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A Revision of Camptocerus Dejean
(Coleoptera: Curculionidae: Scolytinae)
With Notes on its Biology

presented by

Sarah Marie Smith

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5I08 K:IProjIAoc&Pres/ClRC/Dateoue.indd

 

A REVISION OF CAIW’T OCER US DEJEAN (COLEOPTERA: CURCULIONIDAE:
SCOLYTINAE) WITH NOTES ON ITS BIOLOGY

By

Sarah Marie Smith

A THESIS

Submitted to
Michigan State University
in partial fulﬁllment of the requirements
for the degree of

MASTER OF SCIENCE
Entomology

2009

ABSTRACT

A REVISION OF CAWTOCER US DEJEAN
(COLEOPTERA: CURCULIONIDAE: SCOLYTINAE)

By
Sarah Marie Smith

The Neotropical ambrosia beetle genus Camptocerus Dejean was revised. The
monophyly of the genus was tested using 66 morphological characters in a cladistic
analysis. Camptocerus was monophyletic and 30 species are recognized. Six new
synonyms are presented: C. auricomus Blandford (= C. striatulus Hagedorn), C.
inoblitus (Schedl) (= C. morio (Schedl)), C. niger (F abricius) (= C. tectus Eggers), C.
opacicollis (Eggers) (=C. inﬁdelis Wood; = C. uniseriatus Schedl), C. suturalis
(F abricius) (=C. cinctus Chapuis). Two species received new status: C. chappentierae
Schedl and C. hirtipennis Schedl. Eleven new species of Camptocerus are described: C.
coccoformus, C. distinctus, C. dolei, C. igniculus, C. mallopterus, C. noel, C. petrovi, C.
pilifrons, C. pseudoangustior, C. unicornus and C. zucca. An illustrated key, diagnosis,

distribution, hosts and images were provided for each species.

I report observations of Camptocerus attracted to felled Protium amazonicum
Swart in southeastern Peru and the behavior of four species. An unusual behavior for
scolytines was observed as three species aggregated to leaves surrounding potential hosts,
bored into leaves and fed, a rare behavior for Scolytines. . Males competed for mates
with physical contact. Ant predation killed ~50% of colonizing individuals for one

species.

ACKNOWLEDGEMENTS

This work would not have been possible without the guidance and generous
assistance of my major professor, Dr. Anthony Cognato. I thank my thesis committee
members Dr. Deborah McCullough and Dr. Alan Prather for their guidance and advice.
This work was enhanced through the support and advice of my colleagues in the Holistic
Insect Systematic Laboratory: Bernice DeMarco, Dr. Stephanie Dole, Dr. Jiri Hulcr and
Aaron Smith. Krishna Dole and Dr. Matthew Yoder provided assistance with MX. Nick
Barc assisted with specimen curation. I would like to thank Gary Parsons, Tom Atkinson,
Dr. Don Bright, Dr. Alan Gillogly, Dr. Bjarte Jordal, Dr. Milos Knizek, Alex Petrov, Dr.
Robert Rabaglia and Dr. Stephen Wood for their assistance with this research. I would
like to thank my collaborators Dr. Gerardo Lamas (UNMSM), IRENA (Peru), Dr. Donald
Windsor (STRI) and ANAM (Panama) for assistance with collecting efforts. Rocio
Martinez and Dr. Rodrigo Mercader provided Spanish translations and valuable permit
assistance. I am grateful to Dr. Nigel Pitman (ACCA) and the Centro de Investigacion y
Capacitacion Rio Los Amigos for allowing me to conduct studies on the biology of
Camptocerus and for tree identiﬁcation. I would also like to thank the institutions

mentioned herein that loaned material critical to my studies.

I would like to thank my parents, Thomas and Debra Smith, and my grandmother,

Catherine Smith for always encouraging my curiosity and supporting my dreams.

This research was supported by NSF-PEET grant (DEB-0328920), USDA-FS

Early Detection and Rapid Response of Exotic Bark Beetles (07-DG-l 1420004-182)

iii

awarded to Anthony 1. Cognato and graduate research grants ﬁ'om the Michigan State
University Department of Entomology and the Asociacion para la Conservacion de la

Cuenca Amazonica (ACCA) awarded to S.M.S.

iv

TABLE OF CONTENTS

LIST OF TABLES ................................................................................. vi

LIST OF FIGURES ................................................................................ vii

CHAPTER ONE

A REVISION OF CAJW’TOCER US DEJEAN (COLEOPTERA: CURCULIONIDAE:

SCOLYTINAE) ................................. 1
Abstract ....................................................................................... 2
Introduction .................................................................................. 3
Materials and Methods .................................................................... 12
Results and Discussion ................................................................... 36

CHAPTER TWO

OBSERVATIONS OF CAA/ﬂ’TOCERUS DEJEAN (COLEOPTERA:

CURCULIONIDAE: SCOLYTINAE) BIOLOGY IN PERU .......... ' .................... 225
Abstract .................................................................................... 226
Introduction ............................................................................... 227
Materials and Methods .................................................................. 228
Results ..................................................................................... 229
Discussion ................................................................................. 235

LITERATURE CITED .......................................................................... 239

LIST OF TABLES

Table 1.1: Biogeographic distribution of Camptocerus species: 0 = present ........... 135

Table 1.2: Morphological character matrix of 66 characters for 41 species. Characters
and states are described in Materials and Methods ........................................... 137

vi

LIST OF FIGURES

Figure 1.1: Horizontal cross section (A); vertical cross section (B) of Camptocerus
aeneipennis galleries .............................................................................. 143

Figure 1.2: Distribution map illustrating number of Camptocerus species reported for
each country in the Neotropics. Countries without numbers lack records. Camptocerus
range from Veracruz, Mexico (Atkinson & Equihua- Martinez, 1986) to Chaco,
Argentina (Bruch, 1914) ......................................................................... 144

Figure 1.3. Camptocerus suturalis male head illustrating terminology used for the
epistoma and frons ................................................................................ 145

Figure 1.4: Character 7: Male proximal epistoma] margin (A) transverse, concave carina;
(B) tumid above scape insertion; (C) strongly developed transverse carina; (D) weakly
developed transverse carina; (E) no demarcation; (F) transverse carina with strongly
developed lateral areas; (G) short transverse carina .......................................... 146

Figure 1.5: Character 14: Male frons vestiture on impressed area (A) absent; (B) few
setae, less than 20 on impressed area, ﬁ'ons clearly visible; (C) moderate setae; frons
largely covered by setae, but surface visible; (D) dense, frons surface not visible .......147

Figure 1.6: Character 21: Mycangia on female gena: (A) absent; (B) crescent shaped; (C)
oval; (D) triangular ............................................................................... 148

Figure 1.7: Character 25: Male anterior pronotal margin (A) rounded; (B) partially
sulcate; (C) sulcate; (D) transversely impressed .............................................. 149

Figure 1.8: Character 26: Female anterior pronotum margin (A) rounded; (B) partially
sulcate; (C) transversely impressed ............................................................. 150

Figure 1.9: Character 30: Lateral margin of pronotum (A) type A; (B) type B; (C) type
C; (D) type D; (E) type E; (F) type F; (G) type G; (H) type H; (I) type I .................. 151

Figure 1.10: Character 31: Pronotum base (A) straitgh; (B) recurved; (C) bisinuate; (D)
notched .............................................................................................. 152

Figure 1.11: Character 35: Metepistemum shape (A) type A; (B) type B; (C) type C; (D)
type D; (E) type B; (P) type F; (G) type G; (H) type H; (1)1ype 1; (1) type J ; (K) type K;
(L) type L; (M) type M; (N) type N; (0) type 0; (P) type P; (Q) type Q .................. 153

Figure 1.12: Character 36: Mesosternum (A) type A; (B) type B; (C) type C; (D) type D;
(E) type B; (F) type F; (G) type G; (H) type H; (I) type I; (J) type J; (K) type K; (L) type
L; (M) type M; (N) type N; (0) type 0; (P) type P ........................................... 154

vii

Figure 1.13: Character 37: Basal margin of elytra (A) ﬂat; (B) faint carina; (C) faint
carina extending to scutellar apex (D) crenulations ............... 155

Figure 1.14: Character 38: Scutellum shape (A) type A; (B) type B; (C) type C; (D) type
D; (E) type B; (F) type F; (G) type G; (H) type H; (I) type I; (J) type J; (K) type K; (L)

typeL;(M)typeM;(N)typeN;(O)type O;(P)typeP .................................... 156
Figure 1.15: Character 52: Density of elytra] setae (A) glabrous; (B) light; (C) moderate;
(D) dense, elytra] surface not visible ........................................................... 157

FIGURE 1.16: Phylogeny of Camptocerus. Phylogram of one of 26 most parsimonious
trees (432 steps; consistency index, 0.488; retention index, 0.694) generated for 39 taxa
with 66 characters from a heuristic search of 1000 stepwise random additions with TBR
in PAUP“. Numbers above the node indicate bootstrap values (>50) based on 1000
pseudoreplicates. Bremer support values are generated from a heuristic search of 100
random addition sequence replicates ............................................................ 158

Figure 1.17: Pit mycangia on female C. aeneipennis vertex. Pits are 3-5 times the
diameter of other punctures and 3-10 times as deep ......................................... 160

Figure 1.18: Camptocerus suturalis stealing fungus from a C. aeneipennis gallery: (A)
close proximity of entrance holes, with the smaller C. suturalis on top; (B) entrance
tunnel and two maternal galleries, C. suturalis created the bottom gallery;(C) maternal
galleries separated by a thin septum, within the region of fungal staining, C. suturalis is
in the bottom gallery .............................................................................. 161

Figure 1.19: Lateral (A) and dorsal (B) views of Camptocerus aeneipennis, 4.4 — 5.9
mm, male ........................................................................................... 162

Figure 1.20: Camptocerus aeneipennis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ................................................ 163

Figure 1.21: Lateral (A) and dorsal (B) views of Camptocerus angustior, 5.0 — 5.8 mm,
male ................................................................................................. 164

Figure 1.22: Camptocerus angustior male head anterior (A), anterior oblique (B) . . ..165

Figure 1.23: Lateral (A) and dorsal (B) views of Camptocerus annectens, 3.5 mm,

female holotype ................................................................................... 166
Figure 1.24: Camptocerus annectens female holotype head anterior (A), anterior oblique
(B) ................................................................................................... 167
Figure 1.25: Lateral (A) and dorsal (B) views of Camptocerus aterrimus, 3.1 — 4.5 mm,

male ................................................................................................. 168

viii

Figure 1.26: Camptocerus aterrimus male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ......................................................... 169

Figure 1.27: Lateral (A) and dorsal (B) views of Camptocerus auricomus, 3.5 — 4.1 mm,
male ................................................................................................. 170

Figure 1.28: Camptocerus auricomus male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ................................................ 171

Figure 1.29: Lateral (A) and dorsal (B) views of Camptocerus chamentierae, 3.8 — 4.5
mm, holotype male ............................................................................... 172

Figure 1.30: Camptocerus chamentierae holotype male head anterior (A), anterior
oblique (B); female head anterior (C), anterior oblique (D) ................................. 173

Figure 1.31: Lateral (A) and dorsal (B) views of Camptocerus coccoformus, 2.7 mm,
holotype male ...................................................................................... 174

Figure 1.32: Camptocerus coccoformus holotype male head anterior (A), anterior
oblique (B); allotype female head anterior (C), anterior oblique (D) ...................... 175

Figure 1.33: Lateral (A) and dorsal (B) views of Camptocerus costatus, 3.5 — 3.9 mm,
male ................................................................................................. 176

Figure 1.34: Male declivity (A) and female lateral (B) views of Camptocerus costatus
........................................................................................................ l 77

Figure 1.35: Camptocerus costatus male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ......................................................... 178

Figure 1.36: Lateral (A) and dorsal (B) views of Camptocerus distinctus, 5.3 mm,
holotype male ...................................................................................... 179

Figure 1.37: Camptocerus distinctus holotype male head anterior (A), anterior oblique
(B); allotype female head anterior (C), anterior oblique (D) ................................ 180

Figure 1.38: Lateral (A) and dorsal (B) views of Camptocerus doIei, 3.1 — 3.5 mm,
holotype male ...................................................................................... 181

Figure 1.39: Camptocerus dolei holotype male head anterior (A), anterior oblique (B);
allotype female head anterior (C), anterior oblique (D) ..................................... 182

Figure 1.40: Lateral (A) and dorsal (B) views of Camptocerus hirtipennis, 4.9 —- 5.6 mm,
male ................................................................................................. 183

Figure 1.41: Camptocerus hirtipennis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ................................................ 184

ix

Figure 1.42: Lateral (A) and dorsal (B) views of Camptocerus igniculus, 2.5 — 2.8 mm,
male ................................................................................................. 185

Figure 1.43: Camptocerus igniculus male head anterior (A), anterior oblique (B);
allotype female head anterior (C), anterior oblique (D) ..................................... 186

Figure 1.44: Lateral (A) and dorsal (B) views of Camptocerus inoblitus, 2.4 - 3.4 mm,
male ................................................................................................. 187

Figure 1.45: Camptocerus inoblitus male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ......................................................... 188

Figure 1.46: Lateral (A) and dorsal (B) views of Camptocerus latipilis, 2.7 — 3.3 mm,
male ................................................................................................. 189

Figure 1.47: Lateral (A) and dorsal (B) views of Camptocerus Iatipilis, 2.7 — 3.3 mm,
female ............................................................................................... 190

Figure 1.48: Camptocerus latipilis male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ........................................................ 191

Figure 1.49: Lateral (A) and dorsal (B) views of Camptocerus major, 3.4 mm, holotype
female .............................................................................................. 192

Figure 1.50: Camptocerus major holotype female head anterior (A), anterior oblique (B)
........................................................................................................ 193

Figure 1.51: Lateral (A) and dorsal (B) views of Camptocerus mallopterus, 3.8 — 3.9
mm, holotype male ............................................................................... 194

Figure 1.52: Camptocerus mallopterus holotype male head anterior (A), anterior oblique
(B); allotype female head anterior (C), anterior oblique (D) ................................ 195

Figure 1.53: Lateral (A) and dorsal (B) views of Camptocerus mandelshtami, 5.9 - 6.5
mm, male .......................................................................................... 196

Figure 1.54: Camptocerus mandelshtami male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ................................................ 197

