ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Atholus)

± Mesosternum distinctly emarginate, receiving poster-

ior prosternal process ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7(6). Prosternal keel without striae ... . . . . . . . . . . . . . . . . . . . . . . . . 8

± Prosternal keel striate... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8(7). Elytra with striae 1 and 2 complete, third stria present

in basal half, fourth stria in basal one-fourth;

subhumeral striae absent (Fig. 18C); pygidial punctu-

res large, deep, irregularly scattered (Fig. 15C);

propygidium impressed laterally; outer lateral pronotal

stria present in apical one-third or less; known from

Mexico and through Central America to Costa

Rica... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. coronatus Marseul

± Elytra with more than 2 complete striae; pygidial

punctures regularly spaced or absent . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (not servus group)

9(7). Protibiae 2 or 3 dentate, without submarginal carina;

mandibles distinctly dentate; aedeagus with basal

piece longer than tegmen, proximal apodemes very

short; both lateral pronotal striae complete; body

rufescent; meso- and metatibiae very slender; Central

America and northern South America ... . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (H. belti Lewis)

± Protibiae with 3 or more marginal teeth, most

individuals with submarginal protibial carina; mand-

ibles not distinctly dentate; aedeagus with basal piece

shorter than tegmen; median lobe armature present;

proximal apodemes well developed; other characters

varied.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10(9). Outer lateral pronotal stria absent . . . . . . . . . . . . . . . . . . . . 11

± Outer lateral pronotal stria present . . . . . . . . . . . . . . . . . . . 13

11(10). Larger species (3.3±5.5 mm in length); protibia

rounded, in many individuals lacking denticles along

basal portion of margin; mandibles convex above;

third dorsal elytral stria interrupted near apex in many

individuals (Fig. 3D,E); aedeagus with elongate dorsal

median rods and sinuate armature nearly the length of

the tegmen (Fig. 3B,C) .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

± Smaller species (most individuals less than 4.0 mm in

length); protibia strongly tridentate, apical tooth

bispinose; mandibles ¯at above; third elytral stria

complete; aedeagus with armature short, not articu-

lated .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (not servus group)

12(11). Protibia without denticles on basal half of margin, on

apical half only (Fig. 4A); widespread from Mexico to

Venezuela.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. sallei Marseul

± Protibia with denticles along entire length (Fig. 4B);

known only from southern MeÂxico .. . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. matador Caterino, sp.n.

13(10). Pygidium with deep, irregularly spaced punctures

(Fig. 15C) .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

± Pygidial punctures varied in size, but evenly dis-

tributed .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14(13). Supraorbital stria absent; third elytral stria complete in

most individuals; known from Mexico and Guatema-

la .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. bullatus Lewis

± Supraorbital stria present, though abbreviated in some

individuals; third elytral stria interrupted; known from

Amazonia .. . . . . . . . . . . . . . . . . . . . . . . . . . H. diadema Marseul

15(13). Sutural elytral stria absent .. . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

± Sutural elytral stria present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

16(15). Both lateral pronotal striae complete; frontal stria

subcarinate, frons laterally impressed immediately

behind frontal stria; mandibles densely punctate at

base; known from Oaxaca and Puebla ... . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. doyeni Caterino, sp.n.

± Outer lateral pronotal stria abbreviated posteriorly;

frontal stria not carinate.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17(16). Protibia rounded apically (Fig. 13B); known from

Texas to Florida ... . . . . . . . . . . . . H. lagoi Caterino, sp.n.

± Protibiae acute apically; known from Texas, New

Mexico and Arizona south to Oaxaca .. .. . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. comes Lewis complex

18(15). Supraorbital stria present; both lateral pronotal striae

complete .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

± Supraorbital stria absent; lateral pronotal striae

varied... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19(18). Meso-metasternal stria complete across anterior mar-

gin of metasternal disk; pygidial punctation varied,

dense throughout in most individuals; widespread

throughout North and South America .. . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. servus Erichson

± Meso-metasternal stria interrupted medially; pygidium

with deep punctures along apical margin, central

portion of disk raised (Fig. 7E); Costa Rica, PanamaÂ

and northern South America only ... . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. nodatus Lewis

20(18). Mandibles convex above (Fig. 2A); body small,

strongly rounded in outline; proximal apodemes of

aedeagal median lobe broad, abutting along midline

(Fig. 9F); eastern U.S.A ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

± Mandibles ¯attened to depressed above; body shape

varied; proximal apodemes of median lobe thin,

R



separate (Figs 13D, 16A); known from south-western

U.S.A. to Panama ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

21(20). Mesosternum with 2 complete striae along anterior

margin.. . . . . . . . . . . . . . . . . . . . . . . . . . . H. fungicola Schaeffer

± Mesosternum with one complete marginal stria . . 22

22(21). Denticles on apical tooth of protibia separated by the

width of one denticle or more (Fig. 10C) ... . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. defectus LeConte

± Denticles on apical protibial tooth abutting or

separated by less than the width of one denticle

(Fig. 10B) .. . . . . . . . . . . . . . . . . . . . . . . . . . . . H. indistinctus Say

23(20). Fourth elytral stria interrupted in most individuals,

visible both basally and apically (Fig. 18A); meso-and

metatibiae slender .. . . . H. montivagus Lewis complex

± Fourth elytral stria abbreviated, present only apically,

or absent (Fig. 13F±H); meso-and metatibiae broad in

most populations .. . . . . . . . . . . H. comes Lewis complex

Hister sallei Marseul

Hister sallei Marseul, 1854: 584.

Diagnosis. Adults of H. sallei can be separated from all other

New World Histerini by the following combination of

characters: prosternal striae present, a single lateral pronotal

stria present and protibiae rounded, lacking denticles along the

basal half of lateral margin (Fig. 4A).

Description. Average length 4.9 mm (range 4.1±5.5; n = 7);

average width 4.7 mm (range 3.8±5.1; n = 7). Frons more or

less smooth, in some individuals weakly depressed; frontal

stria well impressed, straight anteriorly; labrum almost as long

as wide, rounded apically. Mandibles punctate externally; right

mandible overlapping left almost to base when head in repose;

left mandible with small tooth at base of incisor in some

individuals. Submentum elevated relative to genae, with 4±6

setae; mentum ¯at to convex, anteriorly emarginate, smooth at

middle, laterally setose; stipes with 3 or 4 setae along inner and

lateral margins; penultimate labial palpomere with one long

seta at inner apex, in some individuals with few short setae as

well; terminal palpomere with several short setae on inner

surface.

Pronotum with disk smooth and shining; one lateral stria

present from near base to apex, not connected to marginal stria

L

Fig. 3. A, The aedeagus of Margarinotus fractifrons (LeConte),

dorsal view. The aedeagus in Margarinotus species have armature

which is not independently articulated as in the servus group. The

two arms of the armature rotate upwards during copulations whereas

those of members of the servus group independently rotate laterally.

B, Aedeagus of Hister sallei, dorsal view. C, Aedeagus of H.

matador, dorsal view. D, Left elytron of H. sallei. E, Left elytron of

H. matador.

Fig. 4. A, Left protibia of Hister sallei, posterior view. B, Left protibia of H. matador, posterior view. C, Left metatibia of H. sallei.



anteriorly; interval between stria and pronotal margin more or

less convex; hypomeron without setae; prosternal keel with 2

complete striae in most individuals, rarely absent; prosternal

lobe rounded, with complete marginal stria; protrochanter with

one seta. Protibia (Fig. 4A) with teeth only along approxi-

mately apical half of lateral margin; submarginal protibial

carina present, bearing 3 or 4 denticles.

Elytra (Fig. 3D) lacking outer subhumeral stria; inner

subhumeral stria absent or represented by few punctures near

apex; striae 1 and 2 complete; striae 3 and 4 varied, complete

or abbreviated; stria 5 absent except for trace at apex; sutural

stria present in apical half. Mesosternum broad, with complete

anterior marginal stria; metasternal stria interrupted anteriorly,

laterally extending to metepisternum; mesotibia weakly

curved, expanded apically; metatibia (Fig. 4C) appearing bent

at middle, with rows of marginal spinules separated by

broadened tibial margin near base; basal meso-and metatar-

someres with 2 pairs of spinules ventrally in most individuals

(Fig. 2D).

Propygidium and pygidium coarsely punctate, pygidial

punctures dense, those of propygidum slightly less so.

Aedeagus (Fig. 3B) with basal piece short; tegmen wider

apically, dorsal lobes shallowly medially emarginate; median

lobe with proximal apodemes thin, short; armature sinuate,

nearly as long as tegmen; median rods elongate, tapered.

Biology. Specimens of H. sallei have been attracted to a

variety of rotting organic material, including dung, carrion and

fruit. The single record from `morning glories' is very unusual,

and possibly erroneous. Collecting in Belize suggests that

populations of this species are very susceptible to disturbance,

occurring almost exclusively in primary mesic forest (P.

Kovarik, personal communication).

Remarks. I have seen two similarly atypical specimens, one

female from ChiriquõÂ, Panama and one male from Guanacaste,

Costa Rica, in localities separated by over 400 km. They share

several characters, including extremely broad, nearly rectan-

gular, metatibiae on which the two series of marginal spinules

are widely separated, an elongate, narrow prosternal lobe,

bidentate protibiae, submarginal protibial carina lacking

denticles, elongate mandibles and very short meso- and

metatarsomeres. They are also slightly larger than most

individuals of H. sallei. Only the large geographical separation

and intervening records of typical sallei keep me from

describing these two specimens as a distinct species.

Holotype. Female, `Caracas, Salle, 63¢ [Venezuela]

(MNHN); examined 1995. Other material (see Fig. 5).

MEXICO: Tamaulipas, GoÂmez FarõÂas, 1000 m, viii.1983,

cloud forest (CMNC). San LuõÂs PotosõÂ, 3 mi W Naranjo,

1200¢, vi.1965 (PKC). Veracruz, Canyon RõÂo Metlac, 3200¢,viii.1973 (FMNH); FortõÂn de las Flores, vi.1975 (PKC); one mi

W FortõÂn de las Flores, viii.1962 (EMEC); 5 mi W Catemaco,

viii.1970, on morning glories (FMNH); 6 mi NE Catemaco,

1500¢, rotting fruit, viii.1970; Lake Catemaco, vi.1969,

viii.1960 (CNCI); 33 km N Catemaco Las Tuxtlas, vi.1984

(CMNC); 4 mi N Huatusco, 4200¢, vii.1973 (CMNC). Oaxaca,

5 mi W Tequisistlan, 1200¢, viii.1973 (FMNH). Chiapas,

Palenque, v.1965 (FMNH); Palenque, 100 m, vii.1983, rain

forest (CNCI); Palenque, ix.1974 (PKC); 16 km NW Ocozo-

coautla, 970 m, v.1990 (CMNC); P.N. Sumidero, 1000 m,

v.1990 (CMNC); P.N. Sumidero, vii.1990, faeces pitfall

(PKC); nr Cintalapa, v.1969 (CMNC); 100 km S Palenque,

vii.1983, 230¢, rotting sapote fruit (CMNC). BELIZE: Toledo,

Blue Ck Village, vi.1981 (USNM); Orange Walk, RõÂo Bravo

Cons. Area, ix.1995 (PKC). GUATEMALA: Zacapa, 3.5 km

SE La UnioÂn, 1500 m, vi.1993, cloud forest (CMNC).

HONDURAS: Olancho, P.N. La Muralla, vi.1995 (PKC).

COSTA RICA: Guan. Vol. Tenorio, RõÂo San Lorenzo, 1050 m,

iv.1992; P.N. Guan., 1000±1400 m, v.1992 (INBI); Maritza

Est. Biol., 550 m, v.1993. Puntarenas, Monteverde, 1400 m,

v.1989 (MSCC); Monteverde, vi.1983, viii.1987 (CMNC); S.

Vito, Las Cruces, 1200 m, viii.1982 (CMNC); P.N. La

Amistad, Est. Las Mellizas, 1300 m, vii.1990 (INBI); Est.