Figure 1.55: Lateral (A) and dorsal (B) views of Camptocerus niger, 2.8 — 4.1 mm, male
....................................................................................................... 198

Figure 1.56: Camptocerus niger male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ......................................................... 199

Figure 1.57: Lateral (A) and dorsal (B) views of Camptocerus noel, 6.0 - 7.1 mm, male.
........................................................................................................ 200

Figure 1.58: Camptocerus noel male head anterior (A), anterior oblique (B); female head
anterior (C), anterior oblique (D) ............................................................... 201

Figure 1.59: Lateral (A) and dorsal (B) views of Camptocerus occidentalis, 4.0 — 4.5
mm, male ........................................................................................... 202

Figure 1.60: Camptocerus occidentalis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ................................................ 203

Figure 1.61: Lateral (A) and dorsal (B) views of Camptocerus opacicollis, 2.2 — 3.6 mm,
male ................................................................................................ 204

Figure 1.62: Camptocerus opacicollis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ................................................ 205

Figure 1.63: Lateral (A) and dorsal (B) views of Camptocerus orientalis, 3.2 — 3.7 mm,
holotype male
....................................................................................................... 206

Figure 1.64: Camptocerus orientalis holotype male head anterior (A), anterior oblique
(B); paratype female head anterior (C), anterior oblique (D) .............................. 207

Figure 1.65: Lateral (A) and dorsal (B) views of Camptocerus petrovi, 4.0 mm, holotype
male ................................................................................................ 208

Figure 1.66: Camptocerus petrovi holotype male head anterior (A), anterior oblique (B);
abdomen posterior oblique (C) ........................................................... _ ...... 209

Figure 1.67: Lateral (A) and dorsal (B) views of Camptocerus pilzﬁons, 2.6 — 3.1 mm,
holotype male .................................................................................... 210

Figure 1.68: Camptocerus piliﬁ-ons holotype male head anterior (A), anterior oblique
(B); allotype female head anterior (C), anterior oblique (D) .............................. 211

Figure 1.69: Lateral (A) and dorsal (B) views of Camptocerus pseudoangustz'or, 4.5 -
5.4 mm, male ..................................................................................... 212

Figure 1.70: Camptocerus pseudoangustior male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D) ............................................... 213

Figure 1.71: Lateral (A) and dorsal (B) views of Camptocerus quadridens, 3.4 — 3.7 mm,
male ................................................................................................. 214

Figure 1.72: Posterior oblique (A) view of Camptocerus quadridens male. Lateral (B)
view of female ..................................................................................... 215

xi

Figure 1.73: Camptocerus quadridens male head anterior (A), anterior oblique (B);

female head anterior (C), anterior oblique (D) ................................................ 216
Figure 1.74: Lateral (A) and dorsal (B) views of Camptocerus rectus 3.5 — 4.3 mm, male
....................................................................................................... 217
Figure 1.75: Camptocerus rectus male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ......................................................... 218
Figure 1.76: Lateral (A) and dorsal (B) views of Camptocerus suturalis, 3.6 — 5.0 mm,
male ................................................................................................. 219
Figure 1.77: Camptocerus suturalis male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D) ......................................................... 220
Figure 1.78: Lateral (A) and dorsal (B) views of Camptocerus unicornus, 2.5 mm,
holotype male ...................................................................................... 221
Figure 1.79: Camptocerus unicomus holotype male head anterior (A) .................. 222

Figure 1.80: Lateral (A) and dorsal (B) views of Camptocerus zucca, 4.0 - 4.1 mm,
holotype male ...................................................................................... 223

Figure 1.81: Camptocerus zucca holotype male head anterior (A), anterior oblique (B);
allotype female head anterior (C), anterior oblique (D) ..................................... 224

Figure 2.1: (A) Males of Camptocerus noel aggregating and resting in a leaf near a felled
Protium amazom'cum tree. They made the holes in which they rest. (B) Female
Camptocerus noel resting in a leaf and (C) boring through the leaf with undigested leaf
cuttings .............................................................................................. 237

Figure 2.2: Damage caused by feeding male of Comptocerus noel to a P. amazonicum
leaﬂet ................................................................................................ 238

xii

CHAPTER ONE

A Revision of Camptocerus Dejean (Coleoptera: Curculionidae: Scolytinae)

ABSTRACT

The Neotropical ambrosia beetle genus Camptocerus Dejean was revised. The
monophyly of the genus was tested using 66 morphological characters in a cladistic
analysis. Camptocerus was monophyletic and 30 species are recognized. Six new
synonyms are presented: C. auricomus Blandford (= C. striatulus Hagedom), C.
inoblitus (Schedl) (= C. morio (Schedl)), C. niger (F abricius) (= C. tectus Eggers), C.
opacicollis (Eggers) (=C. inﬁdelis Wood; = C. uniseriatus Schedl), C. suturalis
(F abricius) (=C. cinctus Chapuis). Two species received new status: C. chamentierae
Schedl and C. hirtipennis Schedl. Eleven new species of Camptocerus are described: C.
coccoformus, C. distinctus, C. dolei, C. igniculus, C. mallopterus, C. noel, C. petrovi, C.
pilifrons, C. pseudoangustior, C. unicornus and C. zucca. An illustrated key, diagnosis,

distribution, hosts and images were provided for each species.

INTRODUCTION

Beetles of the subfamily Scolytinae (Coleoptera: Curculionidae) are ubiquitous in
forests world-wide and contribute to the decomposition of dead vegetation. More than
6,000 species (Wood & Bright, 1992) specialize on plant parts, from root to fruit, as food
resources for larvae and adults (Wood, 1982). Scolytines, along with the closely related
subfamily Platypodinae, are collectively referred to as the bark and ambrosia beetles. The
Scolytinae are primarily composed of two distinct ecological feeding groups; bark and
ambrosia beetles. True bark beetles feed on phloem and cambium of live, declining or
dead trees. Ambrosia beetles tunnel into the xylem of a host tree and cultivate a symbiotic
fungal garden on which the adults and larvae feed. This ambrosia] feeding habit has
evolved multiple times within the Scolytinae (Farrell et al., 2001). The primary behavior
of boring under bark and into sapwood hastens decomposition via the introduction of
other xylophagous organisms.

Even though South America possesses one of the most diverse groups of
scolytines in the world (Wood, 2007), the fauna is one of the least known (Bright, 1993;
S. L. Wood, pers. comm). Along with under-studied species diversity, little is known
about South American scolytine biology and ecology, including tree host associations
(Beaver, 1972). One such poorly known group is the tribe Scolytini and speciﬁcally the
ambrosia beetle genus Camptocerus Dej ean (Wood, 2007). This study was undertaken to
taxonomically revise and redescribe Comptocerus (Scolytinae: Scolytini). Phylogenetic

analysis based on adult morphological and ecological characters was incorporated to

determine monophyly of the genus, species validity, species relationships and the
relationship of Camptocerus within the tribe Scolytini.
Natural History

Camptocerus is unique among the genera in the tribe Scolytini because the larvae
are xylomycetophagous (xylem and ambrosial fungus feeding) instead of phloeophagous
(phloem feeding) (Beaver, 1972; Wood, 1972, 1982, 1986, 2007; Petrov, 2007). The
female creates an entrance trmnel into the cambium where the male joins her. Infestation
of moribund host material can start within one day, but typically occurs within 2-4 days
and ceases after 3 to 10 days (Beaver, 1972). Personal observations in Peru indicate that
Camptocerus can continue colonization for more than 26 days (Chapter 2). In a typical
gallery, the entrance tunnel runs in a transverse plane toward the center of the stem. From
this tunnel, the female adds one or two circumferential branches creating a maternal
gallery (Wood, 2007). Females lay single eggs along the maternal gallery in wide niches
cut at right angles to the gallery and on parallel sides. Niches are plugged with boring
dust (Beaver, 1972). Females occupy the inner galleries and males protect the gallery
enhance by plugging the holes with their abdomens (Beaver, 1972; chapter 2).

Larvae hatch and feed on xylem and the black mycelial growth of the ambrosia
fungi lining the niche. Each larva enlarges its niche to a cradle as it grows and is capable
of turning around within the cradle. The 7-10 week life cycle is longer than other
xylomycetophagous scolytines and platypodines and may perhaps be due to the slow
growth of the ambrosial fungus (Beaver, 1972). Females probably inoculate the tree with
ﬁmgal spores stored in their mycangia upon attack (Beaver, 1972), but, the location of the

mycangia have been previously unknown.

Several Camptocerus species have been known to colonize a single host tree and
the species segregate themselves by trunk/branch size, with large species attacking the
trunk and large branches (5- <30 cm) and small species attacking smaller branches (1— 5
cm) (Beaver, 1972; chapter 2). Depending on the site, a single host has been observed to
contain between one and four (Beaver, 1972) and six species of Campiocerus (Smith, per.
0b.).

Camptocerus species also perform leaf feeding, a rare and unusual behavior for
Scolytines. Camptocerus aeneipennis adults have been found feeding on leaves of a
Thrysodium species (Anacardiaceae) in the Panama Canal Zone (von Winning, 1930).
The beetles had chewed the leaf 8 central vein to form circular holes slightly smaller than
the width of their bodies. Each beetle was lying in a hole with its mesostemum lying
between the leaf surfaces, with beads directed toward the leaf’s ventral surface. Once
positioned, the beetles performed window feeding (feeding only on one surface of the
leaf) a few mm from their present hole or up to 20 m away. It is presumed that the
beetles were using the leaves for maturation feeding (von Winning, 1930). This behavior
has also been observed by C. opacicollz's (Eggers), C. noel Smith & Cognato sp. n., C.
pseudoangustior Smith & Cognato sp. 11. (chapter 2), C. aeneipennis and C. aterrimus in
southeastern Peru (S.M.Smith, pers. ob.) and by C. suturalis in Loreto province, Peru (A.
Petrov, pers. com). Although these behaviors were not witnessed, feeding damage from
C. aeneipennis and C. auricomus Blandford were also observed by me along Pipeline Rd
in the Parque Nacional Soberania, Panama Province, Panama. This locality is along the
Panama Canal and in the same area from which this behavior was ﬁrst reported (von

Winning, 1930).

Camptocerus exhibit a unique aggregation technique. It is apparent that
Camptocerus also chew aggregation holes within 2 m of host tree. The behavior is similar
to maturation feeding except that a hole is chewed in the leaf, but the material is not
digested. In addition, there is never window-feeding damage near the hole. This behavior
creates distinctive damage that is easily recognizable by perfectly formed circular holes
located between two leaf veins (S.M.Smith, pers. ob).

Host records for Camptocerus are limited, but Camptocerus have been found
colonizing eleven different tree families with host records reported for half of all species.
Camptocerus are overwhelmingly collected from members of the Burseraceae
(speciﬁcally Protium sp.), but they can also utilize species of Chrysobalanaceae,
Clusiaceae, Euphorbiaceae, Fabaceae Lauraceae, Leguminosae, Meliaceae, Myrtaceae,
Roseaceae, Sapindaceae to a lesser extent. Camptocerus have been observed to
preferentially colonize Protium over other documented hosts (Beaver, 197 2) and Beaver
(1989) concluded that Camptocerus species are host speciﬁc on Protium . Host
speciﬁcity is unusual for ambrosia beetles, which are typically polyphagous (Hulcr et al.,
2007a)

The geographical range of Camptocerus extends from Veracruz, Mexico
(Atkinson & Equihua- Martinez, 1986) to Chaco, Argentina (Bruch, 1914). To date, only
four species have been found in Mexico and Central America. Of those, one is exclusive
to Central America, two are distributed in Central and South America, one is distributed
in Mexico, Central, and South America; 26 species are exclusively in South America
(Table 1.1). Given this distribution pattern, Wood (1982) hypothesized that the genus

originated in South America and has radiated north over time. In addition, the greatest

diversity of Camptocerus species (83%), as well as the other Scolytini, lies in the

Amazon Basin (Wood & Bright, 1992; Wood, 2007) (Fig.1.2).

Systematics

The tribe Scolytini is currently composed of approximately 209 species in four
genera: Camptocerus Dejean 1821, Cnemonyx Eichhoff 1868, Scolytus Geoffroy 1762,
and Scolytopsis Blandford 1896. The tribe likely originated in the Neotropics with all
genera distributed in the New World, except for numerous Palearctic Scolytus (Wood,
1986). All genera are phlonphagous (phloem feeding) with the exception of the
xylomycetophagous Camptocerus. All genera are monogamous except for a few species
of Neotropical Scolytus (Wood, 1982). Members of the tribe are identiﬁed by a single
apical, unarmed spine-like process that curves toward and extends beyond the process of
the inner apical angle of the protibia and a seven-segmented funicle. All display a
strongly sexually dimorphic head ornamented with hair-like setae, with the male frons
variously excavated and female frons ﬂat to convex (excavated in one Scolytus species).
The eye is entire and the posterior area of head is subtrrmcate. The pronotum is unarmed
with a costate lateral margin. In addition, the metapleural suture descends subventrally to
the groove, receiving the groove on the costal margin of the elytra, then turns abruptly
and parallels the groove near the metacoxal process (Wood, 1978, 1982, 1986).

Camptocerus has historically been diagnosed from the other genera of the tribe by
a large, broad, and ﬂat scutellum that is at least 1.5 times wider than long, the absence of

a notch at the elytra] base for scutellar reception, generally larger body size, gradually

ll

ascending abdomen, extensively excavated male frons, suture 1 of antenna] club marked
internally by a septum, a usually ﬁnely sculptured elytra, by meso- and metathoracic
tibiae acutely margined on apical anterior edge without supplemental denticles, and by
xylomeycetophagy (Wood, 1982, 1986, 2007). Camptocerus species show marked sexual
dimorphism of the ﬁ'ons and epistoma; the male frons is often strongly excavated and the
female frons is ﬂat to convex. The male frons is covered by more and longer setae than
the female frons. The epistoma is elongated and rostrate-like in the male. Sexual
dimorphism also occurs in the antenna] scape and funicle and to various degrees in the
elytral declivity. The female scape and frmicle are similar to the male antenna but lack the
long brush-like setae on funicle segments 2-7. It is difﬁcult to distinguish between some
members of Camptocerus and Cnemonyx due to similar morphology. They are most
effectively identiﬁed by phloeophagous (Cnemonyx) or xylomycetophagous
(Camptocerus) feeding habits (S.L. Wood, pers. comm). Unfortunately, many
Camptocerus species have only been collected by passive methods (e. g. canopy fogging).
As a result, species identity of approximately half of the Camptocerus species has been
conﬁrmed by ﬁeld observations of xylomycetophagy; feeding habits of the remaining

species are unknown.

Taxonomic History
Camptocerus was established as a monotypic genus by Dejean who designated
Hylesinus aeneipennis Fabricius as the type species. However authorship of Camptocerus

has been attributed to Latreille in the majority of taxonomic literature (Erichson, 1836;

LaPort de Castelneau, 1840; Lacordaire, 1866; Chapuis, 1869; Girard, 1873; Blandford,
1896; Hagedom, 1910; Blackman, 1943; Blackwelder, 1947; Schedl, 1952; Schedl, 1962;
Beaver, 1972; Wood, 1978, 1982, 1986; Beaver, 1989; Wood & Bright, 1992; Petrov,
2007; Wood, 2007) with three exceptions that also cite Dejean (Lacordaire, 1866;
Beaver, 1972; Wood, 1972) and three that recognize Dejean (Chevrolat, 1838; Hopkins,
1914, 1915). Dejean is the rightful author; he named the genus and assigned a type
species of Hylesinus aeneipennis Fabricius, but he did not provide a written description
of the genus (Dejean, 1821). However, his action validated the name in accordance with
the ICZN code article 12.2.5 (ICZN, 1999). In 1829, Latreille cited Dejean when he
described the genus Camptocerus as “whose males have strongly bent antennae,
furnished outside with long hairs; which are inserted at a notable distance from the eyes,
which are elliptical and oblique.” It seems likely that other authors were unaware of the
Dejean (1821) publication and credit was instead given to Latreille (Lacordaire, 1866).
The unique morphology exhibited by Camptocerus species has led to the creation
and subsequent placement in the group (tribe) Camptocérides with the Paleotropical
genus Diamerus Erichson (Lacordaire, 1866). Ferrari (1867) recognized the
Camptocérides as a subtribe. Chapuis (1869) elevated the Camptocerides back to a tribe
and Blandford (1893) maintained this taxonomic position, until he later sunk the
Camptocérides back to the level of subgroup (subtribe) (Blandford 1896). Blandford
(1896) removed Diamerus from this group and placed the genera Cnemonyx, Loganius
Chapuis, Ceratolepsis Chapuis into the subtribe. Hopkins (1915) elevated the subtribe to
the subfamily Camptocerinae. Blatchey & Leng (1916) sunk the Camptocerinae back to

a subtribe, which they named Camptocerini. Costa Lima (1956) departed from this

classiﬁcation and followed Hopkins (1915) recognizing these genera at the subfamily
level. Wood (1961) appeared to have placed the Camptocerini as a synonym of the tribe
Scolytini.

Prior to Wood’s 1972 anatomical revision, the tribe Scolytini consisted of two
clearly deﬁned genera (Scolytus and Scolympsis) and ﬁve strikingly similar genera
(Camptocerus, Cnemonyx, Loganius, Ceratolepsis, and Coptosomus Schedl).
Camptocerus, Ceratolepsis, and Coptosomus possess meso- and metathoracic tibiae that
are identical to the prothoracic tibiae, with a straight, sharp, and smooth anterior edge of
the apical margin to the base of the outer apical spine. In Cnemonyx and Loganius, the
surface of the tibial anterior edge of the apical margin to the base of the outer apical spine
bears one or more denticles. Species with these denticles also have two to three sutures
on the antennal club marked with setae, while species without denticles lack externally
marked sutures but at least part of suture 1 is marked internally by a septum. Schedl
(1962) noted that the characters used to diagnose these genera were inconsistent and
formed a spectrum of variation. This caused him to synonymize Ceratolepsis with
Camptocerus. However, Wood (1972) determined the genus Camptocerus was a distinct
group without any synonymies; he found the characters used to identify Ceratolepsis
(scales on antennal club), Cnemonyx (crenulations of elytral base), and Coptosomus
(oddly placed antennal suture) were insufﬁcient to clearly deﬁne species groups and
transferred them to the oldest described genus, Cnemonyx. Loganius species were divided
between Cnemonyx and Camptocerus.

Five Camptocerus species were among the ﬁrst New World scolytines to be

described (Illiger, 1807; Blandford, 1896). Since 1801 , nine authors have contributed

10

species. In 1801, Fabricius described Hylesinus aeneipennis, H. fasciatus, H. gibbus, H.
niger, and H. suturalis. In 1821, Dejean created the genus Camptocerus using Hylesinus
aeneipennis as the type species and the remaining species were subsequently placed in
Camptocerus by Wilhelm Erichson (1836). Several taxonomists have described
Camptocerus species including F elicien Chapuis (1869), Walter Fielding Holloway
Blandford (1896), Max Hagedom (1910), Hans Eggers (1928, 1929, 1932, 1943),
Maulsby W. Blackrnan (1943), Karl Schedl (1939, 1952, 1970, 1972, 1973), Stephen L.
Wood (1969, 1972, 2007), and Alex Petrov (2007). To date, only C. bolivae has been
moved to another genus (Cnemonyx). However, several Loganius species have been
transferred to Camptocerus, including C. inoblitus (Wood, 1972), C. major (Schedl,
1952), C. morio (Wood and Bright, 1992), and C. opacicollis(Sched1, 1952; Wood,
1972). Camptocerus terebrator Lacordaire (1833) is a nomen nudum. This species does
not meet the criteria outlined by the ICZN article 12.1 as there is no description,
deﬁnition or indication of the species. Schedl (1976) also indicated that the species did
not meet these requirements, but failed to recognize the name as a nomem nudum.

It is quite puzzling that Camptocerus has one of the longest taxonomic histories of
American scolytines yet remains one of the least understood genera. In Biologia-Centrali
Americana, Blandford (1896) noted that with the exception of C. aeneipennis,
Camptocerus species were rare in collections and few were described. This situation
persists to the present with several species represented by fewer than ﬁve individuals; C.
aeneipennis remains common in collections. Indeed biological information exists for

fewer than half of the known species with most data pertaining to C. aeneipennis.

11

Although the genus Camptocerus was recently reviewed by Wood (2007), an
additional 11 new species have since been discovered. Examination of Dr. Terry Erwin’s
Ecuadorian Amazon rain forest canopy fogging samples collected between 1994 and
2006 yielded seven novel Camptocerus species. Museum loans and recent collecting by
the authors in Guyana and Peru have also led to the discovery of an additional four new
species and previously unknown females for three species, C. charpentierae Schedl, C.
costatus Chapuis and C. quadridens Blackman. Currently, the genus is composed of 30

species, of which 19 are valid and an additional 11 species are described herein.

Materials and Methods

Approximately 1182 Camptocerus specimens representing all taxa from a range
of locations in Mexico, Central and South America were examined for this study. Five
species of Cnemonyx and one species of Scolytopsis were also included in the ingroup.
Five Scolytus species (two Neotropical, one Nearctic, and two Palearctic) were selected
as the outgroup and as the root taxa.

The following entomological collection codens (following Evenhuis, 2007) and

abbreviations are referenced in the text:

AMNH American Museum of Natural History, New York, New York, USA.
BMNH British Natural History Museum, London, United Kingdom.

CASC California Academy of Sciences, San Francisco, California, USA.
CSCA California State Collection of Arthropods, Sacramento, California, USA.
DEI Deutsches Entomologisches Institute, Leibniz-Zentrum ﬁir

Agrarlandschaﬁsforschung, Mﬁncheberg, Germany.

12

ll

FMNH

IRSNB

MECN

MSUC

MUSM

MZUSP

RJRC

SEMC

SMTD

STRI

UCDC

UZMC

ZMMU

Field Museum of Natural History, Chicago, Illinois, USA.

Institute Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium.
Museo Ecuadoriano de Ciencias Naturales, Quito, Ecuador.

Muséum National d'Histoire Naturelle, Paris, France.

Albert J. Cook Arthropod Research Collection, Michigan State University,
East Lansing, Michigan, USA.

Universidad Nacional Mayor de San Marcos, Lima, Peru.

Museum de Zoologia, Universidad de Sao Paulo, Sao Paulo, Brazil.
Naturhistorisches Muesum Wien, Vienna, Austria.

National Museum of Natural History, Smithsonian Institution,
Washington, DC, USA (Including Stephen L. Wood Collection).
Robert J. Rabaglia collection, Maryland, USA.

Snow Entomological Museum, Lawrence, Kansas USA.

Staatliches Museum ﬁtr Tierkunde in Dresden, Germany.
Smithsonian Tropical Research Institute Collection, Balboa, Panama.
Bohart Museum of Entomology, University of California, Davis,
California, USA.

Museum of Zoology, University of Michigan, Ann Arbor, Michigan,
USA.

Natural History Museum of Denmark, University of Copenhagen,
Zoological Museum, Copenhagen, Denmark.

Zoological Museum, Moscow State University, Moscow, Russia.

13

Host plants were compiled from the following publications: Kleine, 1934;
Bondar, 1950; Viana, 1965; Beaver, 1972; Atkinson & Equihua-Martinez, 1986; Maes &
Equihua-Martinez, 1988; Wood & Bright, 1992; Wood, 2007; L. Kirkendall (per.
comm); A. Petrov (per. comm). Distribution records were gleaned from the following

sources: Chevrolat, 1838; Hagedom, 1903, 1910; Kleine 1913, 1914; Bruch, 1914;

Viana, 1914; Nunberg, 1962, Schedl, 1972; Beaver, 1972; Atkinson & Equihua-Martinez,

1986; Maes & Equihua-Martinez, 1988; Wood & Bright, 1992; Bright & Skidmore,
1997, 2002; Wood, 2007; A. Petrov (per. com); T. Atkinson (per. com). Plant
nomenclature was checked using Discover Life (discoverlifecrg) and the International
Plant Names Index (www.ipni.org).

Specimens were examined using a Leica M216 compound microscope,
illuminated with a SCHOTT 150W halogen light source (model ACE ®1). Images were
taken with a Leica DFC 320 and montage images were assembled using Helicon Focus
3.50,5 (HeliconSoﬁ) Following the protocol used by Wood (1982; 2007), measurements
were made using an ocular micrometer on the same microscope and with the same light
source as above calibrated with ROK 150 mm digital calipers (model DC-122A).
Measurements were made perpendicular to the specimen’s dorsal surface. Length was
measured from the anterior margin of the pronotum to the apex of the elytra, excluding
the head. Width was measured at the widest part of the pronotum, the base. Proportions
are the ratio of the length to the width. A maximum of 20 individuals were selected for
each species and encompassed the known species distribution.

Holotypes were designated with red labels. Lectotypes, allotypes and paratypes

were designated with yellow labels.

14

l

Morphological Characters

Traditional bark beetle generic and species level morphological characters were
employed in this study (Hopkins, 1915; Wood, 1978; Kniiek & Beaver, 2004).
Characters were taken from generic descriptions (Hagedom, 1910; Wood, 1982, 1986,
2007), species descriptions, and personal examination of specimens. C. aeneipennis was
used to determine intraspeciﬁc variation within a character due to its abundance in
museum collections and wide geographic range. All characters were based on adult
morphology and feeding habit. External anatomical terminology followed Hopkins
(1909), subsequently used by Schedl (1931) and by Wood (1982, 1986, 2007).
Sculpturing terminology followed Torre-Bueno (1989). The elongated epistoma of male
Camptocerus differed signiﬁcantly from other bark beetles and I created new
terminology to accommodate this structure. The male epistoma was deﬁned as the area
extending from the mandibular margin (here termed the apical epistomal margin) to the
level of antennal insertion (Wood, 1982, 2007) (here termed the basal epistomal margin).
The ﬁons was considered the region of the head originating at the basal epistomal margin
and terminating at the vertex of the head(Fig 1. 3).

Genitalia dissections were performed on male specimens of 35 of 40 taxa.
Dissections were not performed for species not represented by males, or for species only
known ﬁ'om a short series or holotype. Dissections were performed after soaking a
specimen in 70% ethanol for ten minutes. In small species, a hooked #2 stainless steel
insect pin was placed in the coxal cavity and was used to pry back the abdominal stemites

up to tergite 6. The genitalia were then removed and placed in 20% KOH until cleared,

15

usually after 5-7 days and neutralized in 20% glacial acetic acid. For larger species,
preparation was the same and the genitalia were removed by inserting a hooked pin into
the genital opening between the eighth sternite and tergite. The genitalia were removed
with the hook and cleared using the same methods as above. Genitalia were stored in
glass genitalia vials ﬁlled with glycerin or by placing them on mounting cards in the
manner recommended by the loaning institution.

Characters were scored ﬁ'om both sexes unless otherwise noted. Camptocerus
exhibited strong sexual dimorphism; males exhibit a wide spectrum of morphological
features while females display conserved morphology. As a necessary consequence,
characters were predominately male based. Characters were scored for all taxa except for
C. annectens Wood, and C. major (Eggers), which are only known from females. As a
result these species were excluded ﬁ'om the phylogenetic analysis (60% of characters
scored). Characters and character state numbers corresponded to data coded in the data
matrix for each taxon. The characters matrix (Table 1.2) was constructed and edited from
the online database MX (Y oder et al., 2006). Character transformations were evaluated
using MacClade 4.0 PPC (Maddison & Maddison, 2000) and homology of characters and
deﬁnitions of character states were re-examined and modiﬁed if necessary.

A total of 66 characters (24 binary and 42 multistate) were used in this study.
Twenty-two characters were coded from the head including the antennae, 11 characters
from the prothorax, 26 characters from the abdomen including the elytra and scutellum, 1
character from the mesostemum, 5 characters from male genitalia and 1 character based
on feeding habit. Consistency and retention index values generated from MacClade 4.0

are listed next to each character.