Biol. Las Alturas, Coto Brus, x.1991, 1500 m (INBI). LimoÂn,

Res. Biol. Hitoy Cerere, 100 m, iv.1992 (INBI). PANAMA:

ChiriquõÂ, 25 km NNE San Felix, 1500 m, cloud forest, carrion

and human dung trap (FMNH); ChiriquõÂ, La Fortuna, 1150 m,

vi.1995 (SEMC); ChiriquõÂ, 27.7 km W VolcaÂn, 1450 m,

vi.1995 (SEMC); ChiriquõÂ, 20 km N Gualaca, 1350 m,

v.1995; ChiriquõÂ, Sta. Clara, vi.1985, human faeces pitfall

(PKC); ColoÂn, Cana Biol. Sta., 1250 m, vi.1996 (SEMC);

CocleÂ, 7.2 km NE El CopeÂ, 730 m, v,vi.1995 (SEMC).

COLOMBIA: Valle de Cauca (MNHN). VENEZUELA:

Araugua, 1200 m, Maracay, Rancho Grande, viii.1987, cloud

forest (CMNC); Miranda, 700 m, 28 km N AltagracõÂa, P.N.

Guatapo, viii.1987 (CMNC).

Hister matador Caterino, sp.n.

Diagnosis. Hister matador is closely related to H. sallei. It

differs in that the protibial denticles are present along the entire

margin (Fig. 4B).

Description. Average length 4.42 mm (range 3.3±4.9; n=7);

average width 4.2 mm (range 3.4±4.7; n = 7). Frons smooth,

narrow; frontal stria inturned or interrupted anteriorly; labrum

almost as long as wide, rounded apically. Mandibles punctate

R

Fig. 5. Collection records of Hister sallei and H. matador.



externally; left mandible overlapping the right almost to base

when head in repose; left mandible with small tooth at base of

incisor in some individuals. Submentum elevated relative to

genae, with 4±6 setae; mentum ¯at to convex, anteriorly

emarginate, smooth at middle, laterally setose; stipes with 3 or

4 setae along inner and lateral margins; penultimate labial

palpomere with one long seta at inner apex, sometimes with

short setae as well; terminal labial palpomere with several

short setae on inner edge.

Pronotum with disk smooth and shining; one nearly

complete lateral stria, not connecting to marginal stria

anteriorly; interval between stria and pronotal margin more

or less convex; hypomeron without setae; prosternal keel with

2 complete striae in most individuals, rarely none; prosternal

lobe rounded, with complete marginal stria. Protrochanter with

one seta; protibia rounded apically (Fig. 4B), with 4±6

marginal teeth along margin separated by shallow emargina-

tions; submarginal protibial carina usually present but weak,

submarginal denticles present.

Elytra (Fig. 3E) lacking outer subhumeral stria; inner

subhumeral stria absent or visible as punctures near apex;

striae 1 and 2 complete, stria 3 interrupted behind middle; stria

4 visible at base and apex; stria 5 absent except for trace at

apex; sutural stria present in about apical half of elytron.

Mesosternum broad, with complete anterior marginal stria;

metasternal stria almost complete anteriorly, laterally recurved

to basal one-third of metepisternum; mesotibia weakly curved,

expanded apically; metatibia appearing bent at middle, rows of

marginal spinules separated by broadened tibial margin near

base (Fig. 4C); basal meso-and metatarsomeres usually with 2

pairs of ventral spinules.

Propygidial punctures medium sized and evenly scattered;

pygidial punctures smaller, more widely scattered than those of

propygidium. Aedeagus (Fig. 3C) strongly sclerotized; basal

piece short; tegmen slightly widened apically, medial edge of

dorsal lobes straight; median lobe with short, close proximal

apodemes; armature sinuate, as long as tegmen; median rods

elongate, tapered.

Distribution and biology. This species has been attracted to

carrion traps and rotting cactus. Its collection records are

mapped in Fig. 5.

Etymology. The name matador literally translates to `killer.'

Holotype. Male, dissected by the author: `MEXICO:

NayarõÂt, Arroyo Santiago, 3 mi. N.W. of JesuÂs MarõÂa,

VII:6 : 1955, leg. B. Malkin' (FMNH). Paratypes (31). 10,

same data as holotype; one of each sex in MSCC; others

deposited in FMNH. 5, `MEXICO: NayarõÂt, JesuÂs MarõÂa,

30.vii.1955, carrion trap, leg. B. Malkin' (FMNH). 6,

`MEXICO: NayarõÂt, JesuÂs MarõÂa, VII:25±30 : 1955, leg B.

Malkin'/òn carrion bait' (FMNH). 4, `MEX: Oaxaca, 5 mi W

Tequisistlan, 1100¢, ix.61973'/À. Newton, collector'/òn

rotting columnar cacti' (1, FMNH, 3, MSCC). 3, `MEXICO:

Oaxaca, 5 mi. W. Tequisistlan, 1100 ft. viii,23-ix,5.75'/òn

rotting cacti columnar/Opuntia A. Newton' (FMNH). One,

`MEX: Est. Biol. Chamela, Jalisco, vii.20/27±1984, J.A.

Chemsak, J.T. Doyen, at lights' (EMEC). 2, `MEXICO Jalisco

Chamela, 1/8-X-85 F.D. Parker, T.L. Griswold' (EMUS). One,

`MEX. Jalisco, 7 km N. Malacque, 16,19 July 1990. J.E.

Wappes' (PKC). 3, `MEXICO: Morelos, Sierra de Huatla,

Presa Cruz Pintada, RõÂo Quilamula 1050 m `CEAMISH' 18±

27.5'N, 99±02.2¢W, 4±7 Sept 1996 R. Brooks ex. ¯ight

intercept trap' Bar code #¢s SM0048940 (SEMC), SM0048943

(SEMC), SM0048947 (MSCC). One, `MEÂ XICO: Guerrero,

Municipio Zumpango, 12 km S RõÂo Mescala (RõÂo Balsas)

bridge at Mescala on Hwy. 95'/¢(MeÂxico-Acapulco rd.) CanÄon

de Zopilote, 650 m, 17 August 1991 Neil Woodman, ex.

pitfall'.

Hister servus Erichson

Hister servus Erichson, 1834: 147.

Hister nodatus obliteratus Lewis, 1888: 203. Lectotype,

hereby designated: `Chacoj, Vera Paz, Champion'

[Guatemala] (BMNH); examined 1995; previously syno-

nymized in error with H. nodatus Lewis by Bickhardt

(1917). Syn.n.

Hister gibberosus Lewis, 1900: 232. Holotype: `MeÂxico City,

Flohr'/`G. Lewis coll. 1926±369'/Èsperanza' (BMNH);

examined 1995. Syn.n.

Hister meridanus Lewis, 1900: 233. Lecotype, hereby

designated: `Yucatan'/¢G. Lewis coll. 1926±369¢ [MeÂxico];

one paralectotype: same data (BMNH); examined 1995.

Syn.n.

Hister densicauda Casey, 1916: 222. Lectotype, here desig-

nated, a male: no data (nor published type locality); USNM

#38425; examined 1996. Syn.n.

Hister cribricauda Casey, 1916: 222. Lectotype, here desig-

nated, a male: `Sedgewick Co., Kans.'/`Casey bequest

1925'; USNM #38439. Two paralectotypes: `Mt. Hope,

Ks'; all examined 1997. Synonymized with H. densicauda

by Wenzel (1939).

L

Fig. 6. Collection records of Hister servus and H. nodatus.



Diagnosis. The presence of a supraorbital stria, prosternal

striae, 2 complete lateral pronotal striae and ¯attened or

depressed mandibles (Fig. 7C) separate H. servus and H.

nodatus adults from those of all other sympatric Hister. Adults

of H. servus differ from H. nodatus by having an evenly

convex pygidium (Fig. 7B) and complete anterior metasternal

stria, and are larger in size.


average width 3.1 mm (range 2.7±3.7; n = 8). Elongate oval,

widest behind middle. Frons broad, smooth, rounded (Fig. 7C);

frontal stria bisinuate anteriorly, continuous with complete

supraorbital stria; labrum twice as wide as long, weakly

emarginate at apex. Mandibles moderately to strongly

depressed dorsally at base (Fig. 7C), many individuals with

short marginal stria on upper surface of mandible; outer face of

mandible with short stria along lower margin. Submentum

with 2 or 3 setae laterally; mentum ¯at, medially smooth,

laterally setose, emarginate at apex; stipes with 8±10 setae;

penultimate labial palpomere with crown of 5 or 6 long setae at

apex; terminal palpomere with scattered short setae.

Pronotal disk smooth at middle with row of basal punctures;

inner and outer lateral pronotal striae complete, the inner stria

sinuate; prosternal keel with 2 complete striae, the striae

meeting anteriorly and posteriorly in most individuals;

prosternal lobe rounded at apex, gradually tapered to

hypomeron. Protibia with 3 strong teeth and one weaker tooth

at base (Fig. 7A), apical tooth with 2 denticles; small denticles

also present on apical margin; submarginal protibial carina

visible at base, bearing 1 or 2 small denticles, joined to margin

at second basal tooth.

Elytra (Fig. 8C) lacking subhumeral striae; striae 1±4

complete; stria 5 present in apical one-third or less; sutural

stria present in apical half to two-thirds. Mesosternum weakly

emarginate, anterior marginal stria complete, outer marginal

stria visible in anterior corners; metasternal disk smooth;

metasternal stria complete or barely interrupted at midline,

laterally extended to metepisternum; mesotibia curved, with 2

complete rows marginal spinules; metatibia straighter, with 2

rows marginal spinules, basal-most spinules somewhat weaker

or absent; all meso- and meta tarsomeres with one pair of

spinules ventrally.

Propygidium and pygidium densely and normally completely

punctate (Fig. 7B); size of punctures variable. Aedeagus

(Fig. 8A) generally broad though variable in overall propor-

R

Fig. 7. A, Left protibia of Hister servus, posterior view. B, Propygidium and pygidium of Hister servus. C, Head of H. servus. D, Left protibia

of H. nodatus, posterior view. E, Propygidium and pygidium of H. nodatus. F, Head of H. nodatus.



tions; basal piece short; tegmen parallel-sided, dorsal lobes

narrow, meeting along midline; tegmen lobate apically; median

lobe with thin proximal apodemes extending through basal

piece, with continuous sclerotization to apical lobes; armature

about half tegmenal length, curved inward slightly at apex.

Biology. Though a widespread and commonly encountered

species, little has been recorded regarding the habits of H.

servus. It has been found in association with rotting cactus and

carrion. Interestingly it is known from both seasonally dry as

well as generally wet areas.

Lectotype. Female, hereby designated: `Cuba'; ZMHB

#148922. Four paralectotypes, same data as lectotype;

examined 1996. Other material (see Fig. 6). U.S.A.: Wiscon-

sin, Delawan (FMNH). New York, Pelham Bay St. Pk, burnt

fungus. Missouri, Boonesboro, iv.1954 (CDFA). Illinois,

Berwyn (FMNH); Evanston. Indiana, Tippecanoe Co.,

vi.1977; Tippecanoe Co., v.1975 (CUIC); Lafayette Co.

Pennsylvania, Allegheny Co. (CUIC). District of Columbia,

Washington, vi.1893 (USNM); Jeanette, vi.? (CMNH); St.

Vinc. (CMNH). Arkansas, Ft. Smith (USNM). Tennessee,

Friendsville (FMNH); Benton Co., vii.1952. North Carolina,

Chimney Rock (FMNH). South Carolina, Clemson (FMNH).

Arizona, Chiricahua Mts; Globe; Tucson viii.1933 (USNM);

Madera Cyn ix.1955. Texas, Kingsville, V-1920 (LACM);

New Braunfels; Corpus Christi (USNM); Florida, Highlands

Co., Archibold St. Pk, viii.1979 (FSCA); Orange Co., vi.1958

(FSCA); Dade Co., Ross Hammock, vi.1948 (FSCA); Dade

Co., ii.1934; Lake Alfred, iii.1963 (FSCA); Mims, v.1966

(FSCA); Dade Co., Matheson Hammock, iv.1951 (CMNC);

Dade Co., Everglades Nat. Pk (FMNH); Capron (USNM);

Palm Beach Co., Lantana, viii.1986 (FSCA); Dade Co.,

vii.1988 (FSCA). MEXICO: Baja California Sur, Isla Sta.

Catalina, vi.1971 (CASC). Sonora, Advana, 5 mi W Alamos,

vii.1958; Bahia Kino, vii.1980. Sinaloa, 25 mi N MazatlaÂn,

viii.1960 (PKC); 7 mi S CuliacaÂn, viii.1960 (PKC). Nayarit,

JesuÂs MarõÂa, vii.1955; Arroyo Santiago, San LuõÂs PotosõÂ, El

Salto Falls, 12 km NW El Naranjo, 400 m, vii.1990 (SEMC);

nr Mante, Tamaulipas, nr Gomez Farias, 120 m, vii.1992,

rotting leaves (SEMC); Mun, Villa de Casas, 47 mi E Cd.