16

l

Head

1. Antennal club sutures (ci = 1.00; ti = 1.00)
0: zero
1: one
2: two

3: three

2. Female apical epistoma emarginate at medial line (ci = 1.00; ri = 1.00)
0: normal

1: emarginate

3. Male apical epistoma emarginate at medial line (ci = 1.00; ti = 1.00)
0: normal

1: emarginate

4. Relative epistoma size compared to ﬁons (ci = 0.14; ri = 0.65)
0: less than one fourth

1: greater than or equal to one-fourth

5. Vestiture on lateral margin of epistoma (ci = 0.60; ti = 0.33)

0: setae entire

1: setae on lower half

17

2: middle

3: glabrous

6. Male apical epistomal margin (ci = 0.57; ri = 0.73)
0: unarmed
1: short transverse carina
2: tubercule
3: large tubercle extending length of epistoma

4: median lobe

7. Male basal epistomal margin (ci = 0.60; ti = 0.82) (Fig. 1.4)
0: transverse, concave carina
1: tumid above scape insertion
2: strongly developed transverse carina
3: weakly developed transverse carina
4: no demarcation
5: transverse carina with strongly developed lateral areas

6: short transverse carina

8. Excavation of male from (ci = 0.40; ti = 0.57)
0: weak/feeble

1: strong

2: ﬂat

18

9. Male gena (ci = 0.17; ri = 0.00)

0:

1:

triangular structure absent

triangular structure present

10. Lateral margin of male epistoma (ci = 0.50; ri = 0.83)

0:

1:

arcuate costa pointing distally

arcuate costa pointing proximally

: vertical costa
: costa angled inward

: costa absent

11. Male ﬁons excavation size (ci = 0.40; ri = 0.67)

0:

1

2:

excavation does not border ocular margin

: excavation touches ocular margin

excavation absent

12. Male ﬁ'ons armiture (ci = 1.00; ri = 1.00)

0:

1:

unarmed

tubercle

13. Male frontal setae in two bands (ci = 0.25; ri = 0.57)

0:

absent

l9

l4.

0:

1:

15.

: present

Male ﬁons vestiture on excavated area (ci = 0.30; ri = 0.42) (Fig 1.5)
glabrous

few setae (less than 20); ﬁ'ons clearly visible

: moderate setae (frons largely covered by setae); surface obscured but visible

: dense setae; frons surface not visible

Male scape (ci = 0.80; ri = 0.96)

: elongate and narrowly rounded distally
: elongate and narrow

: elongate and expanded distally

: short and round

: elongate and broadly rounded distally

16. Antennal club setae (ci = 1.00; ti = 1.00)

0:

1:

palmately divided

one ﬁlament

17. Female epistoma (ci = 0.50; ti = 0.50)

0:

1:

unarmed

median lobe

20

|l

18. Female ﬁ'ons (ci = 0.43; ti = 0.56)
0: ﬂat
1: weakly impressed
2: convex

3: excavated

19. Armed female frons (ci = 0.33; ri = 0.0)
0: absent

1: carina

20. Medial area of female ﬁons (ci = 0.33; ri = 0.56)
0: ﬂat
1: carina
2: impressed
3: carina and impression

4: groove

21. Female gena mycangium (ci = 0.60; ri = 0.91) (Fig 1.6)
0: absent
1: crescent shaped
2: oval shaped

3: triangular shaped

21

22. Pars stridens (ci = 0.14; ri = 0.50)
0: extends to dorsal margin of eye

1: extends to level of scape insertion

Pronotum

23. Setae an anterior margin of female pronotum between eyes (ci = 0.25; ri = 0.57)
0: absent
1: scales

2: palmately divided into 3 or 4 ﬁlaments

24. Setae an anterior margin of male pronotum between eyes (ci = 0.25; ti = 0.70)
0: absent
1: scales
2: single seta

3: bifid/triﬁd

25. Male anterior pronotal margin (ci = 0.75; ri = 0.90) (Fig. 1.7)
0: rounded
1: partially sulcate
2: entirely sulcate

3: transversely impressed

26. Female anterior pronotal margin (ci = 1.00; ri = 1.00) (Fig. 1.8)

22

0: rounded
1: weakly sulcate

2: transversely impressed

27. Emarginate anterior pronotum (ci = 0.50; ri = 0.50)
0: straight

1: emarginate

28. Transverse medial groove on male pronotal base (ci = 0.50; ri = 0.75)
0: absent

1: present

29. Male pronotum sculpturing (ci = 0.67; ri = 0.86)
0: smooth to granulate
1: rugose
2: strigulate
3: aciculate

4: anterior half strigulate, posterior punctate

30. Lateral margin of pronotum (ci = 1.00; ri = 1.00) (Fig. 1.9)
0: type A
1: type B
2: type C

23

3:typeD
4:typeE
5:typeF
6:typeG
7:typeH
8:type1

31. Pronotum base (ci = 0.33; ri = 0.70) (Fig. 1.10)
0: straight
1: recurved
2: bisinuate

3: broadly emarginate along medial half

32. Pronotum apex (ci = 0.50; ri = 0.67)
0: narrower than base

1: equal width of base

33. Pronotum vestiture (ci = 0.50; ri = 0.47)
0: glabrous
1: anterior and lateral areas
2: anterior margin and angle
3: diagonally extending from anterior margin to median area

4: anterior area

24

5: entire

6: anterior third

7: anterior margin

8: entire except for median section of dorsal area
9: margins

A: lateral margin

Abdomen
34. Metepisternal setae (ci = 0.40; ri = 0.79)
0: biﬁd or triﬁd
l: palmately divided into four or more ﬁlaments

2: variable (scales, single setae and palmately divided seta)

35. Metepisternum shape (ci = 1.00; ri = 1.00) (Fig. 1.11)

0: type A
1: type B
2: type C
3: type D
4: type B
5: type F
6: type G
7: type H
8: typeI

25

9ztypeJ

AztypeK
thypeL
C:typeM
D:typeN
EztypeO
thypeP

GztypeQ
HztypeR

36. Mesosternum shape (ci = 0.94; ri = 0.93) (Fig. 1.12)
0: type A
1: type B
2: type C
3: type D
4: type B
5: type F
6: type G
7: type H
8: type I
9: type J
A: type K
B: type L

26

CztypeM
thypeN
EztypeO

F:typeP

37. Basal margin of elytra (ci = 0.27; ri = 0.58) (Fig. 1.13)
0: ﬂat
1: faint carina
2: carina extending to scutellar apex

3: crenulations

38. Scutellum shape (ci = 1.00; ri = 1.00) (Fig. 1.14)
0: type A
1: type B
2: type C
3: type D
4: type B
5: type F
6: type G
7: type H
8: typeI
9: type J
A: type K

27

thypeL
C:typeM
D:typeN
EztypeO

F: typeP

39. Depressed scutellum (ci = 0.50; ri = 0.83)
0: depressed

1: level of elytra

40. Elytral base with a tumescense from seventh to ninth interstriae (ci = 0.20; ri = 0.56)
0: absent

1: present

41. Discal interstriae punctures (ci = 0.10; ri = 0.31)
0: normal

1: confused

42. Disc sculpturing (ci = 0.33; ri = 0.63)
0: smooth
1: rugose
2: shallow rugosities

3: disc absent

28

43. Discal interstriae I and 2 ending in a large spine (ci = 1.00; ri = 0.0)
0: absent

1: present

44. Strial punctures (ci = 0.33; ri = 0.33)
0: larger than interstriae
1: equal to interstriae

2: smaller than interstriae

45. Carinate costa originating an eighth interstria of male elytra (ci = 1.00; ti = 1.00)
0: absent

1: present

46. Carina originating at base of tenth interstria (ci = 0.50; ri = 0.83)
0: absent
1: shorter than metepistemum
2: longer than metepistemum

3: encompasses elytra

47. Rugae on interior margin of elytra (ci = 1.00; ri = 0.00)

0: absent

1: present

29

48. Elytra (ci = 0.50; ri = 0.50)
0: non-metallic

l: metallic

49. Maximum rows of setae on third discal interstria (ci = 0.33; ri = 0.65)
0: glabrous
1: one row
2: two rows
3: three rows

4: four rows

50. Setae of interstriae (ci = 0.33; ri = 0.63)
0: scales
1: hair-like
2: glabrous

3: scales and hairs

51. Female strial impression (ci = 0.20; ri = 0.50)
0: not impressed
1: weakly impressed

2: strongly impressed

30

ll

52. Density of elytral setae (ci = 0.25; ri = 0.61) (Fig. 1.15)

0:

1:

glabrous

light

: moderate

: dense; elytra surface not visible

53. Elytra vestiture coloration (ci = 0.50; ri = 0.82)

0:

1:

2:

monochrome
variegated bicolor

vestiture absent

54. Declivity sculpturing (ci = 0.33; ii = 0.00)

0:

1

55.

smooth

: recumbent asperities

Male declivity setae (ci = 0.29; ri = 0.50)

: less than 3 times longer than disc
: greater than 3 times longer than disc

: absent

56. Apex of declivital margin (ci = 1.00; ri = 1.00)

0:

1:

smooth

serrate

31

57. Abdomen proﬁle (ci = 1.00; ri = 1.00)

0:

1:

2:

gradually ascending to elytra
abruptly ﬂexed upward at posterior margin of visible segment 2

abruptly ﬂexed upward at anterior margin of visible segment 2

58. Male 2“ visible abdominal sternite (ci = 1.00; ri = 1.00)

0:

1:

2:

not different from other stemites
convex and protuberant at middle and distally distended (see Fig. 1.66 C)

rapidly ascending/vertical

59. Boafy coloration pattern (including head and pronotum) (ci = 0.18; ri = 0.47)

0:

1:

2:

60.

monochrome
bicolored

tricolored

Male stria I impression (ci = 0.25; ri = 0.36)

: not impressed
: slightly impressed
: moderately impressed

: strongly impressed

Male Genitalia

32

61. Apical oriﬁce/ejaculatory canal (ci = 0.50; ri = 0.67)
0: ﬂeshy pore absent

1: ﬂeshy pore

62. Sensory claspers (ci = 0.79; ri = 0.79)
0: scythe shape
1: spatula
2: cactus
3: straight
4: round
5: pointed
6: downward 'V'
7: non-sclerotized
8: ﬂat
9: 'V' shaped
A: tooth
B: square

C: loop

63. Lateral folds of median lobe (ci = 0.67; ri = 0.62)
0: folds thick and uniformly arcuate

1: one fold with an apical curved extension that folds under other fold

2: folds almost join on basal third

33

3: forming a broad point on apical third

4: thin and arcuate distally on basal half

5: folds form an acute point on basal quarter

6: narrow, forming a thicker subacute point in middle

7: thin and straight

8: lateral folds thick nearly touching, broadest and touching on apical third
9: broadest and nearly touching on apical third

A: deep and wrinkled

64. Apex of median lobe (ci = 0.31; ri = 0.36)
0: ﬂat
1: rounded with ﬂat lateral margins
2: mushroom shaped
3: broadly pointed

4: round

65. Relative length of median strut (length until lateral folds start) (ci = 0.18; ri = 0.47)
0: less than 1/4
1: 1/4-1/3

2: greater than 1/3

Feeding habit

66. Feeding habit (ci = 1.00; ri = 1.00)

34

0: phloeophagous

1: xylomycetophagous

Phylogenetic Analysis
The phylogeny was reconstructed using the criteria of parsimony implemented in

PAUP“ 4.0 b10 PPC (Swofford, 1993). A heuristic search was performed with 1,000
stepwise random additions with tree bisection-reconnection (TBR) for 39 taxa (28
ingroup, 11 outgroup). Characters were equally weighted. Bootstrap values were
calculated by performing 1,000 pseudoreplicates with simple additions in PAUP“.
Bremer support values were calculated by creating a constraint tree in TreeRot v.2
(Sorenson, 1999) and analyzed in PAUP“ with a heuristic search using 100 addition-

sequence replicates.

Species Concept

Information for 17 of 30 species was restricted to morphological characters. In
addition, variation among populations could not be assessed for ten species because they
are known from only one locality. Species were deﬁned using the phylogenetic species
concept of Wheeler and Platnick (2000) which recognizes species as the smallest
aggregation of sexual populations diagnosable by a unique combination of character
states. This concept is based on observable (morphological) characters and “is compatible
with phylogenetic study because speciation is associated with character transformation”

(Wheeler & Platnick, 2000).

35

RESULTS

Camptocerus was recovered as monophyletic, but is weakly supported with less
than 50% bootstrap support and a Bremer support value of 2. Phylogenetic analysis
produced 26 most parsimonious trees with a length of 432 steps (Fig. 1.16) and 64 of 66
characters were parsimony informative. There was a moderate amount of homoplasy (CI
= 0.488, R1 = 0.694) and roughly half the characters were consistent with the cladogram.
Moderate support was observed for terminal nodes while most deeper relationships
among clades within the genus were poorly supported (Fig. 1.16). Two Cnemonyx
species, originally described as Ceratolepsis (C. bolivae (Blackman), and C. insignis
Wood), formed a clade, and received the strongest support in the phylogeny (99

bootstrap/ 6 Bremer).

DISCUSSION

Examination of specimens revealed that several characters previously used to
delimit Camptocerus from Cnemonyx are homoplastic (Wood, 1982, 1986, 2007). These
characters include a scutellum that is at least 1.5 times wider than long, and suture 1 of
antennal club marked internally by a septum. The strength of the Ceratolepsis clade and
general lack of support for Cnemonyx suggest that Cnemonyx is in need of revision.
Camptocerus is comprised of ﬁve morphologically distinctive clades (Opacicollis,

Auricomus, Latipilis, Aeneipennis, and Costatus).

36

Members of the “Opacicollis” clade (Camptocerus annectens, C. inoblitus, C.
major, C. opacicollis, and C. unicomus sp. 11.) include three former Loganius species. All
superﬁcially resemble Cnemonyx with their small size (2.2 - 3.6 mm), elongate bodies
(2.0 - 2.4 times as long as wide) and light brown to black coloration. They are idenitiﬁed
by the following characters. Male apical epistomal margin unarmed (large tubercule in C.
unicomus); basal epistomal margin armed with a strongly developed carina; male frons
strongly excavated, ornamented with few erect golden setae lateral to medial line; setae
do not form bands (except C. unicomus); male antennal scape elongate and narrowly
rounded distally. Anterior margin of pronotum rounded in both sexes and the margin
between the eyes lined with scales; base weakly bisinuate (broadly emarginate along
medial half in C. unicomus); lateral margin of pronotum type B (C. unicomus type H).
Metepistemrun shape type D; metepistemum setae palmately divided into four or more
ﬁlaments. Scutellum shape type B (C. unicomus type P); elytra base with a tumescence
from seventh to ninth interstriae; carina originating at base of tenth interstria shorter than
metepistemum (absent in C. unicomus); elytra apex serrate. Female frons always ﬂat,
often with medial area of frons slightly impressed; unarmed; epistoma weakly impressed;
female gena with circular mycangium.