Victoria, 500 m, vii.1986, carrion trap (PKC). Colima, 13 mi W

Pihuamo, along RõÂo Salido, vii.1982, in decaying tree (CDFA).

Veracruz, Santecomapan, vi.?; 16.2 mi N Puente Nacional,

vi.1962; Mexico, Jalapa (FMNH). MichoacaÂn, 8 mi E Apat-

zingaÂn, vii.1958 (DVC). Guerrero, 4.5 km SW Xochipala,

1400 m, vii.1992, thorn scrub (CMNC); Chilpaucingo, vii.1955

(FMNH). Oaxaca, 5 mi W Tequisistlan, 1100 ft, rotting cactus.

BELIZE: Orange Walk, Rio Bravo Cons. Area, La Milpa,

v.1997, FIT (PKC). GUATEMALA: Zacapa, 5 km SW RõÂo

Hondo, 300 m, vi.1993 (CMNC). EL SALVADOR: San

Salvador, vi.1958 (USNM); 10 km E La Libertad, v.?

(CMNC); 20 km E La Libertad (CMNC). NICARAGUA:

Masaya Laguna de Apoyo, xi.1991 (CMNH). COSTA RICA:

Guan., 32.2 km N Liberia, 50 m, v.1993 (SEMC); CanÄas,

vii.1966, vii.1988; Res. Biol. Lomas Barbudal, 30 m, vii.1991

(INBI); P.N. Sta. Rosa, 300 m, vi.1992 (INBI); 8 km SW

Cuajiniquil, Est. MurcieÂlago, vii.1993 (INBI); P.N. Palo

Verde, 10 m, v.1993, vi.1992 (INBI); Ref. Nac. Fauna Silv.

R.L. Rodriguez, 10 m, vi.1991, viii.1991 (INBI). LimoÂn,

Amubri, vii.1975 (EMUS). S.J. EscazuÂ, vii.1988 (EMUS).

PANAMA: sl. (BMNH). VENEZUELA: Edo. Aragua, Cuya-

gua; BolõÂvar, 26 km N Guasipati, vii.?, seasonal humid forest

(CMNC); Suapure, Caura R., viii.? (CUIC). TRINIDAD:

Bryant, ii.1903 (BMNH); Morne Bleu, 2700¢ (CMNC). CUBA:

Sierra Bonilla (FMNH). PUERTO RICO: Ensenada, i.1937.

DOMINICAN REPUBLIC: Independencia, 10 km NW La

Descubierta, Los Pinos, 487 m, carrion trap, xi/xii.1991

(CMNC); Prov. Hato Mayor, P.N. Los Haitizes, iv/vii.1992,

bosque humido (MAIC); Independencia, ESE JimanõÂ, La

Florida S of Lago LimoÂn, iv/vii.1992 (MAIC); Pedernales,

26 km N Cabo Rojo, 730 m, vii.1990 (CMNH).

Hister nodatus Lewis

Hister nodatus Lewis, 1888: 203.

Diagnosis. The presence of a supraorbital stria, prosternal

striae, 2 complete lateral pronotal striae and ¯attened or

depressed mandibles (Fig. 7C,F) separate H. servus and H.

nodatus adults from those of all other sympatric Hister. Adults

of H. nodatus are smaller than those of H. servus, lack the

anterior portion of the metasternal stria and have more strongly

depressed mandibles (Fig. 7f).


Average width 2.5 mm (range 2.1±3.1; n = 8). Frons smooth,

convex, slightly depressed behind laterally bisinuate frontal

stria; frontal stria follows contour of antennal fossa and inner

margin of eye, united with supraorbital stria dorsally; labrum

twice as wide as long, weakly emarginate apically. Mandibles

L

Fig. 8. A, Aedeagus of Hister servus, dorsal view. B, Aedeagus of

H. nodatus, dorsal view. C, Left elytron of H. servus. D, Left elytron

of H. nodatus.



short, concave above, margined dorsolaterally (Fig. 7F); short

stria visible along ventral margin of outer face. Antennal club

slightly oblong, with 3 complete sutures; submentum convex,

with 2 or 3 setae on each side of smooth medial area; mentum

wider than long, emarginate at apex, setose at sides; stipes with

3 or 4 setae; penultimate labial palpomere with 1 or 2 setae at

internal apex; terminal labial palpomere with few short setae.

Pronotum smooth; inner lateral stria complete, sinuate; outer

lateral stria nearly complete, slightly abbreviated at base;

prosternal keel striate, striae curved inward and meeting

posteriorly in most individuals; prosternal lobe long, apically

rounded, tapering sinuately to hypomeron. Protibia (Fig. 7D)

with 3 strong teeth plus a weak tooth basally; apical tooth with

2 denticles; small denticles also present on apical margin;

submarginal carina present, bearing a single basal denticle, and

uniting with margin at second tooth from base.

Elytra (Fig. 8D) lacking subhumeral striae; striae 1±3

complete; stria 4 complete or interrupted at middle; stria 5

absent or just visible at apex; sutural stria present in apical half

to two-thirds. Mesosternum shallowly emarginate, with com-

plete marginal stria; metasternum smooth, convex; anterior

portion of metasternal stria absent, the remainder extending

from near mesocoxa to metepisternum; meso-and metatibiae

slender, curved, with 2 rows of marginal spinules; basal meso-

and metatarsomere with one pair of ventral spinules.

Propygidium (Fig. 7E) depressed on each side, broad

epunctate area on lateral and anterior margins, punctures

longitudinally elongate, large; pygidial punctures deeply

impressed, especially on posterolateral margins; pygidial disk

raised medially, with less densely punctate area on each side.

Aedeagus (Fig. 8B) similar to that of H. servus, slightly

narrower overall; tegmen parallel-sided, dorsal lobes narrow,

meeting along midline; median lobe with thin, elongate

proximal apodemes; armature slightly curved apically.

Biology. Hister nodatus has been found in litter in a riparian

zone and near rotting palm fruit.

Remarks. What I treat as H. nodatus is a fairly widely

distributed entity, possibly containing independent lineages.

Specimens from Ecuador differ slightly in elytral striation and

have denser pygidial punctation though the aedeagus does not

differ from that of the Panamanian or Costa Rican populations.

Samples from the intervening areas will be necessary to

address the discreteness of this variation.

Holotype. Of undetermined sex: `Bugaba, 800±1500 ft,

Champion' [PANAMA] (BMNH); examined 1995. Other

material. (see Fig. 6): COSTA RICA: LimoÂn, Finca La Lola,

vii.1963 (LACM); LimoÂn, Amubri, 70 m, ii.1994 (INBI);

LimoÂn, Sector Cerro Cocori, Fca. de E. Rojas, 150 m, vii.1991

(INBI); LimoÂn, Hamburg Farm; Guan., Maritza Biol. Sta.,

650 m, v.1993 (SEMC); LimoÂn, Hamburg Farm; Guanacaste,

Santa Rosa, vii.1982; Puntarenas, Penin. de Osa, 200 m,

Rancho Quemado, v.1991, iv.1992 (INBI); Puntarenas, P.N.

Amistad, Est. Las Mellizas, 1300 m, v.1990 (INBI); Puntar-

enas, P.N. Corcovado, 100 m, vi.1990 (INBI); San JoseÂ, v.1928

(FMNH); Puntarenas Res. Biol. CararaÂ, Laguna Meandrica,

vi.1990 (INBI); Heredia, La Selva, 3.2 km SE Puerto Viejo,

100 m, v.1992 (SEMC). PANAMA: Canal Zone, Madden

Forest, rotting palm fruits, vi.1976 (FMNH); Barro Colorado

Island, viii.1994 (SEMC); Chepo, v.? (FMNH); Prov. Pan.,

Altos de MajeÂ, x.1975, Burlese litter along stream (PKC);

Prov. Pan., 6.9 km S Gamboa, 80 m, v.1995 (SEMC); ChiriquõÂ,

27.7 km W VolcaÂn, 1450 m, vi.1995 (SEMC); San Blas,

Nusagandi Res., 350 m, v.1995 (SEMC); ColoÂn, P.N. Sober-

ania, v.1995 (SEMC). VENEZUELA: Edo. Aragua, Cuyagua,

10 km E Ocumare, cacao pod litter, ii.1971 (FMNH).

ECUADOR: Pichincha, 47 km S Sta. Domingo, palm fruit

litter, ii.1976 (FMNH)(CNCI); Pichincha, 47 km S Sta.

Domingo, v/vii.1985 (CMNC).

Hister doyeni Caterino, sp.n.

Diagnosis. Among New World Hister possessing prosternal

striae, adults of H. doyeni may be separated by the absence of

the sutural elytral striae (Fig. 9D) in combination with the

complete outer lateral pronotal stria and basally depressed

mandibles.


average width 4.7 mm (range 4.4±4.8; n = 6). Elongate oval,

black and shining; frons broad, smooth, with weak depressions

behind stria on each side; frontal stria nearly carinate, straight

behind labrum, then weakly curving posteriorly to eye; labrum

tapered, straight to emarginate apically, about half as long as

wide; mandibles punctate throughout, weakly depressed

dorsoproximally, with small stria along ventral edge of outer

face; submentum acutely produced anteriorly, laterally setose;

mentum ¯at or with weak longitudinal furrow, weakly

emarginate apically, setose laterally; penultimate labial

palpomere with 4 or 5 setae along inner apex; terminal labial

palpomere with many very short setae; stipes with 8±10 setae.

Pronotal disk smooth at middle, punctate along base; outer

lateral stria united with transverse row of punctures along base;

inner lateral stria sinuate, complete behind head; marginal

pronotal stria also complete behind head or barely interrupted

at middle; prosternal keel with 2 striae divergent anteriorly and

posteriorly; prosternal lobe broadly rounded and deeply

punctate, marginal stria usually complete. Protibia triangular

(Fig. 13A), with 3 or 4 strong teeth, apical tooth with 2

denticles; submarginal protibial carina present at base, without

denticles, joining margin at basal-most tooth.

Elytra (Fig. 9D) lacking outer subhumeral stria; inner

subhumeral stria weakly impressed at apex; striae 1±3

complete, deeply impressed; stria 4 absent or barely visible at

base and/or apex; stria 5 and sutural stria absent. Mesosternum

broadly emarginate; meso- and metasternal disks ¯at and

smooth; metasternal stria absent along anterior metasternal

margin, but present from apex of mesosternal marginal stria,

curving laterad to middle of metepisternum; mesotibia weakly

curved, broad, with 2 rows of marginal spinules; metatibia

broad, almost straight, with 2 rows of strong spinules; meso- and

metatarsomeres short, ®rst tarsomere with 2 pairs of ventral

spinules, remaining tarsomeres with one pair.

Propygidium and pygidium both densely covered with

medium to large punctures, those of pygidium becoming

gradually smaller toward apex. Aedeagus (Figs 1, 9G) with

basal piece about one-fourth entire aedeagal length; tegmen

R



broad, tapered to apex, dorsal lobes slightly emarginate

medially; median lobe with proximal apodemes broad, abutting

along midline; armature about half tegmenal length, hooked

apically, with carina separating basal and apical portions;

median rods short, acute.

Distribution and biology. A few specimens (the largest

known series) were found in association with rotting Opuntia

cactus. No other habits are known. Collection records are

mapped in Fig. 11.

Etymology. This species is named for Professor John Doyen,

collector of one of the paratypes and an outstanding

coleopterist. He has provided much inspiration to my work.

Holotype. Female, dissected by the author: `MEXICO:

Puebla, 2 mi. SW TehuacaÂn, 5300', x.5±75, at light'/`J Powell,

J Chemsak, T Eichlin & T Friedlander' (EMEC). Paratypes

(5). One male, dissected by the author: `MEX: Oaxaca, 98 km

N. Huitzo, Guadelupe de los Ojos, 500 m, viii.19±58, J. Doyen'

(EMEC). One, `CacaloapaÂn, MeÂxico'/`Kil. #226; July 24 1956,

Thorn Forest, Vincent D. Roth, Coll.' (FMNH). 3, one male 2

females, Òaxaca, Mitla Vicinity, 56±6000 ft. ix.8.1973'/òn

rotting cacti, columnar/Opuntia, A. Newton'; one female in

MSCC, others in FMNH.

Hister fungicola Schaeffer

Hister fungicola Schaeffer, 1912: 27.

Hister nanulus Casey, 1916: 224. Lectotype, here designated,

male: `Florida'/ÙSNM 38440', examined 1995; Wenzel,

1939.

Diagnosis. Among adults of eastern U.S. Hister, those of H.

defectus, H. fungicola and H. indistincus are unique in having

prosternal striae in combination with the absence of sub-

humeral elytral striae. Adults of H. fungicola can be easily

recognized by the two complete anterior mesosternal striae,

present in no other known New World Hister.


average width 3.1 mm (range 2.8±3.3; n = 8). Frontal stria

round; frons with minute punctures; labrum about twice as

wide as long; mandibles convex above, weakly margined at

lower outer edge, punctate throughout; antennal club almost

circular, weakly acuminate apically, with 3 evenly spaced

sutures; prementum with 2 long, erect setae medially, shorter

setae laterally; mentum short, about two-thirds as long as wide,

weakly emarginate apically; stipes with 4 or 5 setae;

penultimate labial palpomere with single seta at inner apex,

ultimate segment sparsely covered with short setae.

Pronotum arcuate laterally, shiny with minute, scattered

punctures; inner lateral pronotal stria complete, sinuate at base;

outer lateral pronotal stria absent in basal half to two-thirds;

prosternal keel striate, striae meeting at base in some

individuals, inwardly arcuate, then diverging and apically

free; prosternal lobe short, rounded; prosternal marginal stria

complete, not strongly impressed. Protibia with 3 teeth, the

apical-most bispinose (Fig. 10A); a few denticles present along

apical margin; submarginal protibial carina complete with 1 or

2 minute denticles at base.

Elytra (Fig. 9A) lacking subhumeral striae; striae 1, 2 and 3

complete, fourth stria present in basal one-fourth to half, ®fth

L

Fig. 9. A, Left elytron of H. fungicola. B, Left elytron of H. indistinctus. C, Left elytron of H. defectus. D, Left elytron of Hister doyeni. E,

Aedeagus of H. fungicola, dorsal view. F, Aedeagus of H. defectus, dorsal view. G, Aedeagus of H. doyeni, dorsal view.



stria absent; sutural stria nearly complete. Mesosternum with 2

complete anterior marginal striae; meso- and metasternal disks

smooth medially, with medium-sized punctures laterally;

mesotibia and metatibia weakly curved, with 2 complete rows

of marginal spinules; all tarsomeres with one pair of ventral

spinules.

Propygidium with evenly scattered medium-sized punctures

separated by at least their widths; pygidium evenly covered

with ®ner punctures, about half the size of propygidial

punctures. Aedeagus (Fig. 9E) with basal piece about one-

fourth of entire aedeagal length; tegmen slightly tapered both

basally and apically, with broad, approximate dorsal lobes;

median lobe with proximal apodemes broad, abutting along

midline; median rods short, blunt; armature long, hooked

apically; gonopore situated near base of armature.

Biology. Hister fungicola is known mainly from low

elevation coastal plain localities in the eastern U.S.A. Speci-

mens have been found frequently in Pinus-dominated habitats.

A few specimens (including the type series) have been found in

direct association with rotting fungi (in one case, a Boletus), or

have been attracted to fungus-baited pitfall traps. Specimens

have also been attracted to malt and propionic acid-baited

traps.

Lectotype. Male, here designated (USNM # 42387):

`Jamesburg, NJ, Aug. 31'/`Brooklyn Museum Colln. 1929'/

`Cotype no. 42387 U.S.N.M.'/`Hister fungicola Schaef.'; ex-

amined 1998. Other material (see Fig. 12). U.S.A.: New

Jersey, Lakehurst (CASC); Jamesburg, Pine Barrens, Jenkins

Neck, viii.1974 (USNM). North Carolina, Southern Pines,

R

Fig. 11. Collection records of Hister defectus and H. doyeni.

Fig. 10. A, Left protibia of Hister fungicola. B, Left protibia of H. indistinctus. C, Left protibia of H. defectus.

Fig. 12. Collection records of Hister fungicola and H. indistinctus.



x.1921 (CASC), vii.1951 (CMNC), ix.1951 (CMNC); Eliza-

beth City. South Carolina, Georgetown, vi.1948 (USNM);

Horry Co., viii.1931 (CMNH); Berkeley Co. Florida, Liberty

Co.; MacClenny; Welaka; Glen St Mary, iv.1960 (FSCA);

Jupiter, vi.1966 (FSCA); Dunedin; 4 mi W Archer, iii.1960,

malt & propionic acid trap (CMNC); 3.6 mi N O'Brien,

iii.1956, malt trap (CMNC); Archer, iii.1960, iv.1960 (CNCI);

Ft Walton Beach, iii.1975; Highlands Co., Lake Placid,

Archibold Station, viii.1986, fungus pitfall (PSC); Archibold

Station, x.1984, on rotting Boletus (FMNH); Archibold

Station, vii.1983 (FMNH); Archibold Station, v.1979; Alachua

Co., Newnan's Lake, iv.1960 (FSCA); Alachua Co., 2.6 mi

NW Archer, x.1990 (FMNH); Alachua Co., Gainesville,

vii.1986, fungus; Alachua Co., v.1956, ii.1959, vi.1928

(FSCA); Alachua Co., Gainesville, vi.1959 (pine leaf litter),

viii.1962 (FSCA); Alachua Co., Gainesville, vi.1987, vii.1987,

viii.1986, v.1987 (PSC); Alachua Co., 2.6 mi SW Archer,

x.1990, sandhills (PSC); Levy Co., 3.8 mi SW Archer, vi.1988

(PSC); Marion Co., Rainbow Sprgs, vi.1960, in fungus

(FSCA); Marion Co., Ocala Nat. For., v.1956 (FSCA); Levy

Co., v.1956 (FSCA); Franklin Co., St George Is., Shell Point

area, vii.1972 (FSCA).

Hister indistinctus Say

Hister indistinctus Say, 1825: 35.

Hister debilicinus Casey, 1924: 197. Lectotype male, here

designated: `Southern Pines, NC'/`Casey bequest 1925'/

ÙSNM 38426'/`debilicinus Casey'; two paralectotypes,

same data as lectotype; examined 1995; Wenzel, 1939.

Diagnosis. Among adults of eastern U.S.A. Hister, those of

H. defectus, H. fungicola and H. indistincus are unique in

having prosternal striae in combination with the absence of

subhumeral elytral striae. Hister indistinctus may be

recognized by its single mesosternal stria and protibia with

the apical two denticles close together, usually abutting

(Fig. 10B).


average width 3.1 mm (range 2.8±3.4; n = 8). Frontal stria

variable, semicircular to semihexagonal; labrum rectangular to

emarginate, about half as long as wide; mandibles adentate,

convex above, usually with short striole at outer lateral margin;

antennal club almost circular, weakly acuminate at apex,

sutures more or less perpendicular to longitudinal axis of club;

submentum weakly convex, setose; mentum ¯at to weakly

convex, apically emarginate, laterally setose; penultimate

labial palpomere with single seta at internal apex; stipes with

4 or 5 setae.

Pronotum weakly punctate, row of punctures along basal

margin; 2 lateral striae, the inner complete at base and behind

head, outer stria usually slightly abbreviated at base; prosternal

lobe broadly rounded, punctate, extending laterally to

hypomeron; hypomeron bare. Protrochanter with single seta;

protibia (Fig. 10B) triangular, armed with 3 or 4 teeth, apical-

most tooth with 2 close-set denticles; submarginal protibial

carina present at base only, bearing a few small denticles and

joining margin near middle.

Elytra (Fig. 9B) lacking subhumeral striae; dorsal striae 1±4

complete and strongly impressed, ®fth stria restricted to apical

one-fourth, sutural stria in apical half; transverse metasternal

stria complete or nearly so anteriorly, not extending to

metepimeron laterally; meso- and metatibiae with inner edges

nearly straight, outer edges rounded with 2 rows of marginal

spinules; all tarsomeres with 2 ventral spinules.

Propygidium and pygidium densely punctate, punctures of

pygidium about half the diameter of those of the propygidium.

Aedeagus very similar to that of H. defectus (see Fig. 9F) with

basal piece short; tegmen widest at middle; proximal

apodemes of median lobe thick, approximate medially; median

lobe armature elongate, sinuate; dorsally articulated median

rods present, short.

Biology. A few specimens of H. indistinctus, like H.

fungicola above, have been found in direct association with

the fruiting bodies of fungi: Pleurotus sp., `¯eshy fungus', `gill

fungus', and `¯eshy gill fungus.' A few have been found on or

under the bark of dead trees, both Pinus and Quercus, and on a

sap¯ow of the latter. Only one specimen has been collected at a

carrion-baited trap, and none have been collected in associa-

tion with dung.

Type. The original Thomas Say type is believed lost; I have

followed the protocol of Mawdsley (1993) in selecting and

designating a neotype, the dif®culty of separating many of the

servus group species constituting an exceptional circumstance.

The neotype specimen was selected from the J. L. LeConte

collection in the MCZC. The neotype specimen bears an

orange (or brick red) circle indicating an original collection

site in the `Southern states.' There is also a label with the

number `879', of unknown signi®cance. Other material (see

Fig. 12). U.S.A.: Minnesota, Houston Co. Illinois, Mason Co.,

Sand Ridge St. Pk, under bark (FMNH); Mason Co., Sand

Ridge St. Pk, v.1989, oak tree hollow (PKC); Jersey Co.,

Principio College, iv.1943 (USNM). Indiana, Tippecanoe; Park

Co., Columbus; Warren Co.; Porter Co., Indiana Dunes Nat.

Lakeshore, Inland Marsh, v.1995, FIT (PKC); Jasper Co.,

Jasper Pulaski Wildlife Management Area, vi,vii.1995, mush-

room-baited pitfall (PKC). Pennsylvania, Jeanette, x.?

(CMNH). Maryland, Indian Head, v.1945, sap¯ow on white

oak (USNM). District of Columbia, Washington (USNM).

Kansas, Tanganoxie; Topeka, decaying mushrooms; Douglas

Co., 7.2 km W Lawrence, ix.1992, Omphalotus illudens

(SEMC); same locality, viii.1992, sap¯ow (SEMC). Missouri,

Warrensburg. Virginia, Lee Co., Jones' Ck (USNM). Oklaho-

ma, Latimer Co., 4 mi W Red Oak, iv.1988, beating Quercus

and under bark of dead Quercus (WFBM); Latimer Co.,

iv.1983 (FMNH); Latimer Co., vi.1983, ¯eshy fungus on

Quercus; Latimer Co., vi.1982, iv.1983, vii.1983, iii.1985,

iv.1985, iv.1988, vii.1988 (FSCA); Murray Co., ex Pleurotus

sp. Arkansas, Logan Co., v.1986, ¯eshy mushroom (CNCI).

Tennessee, Ky Lake, nr Model, viii.1968 (CNCI). North

Carolina, Southern Pines, vi.1916 (USNM); Tryon. Texas,

Kirbyville (USNM); Bastrop St. Pk, iv.1959 (CNCI); Mon-

tgomery Co., 7.2 km N Montgomery, v,vi.1987; Nacogdoches

Co., Nacogdoches, iii.1960 (PKC); Sabine Co., 9 mi. E

L



Hemphill, iv.1989, FIT, beech bottom (PKC). Louisiana,

Elizabeth, sticky traps in loblolly pine and under loblolly pine

bark; Rapides Parish, Pineville; Alexandria, xii.1966 (FSCA).

Mississippi, Lucedale, iv.1929 (CUIC). Georgia, Gainesville;

Stone Mt.; Atlanta. South Carolina, Florence; Clemson;

Meredith, iv.1926 (WSUC). Florida, Marion Co., 9 mi SSW

Ocala, v.1975 (FSCA); Baker Co., Glen St Mary, ii.1961,

under bark standing dead pine (FSCA); Crescent City

(USNM).

Hister defectus J.E. LeConte

Hister defectus J.E. LeConte, 1859: 312.

Hister permixtus Zimmerman, in J.L. LeConte, 1869: 252;

Schmidt, 1884c: 159. Lectotype male, here designated

(MCZC): `S.C.'/`890 Z.'/`H. permixtus Zim.'; examined

1998.