The “Auricomus” clade (Camptocerus auricomus, C. niger, C. occidentalis, C.
orientalis, and C. rectus) is the most distinctive clade in Camptocerus with species
readily diagnosed by their dense, variegated, spatulate setae or scales on the elytra
interstriae, strongly bisinuate pronotum base and by two thick bands of setae on the male
frons. Unfortunately, older specimens tend to have abraded vestiture on the elytra and

pronotum which makes identiﬁcation difﬁcult. Males also possess a carina on the basal

37

epistomal margin; lateral margins of epistoma angled inward; antennal scape elongate
and narrowly rounded distally (except elongate and narrow in C. occidentalis). They are
also diagnosed by the following characters. Anterior margin of pronotum rounded in both
sexes and the margin between the eyes lined with scales. Scutellum type C; no carina
originating at the base of the tenth interstria; elytral base never with a tumescence from
seventh to ninth interstriae; metepistemum with setae palmately divided into four or more
ﬁlaments; metepistemum type G; mesostemum type C. Female ﬁons always ﬂat and
unarmed; female gena with triangular mycangium.

The “Latipillis” clade (Camptocerus charpentierae, C. coccoformus sp. n., C.
doeli sp. n., C. igniculus sp. n., C. latipilis, C. mallopterus sp. n., and C. petrovi sp. 11.)
contains several closely related and morphologically similar species. All species are
stout- 1.6 to 2.0 times as long as wide (except C. dolei is elongate 2.1-2.3 times as long as
wide) and vary in color from orange to black. They are identiﬁed by the following
characters. The male apical epistomal margin is armed with a short rectangular carina
(short rounded tubercule in C. dolei); lateral margin of epistoma with vertical costa; basal
epistomal margin tumid at scape insertion; male frons is strongly excavated, with the
excavation bordering ocular margin (moderately excavated and excavation not bordering
ocular margin in C. dolei); ornamented with 2-7 golden setae lateral to medial line; setose
lateral edges; frons sparsely to moderately covered by erect setae; male scape elongate
and expanded distally. The anterior margin of pronotum rounded in both sexes and the
margin between the eyes is glabrous or lined with scales in both sexes; pronotum base
recurved; lateral margin type C; smooth to granulate; base weakly recurved (weakly

bisinuate in C. charpentierae). Scutellum type B (type D in C. dolei); elytra base never

38

with a tumescence from seventh to ninth interstriae; metepistemum type C (type R in C.
dolei); disc covered with shallow rugosities; mesostemum type B (type D in C.
coccoformus, C. mallopterus); carina originating at base of tenth interstria encompassing
elytra; white to yellow scales present on interstria; second visible sternite convex or
protuberant at middle, noticeably smaller in female (absent in C. charpentierae, C. doeli
and C. igniculus, ). Female ﬁ'ons ﬂat, oﬁen with medial area of frons slightly impressed;
unarmed; female gena with circular mycangium (triangular in C. dolez).

Species of the “Costatus” clade (Camptocerus costatus, C. piliﬁons sp. n., C.
quadridens, and C. zucca sp. 11.) share numerous synapomorphies. The clade exhibits
remarkable sexual dimorphism of the elytra. Male C. costatus and C. quadridens posess a
carinate costa originating on the eighth interstria. Camptocerus costatus also have deeply
impressed striae and setae originating from recumbent asperities on the interstriae. C.
quadridens also has a two spines originating from the ﬁrst and second discal interstriae
that extend over the declivity. These characters are not displayed by females. They are
also diagnosed by the following characters. The male apical epistomal margin is
unarmed; lateral margin with an arcuate costa pointing inward, entirely lined with setae;
basal epistomal margin with a short, rectangular, transverse carina (except C. zucca);
male irons strongly excavated and covered by long, dense, erect, yellow-white hairs;
male scape is elongate and broadly rounded distally. Anterior margin of pronotum
rounded with the margin between the eyes in both sexes glabrous; lateral margin type D;
pronotum base recurved. Elytra never metallic; scutellum shape type I; basal margin of
elytra with a weak carina extending to scutellar apex; elytra base never with a tumescence

from seventh to ninth interstriae; carina originating at base of tenth interstria shorter than

39

metepistemum; declivity intrerstriae setae uniserate; apex of declivity margin smooth.
Female frons always ﬂat and unarmed; short, sparse, setae present only on lateral margin
of from; setae present on epistoma; female gena with circular mycangium.

The “Aeneipennis” clade (Camptocerus aeneipennis, C. angustior, C. aterrimus,
C. distinctus sp. n., C. hirtipennis, C. mandelshtami, C. noel sp. n., C. pseudoangustior
sp. n., and C. suturalis) is the most speciose clade in the genus. Species are large (3.1 -
7.1mm) and vary in color from metallic bronze, green and blue to black, red and orange.
Species possess a sundry of synapomorphies and are identiﬁed by the following
characters. Male apical epistomal margin unarmed or armed by a medial tubercule; lateral
margin of epistoma arcuate costa pointing out (vertical costa in C. distinctus); basal
epistomal margin armed with a transverse, concave carina (tumid above scape insertion
in C. distinctus); ﬁons strongly excavated and densely covered with long white to yellow
hair-like setae (moderate in C. distinctus); male scape elongate and narrow. Pronotum
base straight (recurved in C. distinctus); lateral margin type B; anterior margin in males
partially to entire sulcate (round in C. distinctus) and round to partially sulcate in
females; margin between the eyes lined with scales or biﬁd setae in females or with biﬁd/
triﬁd setae or scales in males (glabrous in both sexes of C. distinctus). Elytral base with a
tumescence from seventh to ninth interstriae; metepistemum with biﬁd setae;
metepistemum type B; mesostemum type A; scutellum type A (C. distinctus type H);
elytra metallic (C. aeneipennis, C. angustior, C. noel) or normal; interstriae glabrous
(hair-like setae on C. hirtipennis and C. suturalis); elytra smooth (C. distinctus with rugae
in the sutural interspace); apex smooth. Female frons always ﬂat; medial area often with

an impressed area and with a shallow carina originating on the median line between the

40

eyes and terminating at the epistoma; ornamented with fewer setae than male; female
gena with crescent shaped mycangium (absent in C. suturalis).

Several novel characters were used in this study. The presence and type of
mycangia, shape of the metepistemum, (including the pleural suture and exposed
trianglur plate of postepimerum), the lateral margin of the pronotum and process of the
mesostemum were informative at the species level. Although, previously noted as a
potential reliable character (Hulcr et al., 2007b), this is the ﬁrst time that presence and
type of mycangia have been used for scolytines.

Camptocerus mycangia are semi-circular to circular structures on the female
gena (nonglandular brush mycangium as in Six, 2003) (Fig 1.6) and enlarged punctures
on the female vertex under the anterior margin of the pronotum (nonglandular pit
mycangium as in Six, 2003) (Fig. l. 17). Both structures are present in males, but are
greatly reduced in size. Attempts to verify these structures by cross-sectioning of female
beetle heads were unsuccessﬁrl due the rigidity of the cuticle, despite adherence to
softening protocols used for closely related taxa (Livingston & Berryman, 1972). These
structures are similar in position to those found on Scolytus ventralis (Livingston &
Berryman, 1972) and are present on all Camptocerus species except C. suturalis. Loss of
these structures appears associated with its ambrosia ﬁrngus-stealing habit, mycocleptism
(Hulcr, 2009.) (F ig.1.18).

There are three types of gena mycangia in Camptocerus: crescent shaped (Fig.
1.17 B), oval (Fig. 1.17 C), triangular (Fig. 1.17 D). Crescent-shaped mycangium are
found in all species of the Aeneipennis clade, except C. suturalis. The mycangium

consists of a crescent-shaped groove in the cuticle on the anterior margin the gular

41

striations extending from the level of the ventral aspect of the mandible to the level of
scape insertion. The groove gradually deepens from the anterior margin until it reaches
the posterior margin and abruptly ascends. The groove is posterior margin is lined with
short, thick setae that cover the groove. This type of mycangium is often obscured by the
pronotum and is easily visible when the head either turned or extended. Oval-shaped
mycangium are found in all species of the Opacicollis, Latipillis and Costatus clades. The
mycangium is an oval-shaped impressed area from the posteroventral margin of the eye
and gular striations to a posterior margin lined with dense, short, thick setae. Short setae
arise from the eye and gular striation margins and face the center of the impression. Two
to three rows of long setae originate in the impression and are directed toward the point
where the eye meets the gula striations. In the Opacicollis clade, the posterior margin is
lined with short, sparse setae. Triangular mycangium are found in all species of the
auricomus clade and C. dolei. The position is identical to that of the oval-shaped
mycangium. The gular striation margin and posteroventral margin are lined with four
rows of long, thick setae (twice the width of other setae) covering roughly the ventral half
of the mycangium. Setae are oriented toward center of the impression. The
posterioanterior margin is lined by short, sparse setae. Four rows of long setae originate
in the impression and are directed toward the point where the eye meets the gula
striations.

The enlarged punctures on the female vertex under the anterior margin of the
pronotum are present on all species. Each pit has a short-bristle like seta arising from the

posterior margin and pits are 3-5 times the diameter of other punctures and 3-10 times as

deep.

42

Interspeciﬁc variation in structures (ex. carinae, tubercles), degree and extent of

excavation in the male frons, frons vestiture, elytra vestiture and sculpturing are

extremely useful for classiﬁcation of Camptocerus. Characters such as the type of

mycangia, shape of the metepistemum, the lateral margin of the pronotum and process of

the mesostemum typically diagnose clades within Camptocerus.

Key to the males of the species of Camptocerus

This key does not include species solely known from females. If the specimen

belongs in the “Opacicollis” clade, examine species diagnoses for Camptocerus

annectens Wood or C. major (Eggers) before making a ﬁnal determination.

2(1)

3(2)

Metepistemal setae biﬁd or triﬁd. ......................................................... 2
Metepistemal setae palmately divided into four or more ﬁlaments. ............... 12
Elytra metallic or with metallic luster, glabrous. ....................................... 3
Elytra never metallic, may be glabrous or densely covered with setae. ............. 5

Apical epistomal margin armed with a median tubercle; anterior pronotal margin
declivous, not sulcate; basal half of prontoum slightly less rugose; pronotal
punctures larger, deeper; strial punctures shallow. Elytra metallic bronze. Known
from Andes Mountains ﬁ'om Colombia to Bolivia, 1000-3000m; 5.0 — 5.8 mm

long. ................................................................... C. angustior Eggers

43

4(3)

5(2)

6(5)

7(6)

Apical epistomal margin unarmed; anterior pronotal declivity sulcate, especially

towards sides ................................................................................. 4

Smaller species; ﬁrst discal striae weakly impressed; strial punctures deeper,
larger; lateral declivities on anterior margin of pronotum extending two-thirds of
distance to medial line. Elytra metallic bronze to black with a metallic lustre.
Costa Rica to Argentina; 4.4 - 5.9 mm long. ............ C. aeneipennis (Fabricius)
Larger species; elytral disk smooth, strial punctures shallow, faint; lateral
declivity on the anterior margin of the pronotum extending to median line. Elytra
metallic bronze, green or blue. Bolivia and Peru to Colombia and Suriname; 6.0 —

7.1 mm long. .......................................... C. noel Smith & Cognato, sp. n.

Elytra glabrous, smooth, shining (some setae may be present laterallY); ﬁnely

sculptured, median tubercle on apical epistomal margin ............................... 6
Elytra with setae present at least on interstriae. 8
Body black. ................................................................................. 7

Pronotum bicolored (posterior ﬁfth to two-ﬁfth’s orange, remainder black),
profemur and protibia red orange, remaining legs black; thorax, abdomen and

elytra black. Peru and Ecuador; 5 .9 — 6.5 mm long. . . . . C. mandelshtami Petrov

Smaller size; anterior pronotal declivity weakly sulcate. Peru to French Guiana;

3.1 — 4.5 mm long. ................................................... C. aterrimus Eggers

8(5)

9(8)

10(8)

Larger size; anterior pronotal declivity entirely sulcate. Brazil, Peru and Ecuador;

4.5 — 5.4 mm long. .................... C. pseudoangustior Smith & Cognato, sp. n.

Pronotum base recuved; elytra base with a carina extending to scutellar apex. ....9

Pronotum base straight; elytra base ﬂat or with a faint carina. ...................... 10

Short transverse carina at basal epistomal margin. Each elytron with two discal
spines arising from the ﬁrst to ﬁfth striae, with a sharp spine on the ﬁrst striae and
a blunt spine on the second extending over the declivity, and the declivity
originating in the recess formed by the spines. Color red brown. Panama; 3.4 — 3.7
mm long. .......................................... Camptocerus quadridens Blackman
No carina on the basal epistomal margin. Distinct orange and black color pattern,
pronotum orange with a black mid-dorsal triangle extending from the apex to the
basal third of the pronotum; elytra black with each elytron with an orange oval
spot encompassing the area ﬁ'om the basal margin of the disk and to the basal ﬁfth
of the elytra and from the sutural interspace to the eighth interstriae; ventral
surface of thorax and abdomen black. Ecuador; 4.0 - 4.1 mm long. ...............

..............Camptocerus zucca Smith & Cognato, sp. n.

Elytra black each elytron with eight rugae on the discal sutural interspace to

second interstria, each rugae with a single setae arisng ﬁ'om the center. A large

median tubercle is present on the apical epistomal margin; vertical carinae on the

45

lateral margin of the epistoma; basal epistomal margin tumid above scape

insertion. Ecuador; 5 .3 mm long. ...........................................................

................................................... Camptocerus distinctus Smith & Cognato, sp. n.
- Elytra never with rugae, ﬁnely sculptured; long yellow, recumbent setae present

on both striae and interstriae (often abraded in older specimens). .................. 11

11(10) Larger size; elytra solid black, elytra densely covered with recumbent setae,

 

surface barely visible. anterior pronotal declivity densely covered with long
yellow recumbent setae; anterior margin between eyes lined with biﬁd setae;
rugose basal medial area of pronotum; vestiture emarginate at median line; setae
diagonally extend along the lateral margin ﬁom the anterolateral margin. Brazil,
Ecuador; 4.9 — 5.6 mm long. ........................ Camptocerus hirtipennis Schedl

- Smaller size; head, antennae, thorax, pronotum, legs red-orange to red, abdomen
black; elytra covered with fewer setae, surface largely visible; anterior pronotum
margin between eyes lined with setae palmately divided into 3-4 ﬁlaments; black
elytron each with an orange oval spot extending from the lateral margin of the
elytral interspace to the tenth interstria and from the basal ﬁfth of elytra to the of
apical four-ﬁfths. Peru, Brazil and the Guianas; 3.6 — 5 .0 mm long.