Diagnosis. See diagnosis for H. fungicola above. Hister

defectus differs from H. fungicola in having only a single

anterior mesosternal stria and from H. indistinctus in having

the two denticles on the apex of the protibia separated by

approximately the width of one denticle (Fig. 10C) and in

having the outer lateral pronotal stria abbreviated. Adults of H.

defectus are also smaller on average than those of H. fungicola

or H. indistinctus.


Average width 2.8 mm (range 2.7±3.0; n = 8). Frontal stria

rounded or semihexagonal; labrum about two-thirds as long as

wide; mandibles convex above (Fig. 2A), lightly punctate,

rufous at apices; submentum with 2 long setae, and several

short setae laterally; mentum emarginate apically, with weak

longitudinal carina, setose on anterolateral corners; penulti-

mate labial palpomere with 2 or 3 strong setae at internal apex;

terminal labial palpomere with many short scattered setae;

stipes with 4 setae, 3 along anterolateral margin, one on

internal margin; antennal club round, weakly acuminate

apically, sutures complete, evenly spaced.

Pronotum with sides inwardly arcuate; inner lateral stria

complete from base around anterior margin, often sinuate

basally; outer lateral stria varied, to almost absent, generally

free apically but united with marginal stria in some individuals.

Prosternal keel with 2 inwardly arcuate striae which may be

free or joined at base; prosternal striae ending free anteriorly in

most individuals; prosternal lobe evenly rounded, marginal

stria complete in most individuals, rarely obsolete at extreme

apex; prosternal lobe extended to hypomeron laterally;

hypomeron lacking setae. Protibia (Fig. 28) with 4 teeth,

apical 2 denticles separated by width of one denticle or more;

submarginal protibial carina present at base.

Elytra (Fig. 9C) lacking subhumeral striae; striae 1±4

complete; stria 5 present in apical one-third or less; sutural

stria present in approximately apical one-fourth to half.

Mesosternum broad, about 3 times as wide as long, anterior

marginal stria crenulate, outer marginal stria visible in each

lateral one-third; transverse metasternal stria crenulate, barely

interrupted at middle, laterally reaching almost to outer edge of

metacoxa then curved slightly inward; meso- and metatibiae

slightly curved, each with 2 complete rows of marginal

spinules, anterior stria almost as long as tibia.

Propygidium with evenly scattered medium-sized punctures

separated by greater than their widths; pygidium evenly

covered with punctures about half the size of those of

propygidium. Aedeagus as in H. indistinctus (Fig. 9F) but

slightly more elongate.

Biology. There are several records of H. defectus from

fungus, a preference shared with its close relatives, H.

fungicola and H. indistinctus. However, there are also a few

records from carrion or meat-baited traps.

Lectotype. Male, here designated (MCZC #34890): bears an

orange dot (indicating southern states) and `H. defectus Lec.';

examined 1998. Other material (see Fig. 33). CANADA:

Bousquet (1991) reports this species from Saskatchewan,

Ontario, and Quebec though I have not seen specimens from

these areas. U.S.A.: New York, reported in Mazur (1984).

Kansas, Lawrence. Missouri, Williamsville, viii.1958 (CNCI).

New Jersey, Ballinger's Mill, vii.1916 (USNM); West

Virginia, Lakehurst, W of Sulphur, vii.1914 (USNM).

Virginia, Fairfax Co., Great Falls, vi.1915 (USNM). Maryland,

Sparrow's Pt, v.1932 (CASC). Oklahoma, LeFlore Co., Riche

Mt., 1700¢, vi.1992 (PKC); Latimer Co., vi.1983, cray®sh bait

(FMNH); Latimer Co., xi.1977, ¯eshy gill fungus (FMNH);

Latimer Co. (numerous records from every month between and

including March±December, FSCA). Pittsburgh Co. (FMNH).

Texas, Brazos Co., College Station, ix.1962, in rotting

mushroom (PKC); Anderson Co., Salmon, vii.1975, fungus

(PKC). Arkansas, Washington Co.; Franklin Co.; Hope; Hot

Springs. Tennessee, Ky Lake, nr Model, viii.1968 (CNCI);

Burrville, vi.1954. North Carolina, Southern Pines, ix.1912

(USNM); Raleigh, x.1962 (CMNC); Davidson, on ¯eshy

fungus. South Carolina, Marion; Florence; Gable, vii.1933

(FMNH); Rocky Bottom; Walhalla (USNM); Clemson,

vii.1933 (FMNH). Louisiana, Pollack, vi.1983 (PKC). Mis-

sissippi, Lafayette Co., ix.1988, on mushroom (UMIC);

Calhoun Co., vi.1933, carrion pit trap (UMIC); Wilkinson

Co., Clark Ck Falls, viii.1982 (LSUC). Alabama, Blount

Springs; Alabaster, vi.? (FMNH). Georgia, Gainesville; Dallas,

vii.1942 (USNM). Florida, Alachua Co., rotten hamburger;

Columbia Co., O'Leno St. Pk, x.1987 (FSCA); Pensacola;

Alachua Co., Co. Rd 17b, 0.4 mi S Co. Rd 38, x.1983 (FSCA).

Hister lagoi Caterino, sp.n.

Diagnosis. Members of this species are most similar to those of

the comes/montivagus lineage; together the following char-

acters will separate it from those species: apex of protibia

somewhat rounded (Fig. 13B); pygidium sparsely and shal-

lowly punctate; outer pronotal stria abbreviated; sutural elytral

stria absent (Fig. 13E); mandibles slightly ¯attened dorsally;

aedeagus with subapical gonopore and with stout dorsal

projections (Fig. 13C).


average width 3.3 mm (range 3.2±3.4; n = 5). Oblong, body

widest at middle of elytron. Frons smooth, frontal stria

R



semihexagonal, moderately impressed; labrum trapezoidal,

truncate to weakly emarginate apically; mandibles short,

nearly ¯at above, laterally punctate, incisor edge about as

long as basal portion of mandible; antennal club oblong, widest

near base, sutures slightly oblique; submentum weakly convex

with a few setae along anterior margin; mentum ¯at, wide,

apically emarginate, laterally setose; penultimate labial

palpomere with 2 or 3 setae at apex; terminal labial palpomere

with many short setae; stipes with 5 or 6 setae.

Pronotal disk smooth, sparsely punctate along basal

margin; inner lateral stria complete, somewhat sinuate;

outer lateral stria present only in apical half to one-third of

pronotum. Pronotal keel striate, striae complete in most

individuals; broadly rounded prosternal lobe with complete

marginal stria extending laterally to hypomeron. Protibia

rounded at apical corner, bearing 5 or 6 denticles along

margin (Fig. 13B); submarginal carina visible at base, with

1 or 2 denticles, uniting with lateral margin at second or

third tooth from base.

Elytra (Fig. 13E) lacking subhumeral striae; striae 1 and 2

complete; stria 3 complete in most individuals, occasionally

interrupted brie¯y near apex; stria 4 visible at base and apex but

fragmented or interrupted at middle; stria 5 absent or

represented by a few punctures near apex; sutural stria absent.

Mesosternum broad, smooth; metasternum smooth; metasternal

stria nearly complete anteriorly, but narrowly interrupted at

middle, extending laterally to metepisternum though inter-

rupted in front of metacoxa in some individuals. Mesotibia

curved, apically expanded, bearing 2 rows of spinules; metatibia

slightly curved, with weak marginal spinules at base; all meso-

and metatarsomeres with single pair of ventral spinules.

Propygidium with shallow round evenly distributed punctu-

res separated by approximately their diameters; pygidial

punctures smaller, ®ner, more widely spaced, smaller and less

dense apically. Aedeagus (Fig. 13C) with basal piece approxi-

mately one-®fth entire aedeagal length; tegmen widest near

apex, dorsal lobes wide, slightly emarginate along inner

margin apically; median lobe with proximal apodemes broad,

abutting along midline; median rods, short, blunt; armature

about half tegmenal length, divergent, with oblique dorsal

carina; gonopore tube elongate, with gonopore arising at apex.

Distribution and biology. One of the paratypes was found in

pig dung, a few in oak and other leaf litter litter. However, the

overall scarcity of records in relatively well collected areas

suggests unusual habits for this species. Collection records are

mapped in Fig. 14.

Etymology. I have named this species for Dr Paul Lago, my

undergraduate mentor and a wonderfully enthusiastic entomol-

ogist.

L

Fig. 13. A, Left protibia of Hister doyeni, posterior view. B, Left protibia of H. lagoi, posterior view. C, Aedeagus of H. lagoi, dorsal view. D,

Aedeagus of a member of the H. comes complex, from Zacatecas, Mexico. E, Left elytron of Hister lagoi. F, Left elytron of the H. comes

complex, representative of populations from Oaxaca, Arizona's Pajarita Mts, and some members of the Chiracahuan, Huachuacan, Texan and

Durango, Mexico populations. G, Left elytron of some Chihuahua, Mexico members of the H. comes complex. H, Left elytron of some

Chiricahuan, Huachuacan and Zacatecas, Mexico populations of the H. comes complex.

Fig. 14. Collection records for Hister lagoi and the H. comes

complex.



Holotype. Male, dissected by the author: `TX: Blanco Co.

Pedernales Falls St. Pk 30-v.1985, R. Anderson, coll.'

(FMNH). Paratypes (7). One male, dissected by the author,

`TX: Burleson Co. Cooks Point, 11.vi.1981, leg. B. Summer-

lin, hog manure'/`BBS 19±81' (MSCC). One male, dissected

by the author, `Deland, Volusia Co. FL, 25 DEC 1985, M.

Deyrup'/òak litter vacant lot' (FMNH). One male, Àbilene,

25 mi. SW, Taylor Co., Tex. III::44'/`Col. & Pres. by Henry S.

Dybas'/`Collection of R.L. Wenzel' (FMNH). One male,

dissected by the author, `N. Braunfels, Tex.'/`Collection of

Chas. Schaffer'/`FMNH Ballou coll.'/`H. sp. nov.'/`Hister #27,

R.L. Wenzel 19__' (FMNH). One female, `TEXAS: Blanco

Co., Pedernales Falls St. Pk., vic. Twin Falls, 3-V-1988, R.

Anderson, mixed hdwd. berlese' (PKC). One male, `Florida:

Nassau Co., Ft. Clinch St. Pk., 8-V-1988; P. Skelley, leaf litter

berlese' (PKC). One male, TX: Houston Co., Big Slough Wild.

Area, FR517 & Four C's Hiking Trail, 9.v.88'/`R.S. Anderson,

bottomland hardwood litter berlese' (PKC).

Hister comes Lewis complex

Hister comes Lewis, 1888: 204.

Hister temporalis Fall, 1910: 120. Holotype male: `Baboqui-

varia Mts., AZ, F.H. Snow'; MCZC #24126; examined

1995. Syn.n.

Hister sternalis Casey, 1916: 223. Lectotype, here designated,

a female: `Durango, Dgo., Mex., Wickham'; USNM

#38438. Two paralectotypes, both males, same data as

lectotype; all examined 1997. Syn.n.

Diagnosis. Members of this variable species complex can

generally be recognized by the form of the pronotal striae with

the inner lateral stria complete and basally sinuate and the

outer lateral stria abbreviated and visible in only about the

apical one-third of the pronotum. Additionally, in most

individuals, the mandibles are depressed dorsobasally and the

meso- and metatibiae are triangular, relatively straight and

generally broad (rather than slender as in the similar

montivagus complex). The aedeagus of members of the comes

complex (see Fig. 13D) is characterized by separate, apically

acute dorsal lobes, long armature and short, thin proximal

apodemes. The tegmen is varied signi®cantly in general

proportions. Average length 4.0 mm (range 2.6±4.9; n = 9);

average width 3.5 mm (range 2.5±4.4; n = 9).

Biology. The largest series of H. comes have been taken

during excavations of woodrat (Neotoma sp.) nests. It seems

likely that this will prove to be an important habitat in some

areas. There are also records from rotting Opuntia cactus, fruit

and carrion.

Remarks. What is treated here as H. comes represents a

complex of closely related species and populations. The

complex is broadly distributed from Arizona, New Mexico and

Texas, through Mexico as far south as Oaxaca. There is

substantial variation across this range in genitalic and external

characters, much of which is relatively discrete and indicative

of some degree of reproductive isolation. One form is found in

localities throughout the range. It is relatively small (3±4 mm

in length), exhibits a densely puncatate pygidium and shows

some trace of a sutural stria (Fig. 13F). This combination of

states may represent a plesiomorphic condition from which

several populations have diverged. Specimens from Oaxaca

are quite small (» 3 mm) and have very slender meso- and

metatibiae. A series of specimens from Chihuahua (30 mi NW

of Cd Chihuahua), Mexico is larger than the average, ranging

from 4.5 to 5.0 mm, lacking a sutural stria and frequently

lacking a fourth elytral stria as well (Fig. 13G), with very broad

meso-and metatibiae, broad mandibles and a ®nely punctate

pygidium. Populations from the northern extent of the range, in

southern Arizona and New Mexico are varied in most of these

characters, variation evident even in series from individual

localities. In one area, near the South-western Research Station

in the Chiricahua Mts of Arizona, the so-called plesiomorphic

form co-occurs with a form lacking sutural striae (Fig. 13H)

and without any pygidial punctation whatever. It is possible

that these represent independent lineages. However, this is

dif®cult to assess with existing data. To further complicate

matters, phylogenetic analyses suggest that this complex is

paraphyletic with respect to the montivagus/bullatus lineage.

Isolated species will be distinguished in this complex.

However, at present, the most distinct populations (e.g. those

from Oaxaca or Chihuahua) are represented by only single

series far from other collecting localities and, in my view, their

formal description would be premature.

Holotype. Female, `Tula, Hidalgo' [MeÂxico] (BMNH);

examined by M.S.C., 1995. Other material. (Fig. 14). U.S.A.:

California, San Bern. Co., Mitchell Caverns St. Pk, 4400 ft, iii/

iv.1979 (DVC). Arizona, Yavapai Co., i.17, 2 mi S AZ Hwy.

179, iv.1987, Neotoma nest; Chiricahua Mts, Cave Ck Cyn;

SWRS, nr Portal, viii.1961 (EMEC); SWRS, vi.1950, viii.1955

(CMNC) Chiricahua Mts, 4000 ft, viii.1969 (FMNH); Chir-

icahua Mts, Rucker Cyn, 6000 ft, vii.1979 (CNCI); Chiricahua

Mts, nr Ash Spring, 6300 ft, vii.1979 (CNCI); Chiricahua Mts,

South Fork Cyn, vii.1965 (FSCA); Chiricahua Mts, Idlewild

Cmpgrd, 6000 ft, vii.1972 (USNM); Chiricahua Mts, Stewart

Forest Camp, ix.1952 (FMNH); 5 mi W Portal, ix.1976, pit trap

w/dead snake (CDFA); Portal, viii.1985, fermenting fruit

(CDFA); 2 mi E Portal, vii.1989, Neotoma nest under Prosopis

(CDFA); 2 mi NE Portal (CNCI); Portal (CNCI); Portal,

viii.1906 (USNM); Chiricahua Mts, Paradize, 4700 ft,

viii.1978, Neotoma refuse (DVC); Chiricahua Mts, Ranger

Station at entrance to Cave Ck Cyn, vii.1974, Neotoma

albigula dung chamber (DVC); Pajarito Mts, Sycamore Cyn,

viii.1968 (FMNH); Pajarito Mts, Pena Blanca Canyon,

viii.1972 (USNM); Patagonia Mts; Sta Rita Mts, » 8000 ft;

Sta Rita Mts, Madera Cyn, vii.1962 (DVC); Huachuaca Mts

(FMNH); Douglas, ix.1957 (USNM); Pima Co., Old Tucson,

iv.1971 (USNM); Pima Co., Tucson Mts, iv.1969 (USNM);

Leslie Cyn, 16.5 mi N Douglas, 4700 ft, vii.1976, fruit trap

(BMNH); Whetstone Mts, Cottonwood Cyn, viii.1978 1670 m

(FMNH); Sta. Cruz Co., 4 mi SW Patagonia, 4050 ft, viii.1979

(CNCI). New Mexico, Hidalgo Co., 1 mi W Rodeo, viii.1971

(EMEC); Hidalgo Co., 2 mi NE Rodeo, vii.1989, Neotoma nest

under mesquite (CDFA); Eddy Co., 35 mi SW Carlsbad,

6400 ft, carrion, vii.1975 (CNCI). Texas, Big Bend Nat. Pk,

4000 ft, iv.1959 (CNCI); Big Bend Nat. Pk, 5000 ft, v.1959

R



(CNCI); Big Bend Nat. Pk, Persimmon Gap, v.1959 (CNCI);

Big Bend Nat. Pk, 7500 ft; Ft. Davis; Uvalde Co., BFS Cave,

vi.1985 (PKC). MEXICO: Chihuahua, Majalca Rd, 30 mi NW

Chih., iv.1961 (CNCI); Mesa Del Huracan, 7400 ft, vii.1964

(CNCI). Durango, 5 mi W Durango, 6300 ft, vi.1971 (CMNC);

25 mi W Durango, vii.1964 (CNCI); 44 km SE Durango,

x.1990 (PKC); 4 mi N Tepeapulco, vi.1975 (PKC). Hidalgo,

4 mi N Tepeapulco, vi.1975 (FMNH); Tula. Guanajuato,

viii.1949. Zacatecas, 9 mi S Fresnillo, vi.1956, viii.1954

(EMEC); Puebla, 5 mi N Tehuacan, 6000 ft, Acacia scrub,

vii.1982 (MSCC); 6.5 km N Azumbilla, 2250 m, viii.1987

(EMEC). Veracruz, 19 km NW Cd. Mendoza 1900 m, viii.1987

(EMEC). Oaxaca, Mitla vicinity, rotting columnar/Opuntia,

5600±6000 ft, ix.1973 (FMNH).

Hister montivagus Lewis complex

Hister montivagus Lewis, 1888: 203.

Diagnosis. Members of this species complex are generally

small-bodied (< 3.5 mm). They can be distinguished by the

following characters: the fourth dorsal elytral stria inter-

rupted (impressed only basally and, for a shorter distance,

apically; Fig. 18A) pygidial punctures evenly, densely

spaced; frons with strongly impressed, sinuate frontal stria;

L

Fig. 15. A, Right protibia of the Hister montivagus complex, posterior view. B, Left protibia of H. diadema, posterior view. C, Pygidium of H.

diadema, typical of the three species of the H. bullatus clade.

Fig. 16. A, Aedeagus of Panamanian and Costa Rican populations

of the Hister montivagus complex. B, Aedeagus of H. bullatus,

dorsal view. C, Aedeagus of H. diadema, dorsal view.

Fig. 17. Collection records for the Hister montivagus complex.



apical tooth of protibia bearing three closely set denticles

(Fig. 15A). They are similar to, though smaller than, more

derived members of their lineage (the species H. bullatus,

H. coronatus and H. diadema) in general strial characters

and aedeagal morphology. The irregularly spaced punctation

of the pygidium of members of those species easily

separates them from the montivagus complex. Average

length 3.3 mm (range 2.3±4.6; n = 9); average width 2.8 mm

(range 1.9±3.6; n = 9).

Biology. Specimens of this complex have been found in a

wide variety of situations; many have been burlesed from leaf

litter. There are also records from rotting fruit and from dung

and carrion traps.

Remarks. Additional collections are needed to address the

taxonomic status of many of the populations comprising this

complex. Series from most localities are separately diagnosible

to some degree (see Table 1 for a synopsis of populations

treated in cladistic analysis). Yet most sampled populations are

geographically widely separated. A survey of variation in the

intervening areas is needed. Characters which exhibit sub-

stantial variation include: length of aedeagal basal piece, shape

of dorsal tegmenal lobes, shape of median lobe armature,

elytral striation, body size and propygidial and pygidial

punctation. A better understanding of the variation among H.

montivagus populations is also essential in understanding its

relationships to the related comes and bullatus/coronatus

lineages.

Lectotype. Here designated: `V. de ChiriquõÂ [PanamaÂ], 25±

4000 ft, Champion' (BMNH). Two paralectotypes, same data

as lectotype (BMNH, FMNH); examined 1995. Other material

(Fig. 17). MEXICO: Tamaulipas, GoÂmez FarõÂas, 1000 m, vi/

viii.1983, cloud forest (CNCI). San LuõÂs PotosõÂ, El Salto del

Agua, vi.1975 (PKC). Veracruz, Canyon RõÂo Metlac, 3200 ft,

leaf litter, viii.73 (FMNH); Canyon RõÂo Metlac, 3200 ft,

viii.73, rotting fruit (FMNH); FortõÂn de los Flores, vi.1963

(USNM); Presidio, 1000 ft, vii.1965 (LACM); 3.5 km S Jalapa,

1370 m, v.1991 (SEMC). Hidalgo, 2 mi N Tlanchinol, 5200 ft,

fermenting oak litter, vii.? (FMNH); 4.4 km N Tlanchinol,

1420 m, vii.1992 (SEMC). MeÂxico, 2 mi NE Tenancingo, pine/

madrone/oak, 7100 ft, ix.1973 (FMNH). Morelos, Chapulte-

pec, 1400 m, vi.1942 (FMNH). Jalisco, Nevada de Colima,

6800 ft, ix.1973, pine-oak forest. Oaxaca, 20 mi S Juchatengo,

6000 ft, v.1971, carrion and human dung traps (CNCI);

38.4 km N Telixtlahuaca 1800 m, vii.1992, oak-palmetto

woodland (SEMC); 14.9 km N Sola de Vega 1820 m,

vii.1992 (SEMC). Chiapas, P.N. Sumidero, Coyota Mirador

1700 ft vi.1989 (CMNC); P.N. Sumidero, 1000 m, v/vi.1990

(CMNC); Lago de Calores, vi.1969, vii.1982 (CMNC).

GUATEMALA: Baja VerapaÂz, 8 km S Purulha, 1660 m,

v.1991 (CMNC); Zacapa, 3.5 km SW La UnioÂn, 1500 m,

vi.1993, cloud forest (CMNC); Zacapa, La UnioÂn, 1500 m,

vi.1993 (SEMC); Sacatepequiz, 4.5 km SW San Miguel

Duemas 1760 m, vi.1991 (SEMC); Env. de Guatemala

(MNHN). BELIZE: Orange Walk, RõÂo Bravo Cons. Area, La

Milpa, ix.1995 (PKC). EL SALVADOR: Los Chorros, 4 km S

Santa Tecla, v.1971, ravine litter (CNCI). HONDURAS:

Lempira, 13.1 km NE and 8.3 km E Gracias, Cerro Puca,

1600 m, vi.1994, Liquidambar litter (CMNC); Olancho, P.N. La

Muralla, 14 km N La UnioÂn, 1450 m, viii.1994, wet evergreen

forest (CMNC). COSTA RICA: Guanacaste, SW side VolcaÂn

Cacao, P.N. Guan., 1000±1400 m, v.1992, vi.1990, viii.1991

(INBI); Est. Las Pailas, P.N. RincoÂn de la Vieja, vii.1992

(INBI); Puntarenas, Monteverde, 1400 m, v.1979, viii.1987

(CMNC); Puntarenas, Monteverde, v.1989, vii.1990 (MSCC);

Puntarenas, Monteverde, 1000±1350 m, vi.1994 (INBI); Pun-

tarenas, P.N. La Amistad, Est. Las Mellizas, 1300 m, iii.1990,

v.1990, vii.1990 (INBI); Alajuela, Zarcero, Alfaro RuõÂz 1700 m,

vi.1990 (INBI); San JoseÂ, EscazuÂ, vi.1989 (EMUS). PANAMA:

ChiriquõÂ, 4 km N Sta. Clara, 1500 m, vi/vii.1981 (CMNC);

ChiriquõÂ, 2 km N Sta Clara, v.1977, oak/coffee litter (FMNH);

ChiriquõÂ, Cerro Pelota, 1500 m, vi/vii.1982 (CMNC); ChiriquõÂ,

VolcaÂn de ChiriquõÂ, 4000±6000 ft (BMNH); PanamaÂ, Chepo-

Harti Rd, 400 m, vii.1982 (CMNC); ChiriquõÂ, 20 km N Gualaca,

1350 m, v,vi.1995 (SEMC); ChiriquõÂ, 27.7 km W VolcaÂn,

1650 m, vi.1996 (SEMC).

Hister bullatus Lewis

Hister bullatus Lewis, 1888: 203.

Diagnosis. The irregularly spaced pygidial punctation

(Fig. 15C) readily separates H. bullatus, H. coronatus and H.

diadema from all other New World Hister. Of these H. bullatus

is only sympatric with H. coronatus, which lacks prosternal

striae. In H. bullatus these may be abbreviated anteriorly but

are always present.