...................................................... Camptocerus suturalis (Fabricius)

12(1) Frontal setae in two dense, distinct longitudinal bands near middle of frons,

sometimes merging distally. Basal setae long, and recumbent over anterior setae.

................................................................................................ 13

46

- Frontal setae never in two bands, glabrous, with sparse setae or setae denser or

longer towards the sides of frons. ....................................................... 18

13(12) Apical epistoma with a median tubercle ................................................. 14
- Apical epistoma unarmed, with or without trace of carina on basal epistomal

margin. ...................................................................................... 1 5

14(13) Larger size, body stout. Frontal setae in two thick bands which lie ﬂat against the
frons. Apical epistomal tubercle small, rounded; carina on basal epistomal margin
weakly developed; pronotum base bisinuate. Elytra covered in variegated
spatulate setae (often abraded in older specimens). Mexico to Venezuela and

Trinidad; 3.5 — 4.1 mm long ..................... Camptocerus auricomus Blandford

- Smaller size, body elongate. Frontal setae in two thin bands arcuate above the
ﬁons and combine on the median line between the eyes. Apical epistomal tubercle
large and broad, carina on basal epistomal margin strongly developed and
connecting medially with tubercule; pronotum base broadly emarginate along
medial half. Elytra covered in spatulate setae. Brazil; 2.5 mm long. ...............

...................................... Camptocerus unicomus Smith & Cognato, sp. 11.

15(13) Transverse carina on basal epistoma margin faint or obsolete. Pronotum base

bisinuate. Elytra covered in variegated spatulate setae (often abraded in older

specimens). Bolivia and Brazil to Colombia and the Guianas; 2.8 — 4.1 mm long.

47

............................................................ Camptocerus niger (Fabricius)

- Transverse carina on basal epistomal margin moderately to strongly developed.

................................................................................................ 16

16(15) Carina on basal epistomal margin arcuate, height uniform. Elytra covered in
variegated spatulate setae (often abraded in older specimens). Brazil and Peru; 3.2
— 3.7 mm long. .......................................... Camptocerus orientalis Eggers

- Carina on basal epistomal margin straight, height not uniform. ..................... 17

17(16) Basal epistomal carina highest at middle, nearly tuberculate. Elytra covered in
small, stout variegated spatulate setae (often abraded in older specimens). Striae
distinct, black, interstriae reddish. Ecuador to Venezuela; 3.5 — 4.3 mm long.
...... ,,, Camptocerus rectus Wood

- Carina on basal epistomal margin straight, highest above antennal insertion,
lowest at middle. Elytra black covered in yellow elongate spatulate setae (often
abraded in older specimens). Striae obscured. Bolivia and Peru; 4.0 — 4.5 mm

long. ................................................... Camptocerus occidentalis Eggers
18(12) Basal epistomal margin armed by either a complete transverse or short rectangular

carina; apical epistomal margin unarmed; scape elongate and rounded

distally ....................................................................................... 19

48

- Basal epistomal margin unarmed without complete carina or short rectangular
carina; apical epistomal margin with a medial rectangular carina or unarmed;

scape elongate and expanded distally. .................................................. 22

19(18) Basal epistomal margin armed by an arcuate transverse carina; pronotum base
straight to weakly bisinuate; elytral smooth at base around scutellum. ............ 20

- Basal epistomal margin armed by a short rectangular carina; pronotum base
recurved; elytra with subbasal transverse carina extending to scutellar apex and

transverse rugosities around scutellum ................................................. 21

20(19) Uniserate narrow scales on the elytra interstriae, minute hairs on striae. Strial and
interstrial punctures equal size. Argentina and Bolivia to Ecuador to Costa Rica;
2.2 — 3.6 mm long. ................................. Camptocerus opacicollis (Eggers)
- Very ﬁne and minute hairs on both striae and interstriae. Strial punctures 2-3
tirrres larger than interstriae. Argentina and Brazil; 2.4 — 3.4 mm long. ...............

............................................................ Camptocerus inoblitus (Schedl)

21(19) Elytral declivity strongly impressed below level of disk, occupying apical 4/5 of
elytra; carinate costa on eighth interstriae from basal ﬁfth and extending around
apex to sutural margin; striae moderately impressed; and bristling brown setae
originating from recumbent asperities on the interstriae. Peru and Ecuador to
Brazil, Guyana and Trinidad; 3.5 — 3.9 mm long. .......................................

................................................................................ Camptocerus costatus Chapuis

49

- Elytral declivity and striae unimpressed; carinate costa on the eighth interstria
absent; elytra smooth with short scales on interstriae. Ecuador; 2.6 — 3.1 mm long.

....................................... Camptocerus pilifrons Smith & Cognato, sp. 11.

22(18) Second visible abdominal sternite convex and distally distended or protuberant at

middle (Fig. 1.66 C) ........................................................................ 23

- Second visible abdominal sternite not as above. ...................................... 26

23(22) Elytra densely covered in pale recumbent scales, surface largely covered. Head
and pronotrrm orange, elytra and abdomen black. Ecuador; 3.8 — 3.9 mm long.
................................. Camptocerus mallopterus Smith & Cognato, sp. n.

- Elytra interstriae with small colorless scales, elytra surface clearly visible. ...... 24

24(23) Lateral margin of epistoma with tuft of setae on apical third; head dark red, nearly
black, pronotrrm dark red; abdomen and elytra black. Second visible abdominal
sternite less strongly convex, scarcely protuberant below level of visible sternite

1. Brazil and Ecuador; 2.7 mm long. ......................................................
................................... Camptocerus coccoformus Smith & Cognato, sp. n.

- Lateral margin of epistoma with tuft of setae on apical half; black. ................ 25

25 (24) Smaller species; frontal excavation devoid of setae; 10-12 setae on lateral and

50

dorsal margins combined. Second visible abdominal sternite strongly convex and
protuberant, height of declivity behind much less than length of sternite 3. Brazil

and Ecuador. 2.7 — 3.3 mm long. ........................ Camptocerus Iatipilis Schedl

Larger species; frontal excavation with 1-2 setae lateral to median line, 15-20
setae on lateral and dorsal margins combined. Second visible abdominal sternite
strongly convex and protuberant, height of declivity behind greater than length of

sternite 3. Ecuador; 4.0 mm long. Camptocerus petrovi Smith & Cognato, sp. 11.

26(22) Male frons with6-7 golden setae lateral to medial line; frontal excavation does not
touch ocular margin; elytra base black with an orange patch an orange patch
covering basal third of elytra from the suture to eighth interstriae. Ecuador; 3.1 —

3.5 mm long. ........................... Camptocerus dolei Smith & Cognato, sp. 11.
Male ﬁ'ons with 2-3 golden setae lateral to medial line; ﬁ'ontal excavation borders

ocular margin; elytra color uniform. .................................................... 27

27(26) Larger species. Pronotum red or balck; elytra black. Elytral interstrial scales

semirecumbent, shorter, sparse and pale. Brazil to French Guiana; 3.8 — 4.5 mm

long. ................................................ Camptocerus charpentierae Schedl

Smaller species. Body uniformly red. Longer semi-recumbent, overlapping

golden scales on the elytral interstriae. Brazil; 2.5 — 2.8 mm long.

....................................... Camptocerus igniculus Smith & Cognato, sp. n.

Taxonomy

51

Genus Camptocerus Dej ean
Camptocerus Dejean 1821: 100. Type species: Hylesinus aeneipennis Fabricius.
Diagnosis. Camptocerus is distinguished from other Scolytini genera by the large,
circular mycangia present on the female gena (present in all species except C. suturalis
(F abricius), xylomycetophagy, gradually ascending abdominal proﬁle, the scutellum
ﬂush with the elytral surface, the more strongly excavated male frons, with the
excavation bordering ocular margin (except C. dolei), and from the epistoma to just
below the vertex, the male antennal funicle segments 2-7 ornamented with long setae
which forms the appearance of a brush dense brush on the distal edge 1 to 2 times the
length of frmicle, the absence of crenulations on the elytral base, generally larger body
size (2.2 — 7.1 mm long), female frons ﬂat, never convex, and by the lack of
supplemental denticles acutely margining the on apical anterior edge of the meso- and
metathoracic tibiae.
“Opacicollis” Clade
Camptocerus annectens Wood
(Figs. 1.23, 1.24)
Camptocerus annectens Wood, 2007: 210. Holotype, $2, BRAZIL: Serra de Baturite:
Ceara, (L. Gounelle); Strohmeyer Collection (DEI).
Diagnosis. This species is identiﬁed by the strigulate anterior third of the pronotum, basal
two thirds shining, punctures small; elytra declivity interstriae with uniseriate stout
scales, interstriae punctures as large as striae, and interstriae punctures with their bases

weakly to moderately elevated.

52

Description. Male unknown. Female 3.5 mm long (x = 3.5 mm; n = l); 2.2 times as long

as wide. Head, antennae, thorax, abdomen, elytra, black; legs dark brown.

Female epistoma impressed, 2-5 rows of setae lateral to the median line (mostly
abraded); frons ﬂat, medial area of ﬁons slightly impressed; frons and epistoma shagreen,
punctures large, deep spaced by 1-3 diameters of a punctrrre; frons and epistoma
unarmed; frons glabrous; gena with circular mycangium. Antennal scape elongate and
narrowly rounded distally; antennal funicle segments 5-7 with setae on dorsal margin less
than length of four ﬁrnicular segments; anterior face of the club setose with a partial
septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; apical
third strigulate; posterior two-thirds smooth, shining. Pronotum minutely punctate,
punctures shallow; stout setae present on apical fourth (mostly abraded); base weakly
bisinuate; lateral margin of pronotum type B.

Scutellum shape type B; elytra base with a tumescence from seventh to ninth
interstriae; elytra declivity interstriae with uniseriate stout yellow-brown scales (mostly
abraded); interstriae punctures as large as striae, and interstriae punctures with their bases
weakly to moderately elevated; carina originating at base of tenth interstria shorter than
metepistemum; metepistemum shape type D; metepistemum setae palmately divided into
four or more ﬁlaments. Apex serrate.

Specimens examined. (06‘, 12)
Type material: Holotype Camptocerus annectens, S? (DEI).
Distribution. Brazil (Ceara).

Hosts. Unknown.

53

Biology. Unknown.

Camptocerus inoblitus (Schedl)

(Figs. 1.44, 1.45)

Camptocerus inoblitus (Schedl) 1939: 722 (Loganius). Holotype 6‘: Brazil: Santa
Catharina; viii. 1940, F. Plaumann, Schedl Collection (NHMW).
Camptocerus morio (Schedl) 1952: 348 (Loganius). Holotype 6‘: Brazil: Santa Catharina;
Schedl Collection (NHMW). syn. n.
Diagnosis. This species is diagnosed by the brown color and short, very ﬁne hair-like

setae on both striae and interstriae.
Description. Male 2.4 — 3.4 mm long (i = 3.1 mm; 11 = 20); 2.1 — 2. 4 times as long as

wide. Head, antennae, thorax, abdomen, elytra and legs light to dark brown.

Apical epistomal margin unarmed; epistoma impressed; lateral margin with a
vertical sub-acute carina; tufts of 34 rows of setae along lateral margin; basal epistomal
margin armed with a strongly developed sub-acute arcuate carina; epistoma expanded to
greater than quarter length of head; male frons strongly excavated, excavation borders the
ocular margin; frons ornamented with 3-4 rows of erect pale setae lateral to medial line;
shagreen; scape elongate and narrowly rounded distally; scape with setae on distal third;
setae half length of scape; funicular segments 2-7 lined with brushes of setae the length
of scape; segments 1-7 with setae on dorsal margin less than length of four segments;
anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales;

anterior third strigulate; smooth, shining. Pronotum minutely punctate, punctures

54

shallow; stout setae present on apical fourth; base weakly bisinuate; lateral margin of
pronotum type E.

Scutellum shape type B; elytra base with a tumescence from seventh to ninth
interstriae; elytra interstriae with uniseriate minute pale hairs; striae punctures 2-3 times
larger than interstriae; striae punctures 2-3 times deeper than striae; striae with uniseriate
rows of minute pale hairs; carina originating at base of tenth interstria shorter than
metepistemum; metepistemum shape type D; metepistemum setae pahnately divided into
four or more ﬁlaments. Apex serrate.

Female similar to male except epistoma less impressed, three rows of setae lateral
to the median line; frons ﬂat, medial area of frons slightly impressed; frons with three
rows of setae on lateral margin; frons and epistoma shagreen, punctures small; frons and
epistoma unarmed; frons glabrous; antennal funicle segments 5-7 with setae on dorsal
margin and 3-7 with setae on ventral margins less than length of ﬁve funicular segments;
gena with circular mycangium.

Specimens examined. (216, 33 S2)

Type Material. Holotype, Camptocerus inoblitus, 6‘ (NHMW); Allotype Camptocerus
inoblitus, Q (NHMW); Holotype Camptocerus morio, 6‘ (NHMW).

Other material. BRAZIL: Santa Catharina: Nova Teutonia, [27°11’S, 52°23’W (on
most labels)], 300-500 it, viii.l940, (F. Plaumann), 66‘, 552 (FMNH); xi.1940, 36‘
(FMNH); 1949, 46‘ (FMNH); ex. Sapindaceae sp. 88, 14.x.1949, 252 (FMNH); 4.ii.1950,
36, 8Q (FMNH); ex. ‘trockenes Reisig’ [= dry brushwood], 7.ii.1950, 3Q (FMNH);
1.iii.1950, 16‘, 59 (FMNH); ex. ‘trockenes Reisig’ [= dry brushwood], 10.ii.1950, 29

(FMNH); 7.iii.l950, 26, 752 (FMNH).

55

Distribution. Argentina (Misiones), Brazil (Santa Catharina).
Hosts. Wrocatpusﬁondosus Fr. Allem, Sapindaceae sp.

Biology. Unknown.