R

Fig. 18. A, Left elytron of the Hister montivagus complex (in few

populations the 4th dorsal stria is complete). B, Left elytron of H.

bullatus. C, Left elytron of H. coronatus.




average width 3.6 mm (range 3.1±4.0; n = 4). Elongate oval;

frons smooth, depressed behind bisinuate frontal stria; labrum

two-thirds as long as wide, slightly tapering towards apex;

mandibles slightly depressed dorsoproximally, with short stria

at ventral edge of outer face; submentum laterally setose;

mentum as long as wide, shallowly emarginate at apex; setose

at sides; stipes with 4 or 5 setae; antennal club large, with

slightly oblique sutures.

Pronotal disk smooth; inner lateral stria crenulate and

complete; outer lateral stria present only in apical half to one-

third; prosternal keel with striae abbreviated anteriorly in some

individuals; prosternal lobe rounded, punctate, laterally

narrowed but reaching hypomeron. Protibia rounded, teeth

weakly developed, with 7 or 8 denticles along lateral and

apical margins (Fig. 15B); submarginal protibial carina pre-

sent, with 2 or 3 denticles near base, united with margin near

second tooth from base.

Elytra (Fig. 18B) lacking subhumeral striae; striae 1±3

complete in most individuals; stria 3 more weakly impressed

near apex; stria 4 interrupted at middle, present in basal one-

third and apical one-®fth; stria 5 absent or barely visible at

apex; sutural stria present in apical half to two-thirds.

Mesosternum broad, marginal stria complete, outer stria very

short; metasternal disk smooth, marginal stria complete

anteriorly, crenulate, recurved forward laterally to basal one-

third of metepisternum; meso- and metatibiae weakly curved,

with 2 complete rows of marginal spinules.

Propygidium depressed laterally, raised at lateral edges

(Fig. 15C); punctures large, dense, shallow, round to poly-

gonal; pygidium with irregularly spaced deep round punctures.

Aedeagus (Fig. 16B) with basal piece between one-fourth and

one-third entire aedeagal length; tegmen parallel-sided;

gonopore-bearing tube elongate, gonopore situated near apex

of tegmen; median lobe armature short, divergent, dorsal lobes

of tegmen broad, inner margins straight, approximate.

Biology. Nothing is known of the habits of this species.

Holotype. Male, `Capetillo, Guatemala, C. Champion'

(BMNH); examined by M.S.C., 1995. Other material

(Fig. 19). MEXICO: Tamaulipas, GoÂmez FarõÂas, viii.1983

(CNCI). GUATEMALA: Chimal, S.P. Yepocapa, v.1948,

4800¢, under fungus-covered log (FMNH). HONDURAS:

Olancho, 14 km N La UnioÂn, P.N. La Muralla, viii,ix.1994

(CMNC).

Hister coronatus Marseul

Hister coronatus Marseul, 1861: 545.

Diagnosis. Members of this species are unique in the H. servus

group in that they lack prosternal striae. They differ from all

other such Hister by their irregularly spaced pygidial

punctation (Fig. 15C) and pattern of elytral striae (Fig. 18C).


average width 4.2 mm (range 3.5±4.7; n = 8). Adults of this

species differ from those of H. bullatus only in the following

characters: body size larger; supraorbital stria present;

prosternal keel lacking striae; prosternal lobe narrower;

protibial teeth well developed; outer lateral pronotal stria

visible in apical one-third or less; elytral striae with 1st and

2nd striae complete, 3rd stria interrupted at middle, 4th stria

short, just visible at elytral base, 5th and sutural striae absent

from most individuals; propygidial punctures small, more

widely separated. Aedeagus identical to that of H. diadema

(Fig. 16C).

Biology. There is nothing known of the habits of this

species.

Types. The original type may be lost. It could not be

located by R. L. Wenzel (personal communication) nor by

me in the Paris Museum. R. L. Wenzel (personal

communication) has labelled two specimens in the B.C.A.

collection (BMNH) as a lectoholotype and lectoparatype,

though these designations were never published. The data

on the specimens is consistent with that of the original

description. Marseul (1861) cited the type data as `MexõÂque

(SalleÂ).' The specimens bear the data `Cordova, Mex.

[southern Veracruz], Salle, coll.' I hereby formalize

Wenzel's lectotype designations though the possibility exists

that an original Marseul type may turn up in the Paris

Museum of Natural History. Other material (Fig. 19).

GUATEMALA: Chimal, S.P. Yepocapa, 4800 ft, v.1948

(FMNH). COSTA RICA: Guanacaste, P.N. Sta. Rosa,

300 m, vi.1992 (INBI); Guanacaste, 35 km SE RõÂo Naranjo,

xi.1991 (EMUS); Guanacaste, Vol. Rincon de la Vieja, S.

Slope, vi.1993 (MSCC); LimoÂn, Res. Biol. Hitoy Cerere,

100 m, vii.1991 (INBI); LimoÂn, Sector Cerro Cocori, 150 m,

vii.1993 (INBI); Puntarenas, Res. Biol. CararaÂ, Est.

Quebrada Bonita, 50 m, vi.1993 (INBI). PANAMA: reported

in Mazur (1984).

Hister diadema Marseul

Hister diadema Marseul, 1854: 559.

Hister striatipectis Lewis, 1899: 16. Holotype, probably male:

`Chapada Forest'[BRAZIL]/`G.Lewis coll. 1926±369'/

`H.H. Smith 1898'/`Nov.' (BMNH); examined 1995. Syn.n.

L

Fig. 19. Collection records of Hister bullatus and H. coronatus.



Diagnosis. Adults of this species are very similar to those of H.

coronatus but differ in having prosternal keel striae and a

sutural stria in the apical half to one-third of the elytron. They

exhibit irregularly spaced pygidial punctures (Fig. 15C) similar

to those found in H. bullatus and H. coronatus. The

distribution of H. diadema, however, does not appear to

overlap with that of these other species.

Description. Average length 5.1 mm (range 4.6±5.5;

n = 4); average width 4.8 mm (range 4.3±5.1; n = 4).

Individuals differ from H. bullatus in the following

characters: body size larger; supraorbital stria varied,

complete or interrupted; outer lateral pronotal stria short

as in coronatus, but closer to pronotal margin, in some

individuals with additional strioles posteriorly; protibial

teeth well developed; apical fragments of inner subhumeral

elytral striae visible in some specimens. Aedeagus as in

Fig. 16(C). Some specimens exhibit more than one pair of

prosternal keel striae (those formerly designated as H.

striatipectis). However, this appears to represent only

intraspeci®c variation.

Biology. Nothing is known of the biology of this species.

Holotype. Probably female, `Cayenne' [French Guiana]

(MNHN); examined by MSC 1995. Other material

(Fig. 20). BRAZIL: Sta. Catarina, Nova Teutonia, x.1950,

300±500 ft (FMNH); Para, Barcarena, vi.1991 (MSCC);

Amazonas, Pobas (MNHN); SaÄo Paulo (MNHN); Chapada

Forest (BMNH). BOLIVIA: Region ChapareÂ, i.1950, 400 m

(FMNH).

Phylogenetic analysis and results

Parsimony searches resulted in 395 equally parsimonious

cladograms of 544 steps (CI = 0.494; RI = 0.703). The majority

of these alternatives result from various resolutions among the

H. comes/montivagus OTUs. The resulting consensus clado-

gram (Fig. 21) is otherwise quite well resolved. The consis-

tency index indicates substantial homoplasy, though

subsequent analyses with two representatives each of the

comes and montivagus complexes resulted in slightly higher

internal support (CI = 0.525). Individual character consistency

indices are given in Table 2.

The basal topology does not re¯ect presupposed relation-

ships among the ingroup and the outgroup taxa. The

monophyly of Spilodiscus + H. sallei + H. matador hypothe-

sized by Caterino (1998) is not supported by any of the most

parsimonious cladograms. Forcing this trio to be monophyletic

at the base of the servus group requires an additional twelve

steps. However, constraining this group to monophyly basal to

the putative outgroup taxa (H. curtatus, H. dupurator and H.

civilis) requires only one additional step. Though Spilodiscus

and the sallei/matador lineages share a nearly identical

aedeagal morphology, their subtending branches are quite

long (c. ten changes each; not shown for Spilodiscus).

Spilodiscus, in particular, is highly divergent in external

morphology and the two lineages differ in numerous characters

of elytral and sternal striation. If these groups are, in fact,

closely related, phylogenetic signal in the genitalic characters

is being obscured by external autapomorphies. Additional

characters will be needed for further clari®cation.

There is good support for the monophyly of the remainder of

the ingroup (node c). This clade is found in all most

parsimonious reconstructions and is supported by ®ve

character transitions, including the gain of an external

mandibular stria (ch. 5), a reduction in tarsal setae from three

or more on the basal tarsomere to two (ch. 21), a transforma-

tion of the lateral tegmenal cleft from slit-like to socket-like

(ch. 27) and a shortening of the armature of the aedeagal

median lobe (ch. 30). Bootstrapping the entire data set gives

only slightly greater than 50% support. However, bootstrap

runs in which only one to two representatives each of the

montivagus and comes complexes were included increased this

support to 60±70%.

Within the restricted servus group, there are three main

lineages, each of which is relatively well supported. A

group of four species above node f (H. doyeni, H. defectus,

H. indistinctus and H. fungicola; the defectus lineage for

the purposes of this discussion) is present in all most-

parsimonious reconstructions and is supported by two

aedeagal synapomorphies, dorsal projections present and

stout (ch. 29), and proximal apodemes thick, abutting (ch.

31), a loss of the basal row of protibial setae (ch. 11), and

a change in elytral striation (ch. 16). A lineage comprising

H. servus and H. nodatus (node e) is likewise found in all

most parsimonious reconstructions, with 86% bootstrap

support and a large decay index. These species are united

by six characters, including two aedeagal characters: dorsal

tegmenal lobes narrow and medial (ch. 26) and median

R

Fig. 20. Collection records of Hister diadema. The open circle in

Amazonas represents a state record for which the speci®c locality,

Pobas, could not be located.



lobe armature divergent and lacking an oblique carina (ch.

30). The third major lineage comprises all of the OTUs

above node i, found in 88% of most parsimonious

reconstructions. All possible resolutions of these three main

lineages are equally parsimonious with no one resolution

supported by > 50% bootstrap replicates. The resolution

shown in Fig. 21 was chosen because it appeared in 62% of

all most parsimonious reconstructions.

Relationships within these groups can be resolved with

considerably greater certainty. Relationships among the four

OTUs of the defectus lineage (node f) are supported as shown in

the majority of most parsimonious reconstructions. The

inclusion of H. doyeni is not supported by bootstrapping.

However, character support for its inclusion, particularly the

thick proximal apodemes (ch. 31), is strong. Although H.

defectus and H. indistinctus are not found as sister species in all

reconstructions (thus only 51% consensus support), no

con¯icting resolution was found; the less than unanimous

support results from a trichotomy frequently reconstructed

among H. indistinctus, H. defectus and H. fungicola. None-

theless, H. indistinctus and H. defectus share a nearly identical

aedeagal morphology and their sister species status seems

likely.

Monophyly of the comes/montivagus lineage (all OTUs

above node i) is supported by one unambiguous synapomor-

phy, the elongated gonopore tube (ch. 32). The basal position

of H. lagoi seems relatively well founded; the monophyly of

the remaining OTUs is supported by two characters: mandibles

¯at or depressed above (ch. 4) and dorsal tegmenal lobes

narrowed, separate medially (ch. 26). Little resolution is

supported among members of the comes and montivagus

complexes. Maximum parsimony supports neither complex as

monophyletic, though a subset of the populations of H.

montivagus appears to be. Instead the comes complex appears

to be paraphyletic with respect to group including the

montivagus complex and the bullatus/coronatus/diadema

lineage (node l). The montivagus complex, in turn, appears

to be paraphyletic with respect to the bullatus/coronatus/

diadema lineage. While one of the comes populations appears

to be basal to the remainder of this lineage, node k is only

supported by one character, the tegmen's lateral surface being

longitudinally concave. In light of the considerable variation in

tegmenal shape in the comes complex, I would not consider

this strong evidence for a basal position of comes3. In the case

of the montivagus complex, the resolution among its popula-

tions obtained is similarly suspect. Several montivagus OTUs

are represented only by females (montivagus1, 2 and 3). An

L

Fig. 21. The majority rule consensus of 395 cladograms of length 544 (C.I. = 0.494). Numbers above the branches indicate majority rule

consensus indices/decay indices/bootstrap support. Decay indices were calculated excluding all but two populations each of the H. comes and H.

montivagus complexes. Bootstraps were calculated for the entire data set over 100 replicates. Numbers below the branches indicate the characters

supporting that branch. Known ranges to the right of the cladogram are approximate. C.R. = Costa Rica; S. Am. = South America; C.