Camptocerus major (Eggers)
(Figs. 1.49, 1.50)
Camptocerus major (Eggers), 1929: 60 (Loganius). Holotype E2: Peru: [Junin]
Chanchamajo; Eggers Collection (NHMW).
Diagnosis. The female is diagnosed by the strigulate apical third of pronotum and
punctate posterior; short, ﬁne setae on the declivity interstriae and shorter, thinner setae

on the striae.
Description. Male unknown. Female 3.4 mm long (i = 3.4 mm; n =1); 2.3 times as long

as wide. Head, antennae, thorax, abdomen, elytra and legs dark brown, nearly black.
Female epistoma impressed, 4-5 rows of setae; frons ﬂat, medial area of ﬁ'ons
slightly impressed; frons with 4-5 rows of setae lateral to median lines; frons and
epistoma shagreen, punctures on lateral margins larger, deeper; spaced by 1-3 diameters
of a puncture; frons and epistoma unarmed; ﬁ'ons glabrous; gena with circular
mycangium. Antennal scape elongate and narrowly rounded distally; antennal funicle
segments 5-7 with setae on dorsal margin and 3-7 with setae on ventral margins less than
length of ﬁve funicular segments; anterior face of the club setose with a partial septum.
Anterior pronotal margin rounded, margin between eyes lined with scales; apical

third strigulate; posterior two-thirds smooth, shining. Pronotum minutely punctate,

56

punctures shallow; stout setae present on apical fourth; base weakly bisinuate; lateral
margin of pronotum type B.

Scutellum shape type B; elytra base with a tumescence from seventh to ninth
interstriae; elytra smooth; elytra interstriae with uniseriate minute pale setae; interstriae
punctures smaller than striae; striae punctures 2-3 times deeper than striae; striae with
uniseriate rows of minute hairs; carina originating at base of tenth interstria shorter than
metepistemum; metepistemum shape type D; metepistemum setae palmately divided into
four or more ﬁlaments. Apex serrate.

Specimens examined. (06‘, 19)

Type Material. Holotype Loganius major, S? (NHMW).
Distribution. Brazil, Peru (Huanuco, Junin).

Hosts. Unknown liana.

Biology. Unknown.

Camptocerus opacicollis (Eggers)
(Figs. 1.61, 1.62)

Camptocerus opacicollis (Eggers), 1929: 61 (Loganius). Holotype 6‘: Bolivia: [Cercado],
Cochabamba; Eggers Collection (NHMW).
Camptocerus aquilus Wood 1972: 244. Holotype 6‘: Brazil: Goias [=Goias]: near Aldeia
Caraya, on or near the Rio Araguaia: 12°49’S 51°46’W; (BMNH).
Synonymy: Wood 1985: 266.
Camptocerus inﬁdelis Wood, 1969: 11. Holotype 82: Costa Rica: Peralta: Cartago, 1500

ft, 10.iii.1964, S.L. Wood, unknown vine; NMNH. syn. n.

57

Camptocerus uniseriatus Schedl, 1972: 54. Holotype 6‘: Brazil: Guanabara [=Rio de
Janeiro], Corcovado, x.1969, Alvarenga & Seabra (NHMW). syn. n.
Diagnosis. Both sexes are diagnosed by the larger, deeper punctures on the pronotum,

and uniseriate narrow scales on the elytra interstriae.
Description. Male 2.2 — 3.6 mm long (It = 2.9 mm; 11 = 17); 2.0 — 2.4 times as long as

wide. Head, antennae, thorax, and elytra dark brown to black. Gena, abdomen and legs
light to dark brown.

Apical epistomal margin unarmed; epistoma impressed; lateral margin with a
vertical sub-acute carina; 4-5 rows of setae lateral to median line; basal epistomal margin
armed with a strongly developed sub-acute arcuate carina; epistoma expanded to greater
than quarter length of head; male ﬁons strongly excavated, excavation borders the ocular
margin; frons ornamented with 5-6 rows of erect pale setae lateral to medial line;
shagreen; scape elongate and narrowly rounded distally; scape with setae on distal two-
thirds; setae length of scape; funicular segments 2-7 lined with brushes of setae 1.5 times
the length of scape; segments 1-7 with setae on dorsal margin less than length of six
segments; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales;
anterior third strigulate; smooth, shining. Pronotum densely punctate, punctures shallow;
stout setae present on apical fourth; base weakly bisinuate; lateral margin of pronotum
type B.

Elytra smooth; scutellum shape type B; elytra base with a tumescence from
seventh to ninth interstriae; each strial puncture with a hair-like seta arising from basal

margin, covering puncture; interstriae punctures equal size of striae; as deep as striae;

58

interstriae with uniseriate rows of narrow scales; carina originating at base of tenth
interstria shorter than metepistemum; metepistemum shape type D; metepistemum setae
palmately divided into four or more ﬁlaments. Apex serrate.

Female similar to male except epistoma less impressed, three rows of setae lateral
to the median line; ﬁ'ons ﬂat, medial area of frons slightly impressed; frons with four
rows of setae on lateral margin; frons and epistoma shagreen, punctures small; frons and
epistoma unarmed; frons glabrous; antennal funicle segments 4-7 with setae on dorsal
margins and segments 5-7 with setae on ventral margins less than length of four funicular
segments; gena with circular mycangium.

Specimens examined. (336, 4652)

Type Material. Holotype Camptocerus opacicollis, 6‘ (NHMW). Allotype Camptocerus
opacicollis, Q (NHMW) BOLIVIA: Cochabamba, (Germain), 1907, (H. Donckier).
Holotype Camptocerus aquilus 6‘ (BMNH). Paratypes Camptocerus aquilus BRAZIL:
[Mato Grosso]: 12° 31’S, 51° 46’W, RS-RGS, 12-18.xi.1968, (R.A. Beaver), 16‘, 1S?
(NMNH); Holotype Camptocerus uniseriatus, 6‘ (NHMW). COSTA RICA: Holotype
Camptocerus inﬁdelis, $2 (NWH); Paratype Camptocerus inﬁdelis Peralta: Cartago,
1500 ft, ex. unknown vine 10.iii.1964, (S.L. Wood), S? (NMNH).

Holotype Camptocerus uniseriatus, 6 (NHMW).

Other material. BRAZIL: Amazonas: 69 km N Manaus, (G. Stevens), 7.xii.1979, 36‘, 3Q
(NMNH); Goyas [=Goias]: Jatahy, ix-xi.1897, 4Q (DEI); ECUADOR: Napo: Reserva
Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778, -76.433333, 220 m, ex. canopy
fogging, i.1994, (T.L. Erwin et al.), lot 610, 192 (NMNH); Tiputini Biodiversity Station,

—0.631944, -76.144167, 220-250 m, ex. canopy fogging, ii.l999, (T.L. Erwin et al.), lot

59

2034, 12 (NMNH); lot 2053, 16‘ (NMNH). PERU: Madre de Dios: Los Amigos
Biological Station, -12.56916, -70.100114, 250 m, ex. Protium amazonicum, 17-
25.v.2008, (Smith, Hulcr), 76‘; 6S? (MSUC); 26-27.v.2008, 196‘; 2692 (MSUC);
Tampopata, 15 km NE Puerto Maldonado, Maldonado Reserva Cuzco Amazonico,
12°33’S 69°03’W, 200 m, ex. ﬂight intercept trap, 15.vi.l989, (J. S. Ashe, A. Leschen),
IQ, (SMEC).

Distribution. Argentina, Bolivia (Cercado), Brazil (Amazonas, Goias, Mato Grosso, Rio
de Janeiro), Costa Rica, Ecuador (N apo), Peru (Madre de Dios).

Hosts. Hirtella sp., Protium sp. (“Almesca branca” [Brazil], “Almesca vermelha”
[Brazil], “Breu branco” [Brazil]) (Beaver, 1972). Protium amazonicum Swart. Unknown
1 cm diameter tree sapling (Wood, 1969)

Biology. Beaver (1972) provides a detailed account of life history information for C.
aquilus and Wood (1969) describes the gallery of C. inﬁdelis, both are synonyms for C.

opacicollis.

Camptocerus unicomus Smith & Cognato, sp. 11.
(Figs. 1.78, 1.79)
Type Material. Holotype, 6‘, BRAZIL: ‘Estado Amazonas’ 010BR Chplac, km 31,
18.vi.1976, (E. Ruﬁno) (NMNH).
Diagnosis. The male is diagnosed by the large and broad tubercle on the distal epistomal
margin medially connecting with carina on basal epistomal margin, two narrow bands of

long yellow setae arising above the dorsolateral margin of the eyes which curve above the

60

hens and combine on the median line between the eyes; pronotum base broadly

emarginate along medial half and uni- and diseriate spatulate setae on interstriae.
Description. Male 2.5 mm long (i = 2.5 m; n = 1); 2.1 times as long as wide. Head,

antennae, thorax, legs, abdomen and elytra light to dark red brown.

Apical epistoma with a large and broad medial tubercle; lateral epistoma margin
with a faint costa angled inward; epistoma with six diagonal rows of setae ﬁ'om proximal
epistomal margin to tubercle; proximal epistomal margin armed with a strongly
developed carina connecting medially with epistomal tubercule; epistoma weakly
excavated; frons strongly excavated, two narrow bands of long yellow setae arise above
the dorsolateral margin of the eyes and curve above the frons and combine on the median
line between the eyes; frons ornamented with 7-8 rows of small and ﬁne and long thick
erect golden setae lateral to medial line; scape elongate and narrowly rounded distally;
scape with setae on distal third; setae less than half length of scape; ﬁrnicular segments 2-
7 lined with brushes of setae 1.5 times the length of scape; segment 7 with setae on dorsal
margin less than length of two segments; anterior face of the club setose with a partial
septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; surface
minutely, densely punctate; punctures shallow; stout setae present on apical fourth; lateral
margin of pronotum type H; base broadly emarginated along medial half.

Scutellum shape type P; interstriae densely covered with uni- and diseriate
spatulate setae; each setae arising from granules on interstria; each strial puncture with a
hair-like seta arising from basal margin, covering puncture; elytra base with a tumescence

from seventh to ninth interstriae; carina absent at base of tenth interstria shorter than

61

metepistemum; metepistemum shape type D; metepistemum setae palmately divided into
four or more ﬁlaments. Apex serrate.

Female unknown.

Etymology. Uni- (L) = one, -comus (L) = born. In reference to the large tubercle on the
distal epistomal margin.

Specimens examined. (16‘, 09)

Distribution. Brazil (Amazonas)

Hosts. Unknown.

Biology. Unknown.

“Auricomus” Clade
The “Auricomus” Clade is the most distinctive clade in Camptocerus with
species readily diagnosed by their distinctive dense, variegated, spatulate setae or scales
on the elytra interstriae, strongly bisinuate pronotum base and by two thick bands of setae
on the male frons; scutellum type C; no carina originating at the base of the tenth

interstria; metepistemum type G; mesostemum type C.

Camptocerus auricomus Blandford
(Figs. 1.27, 1.28)
Camptocerus auricomus Blandford, 1896: 125. Type locality: Panama: Chiriqui, V. dc
Chiriqui, below 40000 ft. Champion; Lectotype 6 (here designated). (BMNH).
Camptocerus striatulus Hagedom, 1903: 547, Type locality: French Guiana: Camopi,

1900, F. Geay; MNHN. Holotype SB, MNHN. syn. n.

62

Diagnosis. The male is diagnosed by its median small rounded tubercle on the apical
epistomal margin, an sub-acute recurved carina on the basal epistomal margin, ﬁons
strongly excavated with dorsal-lateral margin of the eye on excavated area of the frons,
the rounded margins of the frons excavation, two thick bands of setae on the male frons,
elytra interstriae covered in 2 to 3 rows of variegated uniseriate scales, and thinner and
pointed setae on apical quarter.

Female is distinguished from C. niger by and hair-like setae on apical quarter of
declivity and from C. orientalis by the presence of a circular, weakly impressed region

between upper level of eyes.
Description. Male 3.5 — 4.1 mm long (a = 3.9 mm; 11 = 20); 1.8 — 2.0 times as long as

wide. Color uniform; head, thorax, abdomen, elytra, antennae and legs light orange to
black. Elytra interstriae covered in tan scales with dark brown scales often forming a ‘X’
shape. Occasionally the sides of the ‘X’ are ﬁlled with brown scales forming a triangle on
each elytron.

Apical epistomal margin with a small rounded tubercle; lateral margins of
epistoma angled inward, lined with a single row of setae; a sub-acute, transverse recurved
carina on the basal epistomal margin; epistoma weakly excavated, expanded to less than
quarter length of head; ventral margin of frons elevated above epistoma; frons strongly
excavated with dorsolateral margin of the eyes on ﬁons excavation, lateral and dorsal
margins are rounded; frons densely covered with two thick bands of long yellow-brown
setae, clearly divided between the median line with lateral ﬁfths of frons and medial ﬁfth
glabrous; single row of setae lining lateral margins; frons surface shagreen; antennal

scape elongate and narrowly rounded distally; scape with a lateral row of setae on ventral

63

 

margin, setae less than half length of scape; funicular segments 2-7 lined with brushes of
setae 1.5 times the length of scape; segments 5-7 with setae on dorsal margin less than
length of four segments; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; medial
area of anterior third of pronotum covered with long bristles on the anterolateral areas;
base and disc covered with ﬁne, short hair-like setae; short bristles form two to three
rows above lateral carinae; pronotum strigulate on apical third and punctate basally;
punctures and vestiture on median line; densely and minutely punctured; punctures deep;
carina on lateral margin type F; base strongly bisinuate.

Declivity occupying apical half of elytra; each interstria densely covered with
three rows of semi-erect spatulate setae on disc, and two rows on declivity; scales
transition to uniseriate thinner and pointed setae on apical quarter; interstriae three times
width of striae; interstriae punctures strongly confused; striae weakly impressed; each
puncture with a hair-like seta arising from basal margin, covering puncture; scutellum
type C; no carina originating at the base of the tenth interstria; metepistemum with setae
palmately divided into four or more ﬁlaments; metepistemum type G; mesostemum type
C; apex smooth.

Female similar to male except epistoma less impressed and unarmed; epistoma
covered by ﬁve rows of setae; frons ﬂat, with a circular, weakly impressed region
between upper level of eyes; surface shagreen; 3-4 rows of setae lateral to median line;
antennal funicle segments scattered with setae less than length of six funicular segments;

gena with triangular mycangium; anterior third of pronotum covered with long, dense

64

hair-like setae on anterolateral areas; long bristles absent; base and disc covered with
ﬁne, short hair-like setae.