Am. = Central America; Pan. = Panama; Guat. = Guatemala; S. M. = Sierra Madre de Mexico (Or. = Oriental; Occ. = Occidental).



assessment of the male's genitalic states might provide

additional resolution. These results suggest that the montivagus

complex is paraphyletic with respect to a clade composed of H.

bullatus, H. coronatus and H. diadema. However, establishing

which montivagus populations are more closely related to the

bullatus clade is not possible with these data, though a few

characters, including spermathecal shape (ch. 34), and

propygidial shape (ch. 22), are suggestive.

The monophyly of the three species of the H. bullatus clade

is supported by all analyses (68% bootstrap support; decay

index of six steps), though resolution among them is not clear.

A clade comprising H. coronatus and H. diadema seems most

likely. Both of these are larger in size than either H. bullatus or

any of the possible H. montivagus progenitors. Additionally,

both of these species possess a supraorbital stria in at least

some individuals, whereas no specimens of H. bullatus exhibit

this (nor do any H. montivagus populations.) Geography would

also seem to favour a coronatus/diadema lineage (see Figs 19,

20). The data set including only two populations each of H.

comes and H. montivagus supports the monophyly of

coronatus/diadema in all most parsimonious reconsructions

at a decay level of ®ve steps. In the full analysis, however, that

resolution requires one step more than a bullatus/coronatus

lineage. At any rate the three species are clearly very closely

related. Additional data will be necessary to resolve their

interrelationships.

Discussion

Several interesting biogeographic situations emerge from

examining the distributions of the H. servus group species in

light of the cladogram. The approximate distributions of the

OTUs are presented with the cladogram in Fig. 21. Areas of

endemism follow Liebherr (1994), with the addition of areas

for species with U.S.A. and South American distributions.

Many of the taxa in this analysis share areas and it is probably

inappropriate to try to look for one vicariance history to

explain the entire group's distribution. Thus, I have not tried to

derive a fundamental area cladogram for all of the included

taxa. Nonetheless, several disjunctions should be noted.

An interesting disjunction is observed between H. doyeni

and the more derived members of the defectus lineage. Hister

doyeni is known only from the highlands of Oaxaca and

southern Puebla, while the other members of the defectus

lineage are resticted to the eastern and south-eastern U.S.A. It

is a clean distributional break with no known intervening

records for any of the species. A somewhat similar pattern is

described by Noonan (1988) but, in all cases, phylogenetically

intermediate taxa are known from north-eastern Mexico.

Future collectiong in north-eastern Mexico may reveal

additional members of this clade.

An interesting recurrent trend is related (sister) taxa

differing in habitat preference, with several phylogenetic

disjunctions between drier, seasonal, subtropical habitats and

moister, tropical areas. This occurs in the following lineages:

comes/montivagus, defectus/doyeni, matador/sallei and servus/

nodatus. In the case of the comes/montivagus lineage, the

comes complex is restricted to semiarid areas of the U.S.A. and

western Mexico while the more derived montivagus/bullatus

lineage is found primarily in eastern Mexico south throughout

the moist highlands of Oaxaca and Central America. Interest-

ingly, the most basal montivagus population occurs in Oaxaca,

one of the few areas where H. montivagus and H. comes are

sympatric, a possible indication of a southern vicariance. This

is broadly consistent with the general area hypothesis of

Liebherr (1994) in which the drier areas of western Mexico and

the southern U.S.A. were found to be biogeographically

paraphyletic with respect to the moister tropical areas of

Mexico and Central America. A more thorough understanding

of relationships among these complexes may provide an

excellent system for examining relationships among these

diverse biogeographic regions.

The occurrence of only two of the species of the H. servus

group (H. diadema and H. nodatus) in South America is

interesting and merits some discussion. Because these species

show very different distributional patterns in South America it

might be argued that they arrived there via different pathways

or at different times. Hister nodatus and H. sallei appear to

have experienced the Andes as a barrier, not having penetrated

further into the continent than their western and northern

foothills. While this suggests recency of colonization, H.

nodatus has differentiated in Ecuador and its occurrence there

may well date to the earliest modern connection between

Central and South America (3±5 Ma; Donnelly, 1985). Hister

diadema, on the other hand, has been recorded only from the

southern and south-western fringes of the Amazon basin,

suggestive of a pre-Andes colonization.

The most fertile ground for future studies in this group will

be found in the interrelationships of the comes and montivagus

lineages. The strong morphological continuity and concomi-

tant dif®culty of reconstructing cladistic relationships within

each of these complexes indicates the possibility that they each

may indeed have some level of genetic continuity. Yet the

derivation of the bullatus clade from within this complex is

strongly supported by these data. An exploration of the

distribution of the genetic diversity underlying such a situation

would be very interesting. Furthermore, an understanding of

the historical and present connections within and between

these might enhance our understanding of recent Mexican

biogeography.

Ultimately, the ambiguities in the results presented here

may stem from the lack of an appropriate outgroup to the

servus lineage. A reanalysis with a clearer idea of outgroup

relationships will be necessary to determine, for example,

whether the sallei/matador and/or Spilodiscus linages are

closely related to the H. servus group or not, or, indeed, if

they are closely related to each other. Nonetheless, the

monophyly of the H. servus group (without H. sallei, H.

matador or Spilodiscus) is supported by this analysis, and

the group may appropriately serve as a terminal OTU for

future studies aimed at understanding higher level relation-

ships within Histerini. While such a study is not presently

possible, a continued focus on species level revision which

identify and characterize monophyletic lineages will even-

tually make it attainable.

R



Acknowledgements

I would like to thank A. Newton, R. Wenzel and P. Parrillo of

the Field Museum of Natural History for their assistance

during a visit to the museum in 1995. The Field Museum's

Thomas J. Dee fund also helped make that trip possible. The

CanaColl Foundation provided funds for a visit to the

Canadian National Collection where I received valuable

assistance from D. Bright, Y. Bousquet, A. Smetana and S.

LaPlante. Sigma Xi provided funds for studies at BMNH and

MNHN. Numerous additional persons and institutions pro-

vided loans which made this study possible: C. Barr (EMEC),

D. Kavanaugh and R. Brett (CASC), D. Furth, G. Hevel and G.

House (USNM), W. Hanson (EMUS), R. Anderson and F.

Genier (CMNC), J. Beard and E. Dubois (BMNH), N. Berti

(MNHN), A. Solis (INBC), P. Skelley (FSCA, PESC), R. Zack

(WSUC), F. Merickel (WFBM), M. Ivie (MAIC), J. Liebherr

and R. Hoebeke (CUIC), P. Perkins (MCZC), P. Lago (UMIC),

R. Davidson (CMNH), F. Andrews (CDFA), M. Uhlig

(ZMHB), A. Tishechkin, D. Verity (DVC) and P. Kovarik

(PKC). Special thanks are due P. Perkins for help with

LeConte types. For comments on early versions of this

manuscript I am grateful to D. Kavanaugh, J. Doyen, N.

Johnson, F. Sperling, P. Kovarik and R. Wenzel. This study

was funded in part through a California Agricultural Experi-

ment Station Grant to F. Sperling.

References

Arnett, R.H., Samuelson, G.A. & Nishida, G.M. (1993) The Insect and

Spider Collections of the World. Sandhill Crane Press, Gainesville.

Bickhardt, H. (1917) Histeridae. Genera Insectorum (ed. by P.

Wytsman), pp. 113±302, La Haye.

Bousquet, Y. (1991) Checklist of the Beetles of Canada and Alaska.

Agriculture Canada, Ottawa.

Bremer, K. (1994) Branch support and tree stability. Cladistics, 10,

295±304.

Casey, T.L. (1916) Memoirs on the Coleoptera, VII. Lancaster,

Pennsylvania.

Casey, T.L. (1924) Memoirs on the Coleoptera, IX. Lancaster,

Pennsylvania.

Caterino, M.S. (1998) A phylogenetic revision of Spilodiscus Lewis.

Journal of Natural History, 32, 1129±1168.

Donnelly, T.W. (1985) Mesozoic and Cenozoic plate evolution of the

Caribbean region. The Great American Biotic Interchange (ed. by F.

G. Stehli and S. D. Webb). Plenum Press, New York.

Erichson, W.F. (1834) Uebersicht der Histeroides der Sammlung.

Jahrbuch der Insektenkunde (ed. by J. C. F. Klug). Berlin.

Fall, H.C. (1910) Miscellaneous notes and descriptions of North

American Coleoptera. Transactions of the American Entomological

Society, 36, 89±197.

Horn, G. (1873) Synopsis of the Histeridae of the United States.

Proceedings of the American Philosophical Society, 13, 273±360.

LeConte, J.E. (1845) A monography of the North American

Histeroides. Boston Journal of Natural History, 5, 32±86.

LeConte, J.E. (1859) Description of new species of the coleopterous

family Histeridae. Proceedings of the Academy of Natural Sciences,

Philadelphia, 11, 310±317.

LeConte, J.L. (1869) Synonymical notes on Coleoptera of the United

States, with descriptions of new species, from the MSS. of the late

Dr. C. Zimmerman. Transactions of the American Entomological

Society, 2, 243±259.

Lewis, G. (1888) Histeridae. Biologia Centrali-Americana, Insecta,

Coleoptera (ed. by F. D. Godman and O. Salvin), pp. 182±244.

London.

Lewis, G. (1899) On new species of Histeridae and notices of others.

Annals and Magazine of Natural History, 4, 1±29.


Annals and Magazine of Natural History, 5, 224±254.


The Annals and Magazine of Natural History, 17, 337±344.

Liebherr, J.K. (1994) Biogeographic patterns of montane Mexican and

Central American Carabidae. The Canadian Entomologist, 126,

841±860.

Maddison, W.P. & Maddison, D.R. (1992) MACCLADE, Version 3.0.6.

Sinauer, Sunderland.

Marseul, S.A. (1854) Essai Monographique sur la Famille des

HisteÂrides (Suite). Annales de la Societe Entomologique de France,

2(ser. 3), 161±311, 525±592, 671±707.

Marseul, S.A. (1861) SuppleÂment aÂ la Monographie des HisteÂrides.

Annales de la Societe Entomologique de France, 1(ser. 4), 141±184,

509±566.

Mawdsley, J.R. (1993) The entomological collections of Thomas Say.

Psyche, 100, 163±171.

Mazur, S. (1984) A world catalogue of Histeridae. Polskie Pismo

Entomologiczne, 54, 1±379.

Noonan, G.R. (1988) Biogeography of North American and Mexican

insects, and a critique of vicariance biogeography. Systematic

Zoology, 37, 366±384.

Paykull, G. (1811) Monographia Histeroidum. Upsaliae.

Say, T. (1825) Descriptions of new species of Hister and Hololepta

inhabiting the United States. Journal of the Academy of Natural

Sciences, Philadelphia, 5, 32±47.

Schaeffer, C. (1912) Three new species of Hister. Bulletin of the

Brooklyn Entomological Society, 8, 26±27.

Schmidt, J. (1884) NachtraÈge und Berichtigungen zum Catalogus

Coleopterorum von M. Gemminger und E. v. Harold, betreffend die

Familie der Histeridae. Berliner Entomlogische Zeitschrift, 28, 147±

160.

Swofford, D.L. (1993) PAUP: Phylogenetic Analysis Using Parsimony,

Version 3.1.1. Illinois Natural History Survey, Champaign.

Wenzel, R.L. (1939) A new genus and several new species of North

American Hister. Annals of the Entomological Society of America,

32, 384±394.

Wenzel, R.L. (1944) On the classi®cation of histerid beetles.

Fieldiana, Zoology, 28, 51±151.

Wenzel, R.L. & Dybas, H. (1941) New and little known Neotropical

Histeridae (Coleoptera). Fieldiana, Zoology, 22, 433±472.

Accepted 18 November 1998

L



Documents

Taxonomy and phylogeny of the Hister servus group (Coleoptera: Histeridae): a Neotropical radiation