Specimens examined. (376, 27 SB)

Type Material. Paralectotypes Camptocerus auricomus 26 19 Panama: Chiriqui, V. dc
Chiriqui, below 40000 ft. (Champion) (BMNH). Paralectotypes here designated.
Holotype Camptocerus striatulus, Q (MNHN).

Other material. COSTA RICA: Limon: Reventazon ebene, Hamburgfarm, 24.ii. 1937,
(F. Nevermann), 36 (DEI); Peralta: Cartago, 500 m, 4” limb, 10.iii.1964, (S.L. Wood),
56, 59 (MSUC); Puntatenas: Gromaco, Rio Colo Brus, 1 quarter” trail cutting,
23.vii.1963, (S.L. Wood), 16, IS? (MSUC); No speciﬁc locality: ‘Rio Damites’, 200m,
18.ii.1964, (S.L. Wood) 16, 1S? (MSUC). GUATEMALA: Petén: Tikal, 100m, at light
at camp, 10.iv.1956, (T.H. Hubbell, I.J. Cantrall), l6 (UMMZ). MEXICO: Veracruz:
pranapa (Hidalgotitlan), 27.iv.1982, ex. Protium copalifera [likely Protium copaI],
(T.H. Atkinson), 16, 19 (CAS). PANAMA: Canal Zone: Barro Colorado Island, ex. trap,
17.ii.1978, (Henk Wolda), 16 (STRI); Madden Forest, 3” limb, 2.i.1964, (S.L. Wood) 16,
192 (MSUC); Chiriqui, V. dc Chiriqui, below 40000 ft. Champion 36 (BMNH);
Panama, Parque Nacional Soberania, Pipeline Rd, 9.166667, -79.75, 95 m, 3.ix.2008,
(S.M. Smith, A.D. Smith, A.R. Gillogly), 56, SS? (MSUC), 56, SS? (STRI).
VENEZUELA: Mérida: 20 km SW Vigia, 50 m, ex. limb, 21 .xi.1970, (S.L. Wood), 76,
692 (MSUC).

Discussion. C. striatulus is recognized as a synonym of C. auricomus due to the presence

of thinner and pointed setae on apical third of declivity.

65

 

Distribution. Costa Rica, French Guiana, Guatemala, Mexico (Veracruz), Nicaragua,
Panama, Trinidad, Venezuela (Mérida).

Hosts. Protium copal (Schltdl. & Cham.) Engl., Cedrela sp., Protium sp., Protium
glabrum (Rose) Engl., Protium panamense (Rose) I.M. Johnst, Rheedia edulis
[=Garcinia intermedia (Pittier), Hammel].

Biology. Atkinson & Equihua Martinez (1986) provide a detailed account of host use and
gallery description. C. auricomus excavate a short circumferential gallery below the bark
on both sides of the entrance trrnnel. Males use these galleries and the entrance tunnel to

turn around without leaving the gallery (Smith, per. 0b.).

Camptocerus niger (F abricius)
(Figs. 1.55, 1.56)

Camptocerus niger (Fabricius) 1801: 393 (Hylesinus). Lectotype 6: Essequibo [River]
[=Guyana] (UZMC). Lectotype designated Wood 1982: 415.
Camptocerus squammiger Chapuis, 1869: 259. Lectotype 6: Cayenne [French Guiana]
(IRSNB). Lectotype here designated. Synonymy: Eggers 1934: 27.
Camptocerus tectus Eggers 1943:245. Holotype 6: Brazil: Pemambuco: Sarra da
Bemada, 5.vi.1894, Duhant; Strohmeyer Collection, DEI. syn. n.
Diagnosis. This species is distinguished by unarmed apical epistomal margin, faintly
impressed epistoma, nearly indistinct recurved carina on the basal epistomal margin and

by two thick bands of setae on the male frons.

66

The female is distinguished from C. auricomus by the absence of hair-like setae
on declivital interstriae and from C. orientalis by frons with a circular, weakly impressed

region between upper level of eyes.
Description. Male 2.8 — 4.1 mm long (x = 3.5 mm; 11 = 19); 1.8 — 2.1 times as long as

wide. Color uniform, head, thorax, abdomen, elytra, antennae and legs light orange to
dark brown. Elytra interstriae covered in tan scales with dark brown scales creating a
variable pattern, oﬁen forming an ‘X’ shape. Occasionally the sides of the ‘X’ are ﬁlled
with brown scales forming a triangle on each elytron.

Apical epistomal margin unarmed; lateral margins of epistoma angled inward,
lined with a single row of setae; a weakly developed (nearly indistinct) sub-acute
recurved carina on the basal epistomal margin; epistoma faintly excavated; expanded to
less than quarter length of head; ventral margin of frons elevated above epistoma; ﬁons
strongly excavated with dorsal-lateral margin of the eye lies on excavated area of the
frons; lateral and dorsal margins are rounded; densely covered with two thick bands of
long yellow setae; setae are clearly divided between the median line; setae touch
proximal epistomal margin; lateral ﬁfths of frons glabrous; single row of setae lining
lateral margins; frons surface shagreen; antennal scape elongate and narrowly rounded
distally; scape with a row of setae on ventral margin; ventral margin of funicle segments
2-7 lined with brushes of setae 1.5 times the length of scape; dorsal margin with setae
less than four funicle segments; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; medial
area of anterior third of pronotum covered with long bristles on the anterolateral areas;

base and disc covered with ﬁne, short hair-like setae short bristles form two rows above

67

lateral carinae; antennal funicle segments scattered with setae less than length of six
funicular segments; pronotum strigulate on apical third and punctate basally; punctures
and vestiture on median line; densely and minutely punctured; punctures deep; carina on
lateral margin type F; base strongly bisinuate.

Declivity occupying apical half of elytra; each interstria densely covered with
three rows of semi-erect spatulate setae on disc, and one to two rows on declivity;
interstriae three times width of striae; interstriae punctures strongly confused; striae
weakly impressed; each puncture with a hair-like seta arising from basal margin,
covering puncture; scutellum type C; no carina originating at the base of the tenth
interstria; metepistemum with setae palmately divided into four or more ﬁlaments;
metepistemum type G; mesostemum type C; apex smooth.

Female similar to male except epistoma less impressed and unarmed; epistoma
covered by ﬁve rows of setae; frons ﬂat, with a circular, wealdy impressed region
between upper level of eyes; surface shagreen lateral margins may be strigulate; 3-4 rows
of setae lateral to median line; gena with triangular mycangium; anterior third of
pronotum covered with long, dense hair-like setae on anterolateral areas; long bristles
absent; base and disc covered with ﬁne, short hair-like setae.

Specimens examined. (346, 242, 1 unknown sex)

Type Material. Lectotype Camptocerus niger, 6 (MZMC); Paralectotypes Camptocerus
niger, NO LOCALITY: 3 SE, 1 unknown sex (MZMC); Paralectotypes Camptocerus
squammiger Q Cayenne [French Guiana] (IRSNB); Holotype Camptocerus tectus, 6
(DEI); Paratypes, Camptocerus tectus BRAZIL: Pemambuco: Sarra da Bemada,

5.vi.1894, (Duhant) 23 (DEI).

68

 

Other material. BOLIVIA: Beni: Cosincho Region, (G.L. Harrington), 16 NMNH); No
specific locality ‘Aragua’, 19 (NHMW). BRAZIL: Amazonas, Amazonas Parque, 30 lcrn
E Manaus, Amazon River, 110 ft, 17.ii.1981, (Chen-wen Young), is, 122 (RJRC) 19
(MSUC); AM 010 km 26, Reserva Ducke, ex. malaise trap, 21.ii.1978, (Jorge Arias), 16,
IS? (NMNH); Santa Catharina, Corupa (Hansa Humbolt), xi.1945, (A. Maller), l6
(AMNH). COLOMBIA: Cauca, 10 km SE Tambo, ex. ‘Anime’ [= local name of
Protium sp.], 9.vii.1970, (S.L. Wood), 76, 109 (MSUC); Putumayo: Mocoa, 26.vi.1974,
(M. Cooper), B.M. 1964-548, 19 (BMNH). ECUADOR: Napo: Reserva Ethnica
Waorani, 1 km S Onkone Gare Camp, -0.652778, -76.433333, 220 m, ex. canopy
fogging, i.1995, (T.L. Erwin et al.), lot 996, 16 (NMNH); x.1995, lot 1226, 16, 182
(NMNH); vii.l995, lot 1548, 19 (NMNH); Pastaza: Cononaco, -1.55, -75.583333, ex.
malaise trap, 29.v.1976, (J. Cohen), 26 (NMNH); FRENCH GUIANA: No specific
locality: ‘Cayenne’ 16 (BMNH); GUYANA: No speciﬁc locality: ‘Essequibo’ [River]
[=Guyana], 26 (MZMC). PERU: Loreto: 1.5 km N Teniente Lopez, -2.583333, -76.1,
210-240 m, ex. ﬂight intercept trap, 9-22.vii.l993, (R. Leschen), 192 (NMNH); 70 km
SSW from Iquitos to Nauta, 150 m, 5.iii.2008, (A. Petrov) 66, (MSUC). SURINAME:
Commewijne: Akintosoela, 32 km SE Suriname River bridge, road to Redi Doti,
5.271389,-70.2517, 40 m, ex. ﬂight intercept trap, 29.vi.-3.vii.1999, (Z.H. Falin, B.
DeDijn, A. Gangagin), l6 (SMEC); No speciﬁc locality: ‘Surinam’, 4S? (MZMC);
VENEZUELA: Aragua: Rancho Grande, 1500-1900m, 8.v.1970 (O ’Brien, Marshall)

16 (CAS). N0 SPECIFIC LOCALITY: ‘America meridionali’ [=South America], 26

(IRSNB).

69

 

Discussion. Camptocerus tectus is here designated as a synonym of C. niger based on
lack of a transverse carinae on the male frons and identical male genitalia. When Eggers
described C. tectus, only specimens of C. auricomus were examined for comparison and
not C. niger (Eggers, 1943).

Distribution. Bolivia (Beni), Brazil (Amazonas, Mato Grosso, Pemambuco, Santa
Catharina), Colombia (Cauca, Putumayo), Ecuador (N apo, Pastaza), French Guiana,
Guyana, Peru (Loreto), Suriname, Venezuela (Aragua).

Hosts. Hirtella sp., “Iratinga” (Beaver, 1972), Licania kunthiana Hook. f., Protium spp.
(“Almesca branca” [Brazil], “Almesca vermelha” [Brazil], “Breu branco” [Brazil])
(Beaver, 1972).

Biology. Beaver (1972) provides a detailed life history. C. niger females excavate a short
circumferential gallery below the bark on both sides of the entrance trmnel, identical to
those produced by C. auricomus and C. rectus. C. niger was also reported to partially
block their entrance tunnels with boring dust, a potential defense against predators

(Beaver, 1972), which can cause signiﬁcant Camptocerus mortality (chapter 2).

Camptocerus occidentalis Eggers
(Figs. 1.59, 1.60)
Camptocerus occidentalis Eggers, 1928: 91. Lectotype 6: Bolivia (NMNH).
Lectotype designated by Anderson & Anderson 1971: 23.
Diagnosis. The male is distinguished by lateral margins of epistoma with an arcuate costa
angled distally; basal epistomal margin with a straight carina hightest above scape

insertion and lowest at the middle, elongate and narrow scape.

70

The female is diagnosed by the elongate and narrow scape.
Description. Male 4.0 — 4.5 mm long (i = 4.3 mm; 11 = 5); 2.0 — 2.1 times as long as

wide. Color variable; head, antennae, pronotum, thorax, abdomen, and legs red-brown to
dark brown; elytra orange-red to black. Elytra interstriae covered in variegated tan and
dark brown scales forming highly variable patterns.

Apical epistomal margin unarmed; lateral margins of epistoma with an arcuate
costa a angled outward, lined with 6 rows of setae lateral to median line; basal epistomal
margin with a straight carina hightest above scape insertion and lowest at the middle;
epistoma strongly excavated; expanded to greater than quarter length of head; frons
strongly excavated with excavation bordering the ocular margin and the posterior margin
of the carina; dorsolateral margin of eyes not impressed; lateral margins sub-acute; frons
densely covered with two thick bands of long pale yellow setae; setae are clearly divided
between the median line; frons shagreen; setae touch proximal epistomal margin; lateral
quarters of frons and medial ﬁfth glabrous two to three rows of long setae lining lateral
margins; frons surface shagreen; antennal scape elongate and narrow; scape with a row of
long setae on ventral margin; ventral margin of funicular segments 2-7 lined with brushes
of setae 2 times the length of funicle; posterior margin of funicle segments with setae less
than four segments in length; setae on both ventral and dorsal margin equal thickness;
anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; medial
area of anterior third of pronotum covered with long, slender bristles on the anterolateral
areas; base and disc covered with ﬁne, short hair-like setae; short setae form two rows

above lateral carina; pronotum strigulate on apical third and punctate basally; median line

71

devoid of punctures and vestiture; densely and minutely punctured; punctures deep;
carina on lateral margin type F; base strongly bisinuate.

Declivity occupying apical half of elytra; each interstria densely covered with
three rows of semi-erect thin spatulate setae on disc, and one to two rows on declivity;
ﬁrst interstria with uniseriate spatulate setae on apical third; interstriae three to ﬁve times
width of striae; interstriae punctures strongly confused; striae weakly impressed; each
puncture with a hair-like seta arising from basal margin, covering puncture and
terminating at basal margin of apical puncture; scutellum type C; no carina originating at
the base of the tenth interstria; metepistemum with setae palmately divided into four or
more ﬁlaments; metepistemum type G; mesostemum type C; apex smooth.

Female similar to male except epistoma unarmed, moderately impressed; covered
by four rows of setae; frons ﬂat; surface shagreen; 5-7 rows of setae lateral to median
line; unable to examine antennae; gena with triangular mycangium; anterior third of
pronotum covered with long, dense slender hair-like setae on anterolateral areas; long
bristles absent.

Specimens examined. (66, 29)

Type Material. Lectotype Camptocerus occidentalis, 6 (NMNH); Paratypes BOLIVIA:
No speciﬁc locality, ‘Bolivia’, 16 (BMNH) 2S2 (NMNH).

Other material. PERU: Junin: T arma, Utcuyacu & Agua Dulce, iii.1948, (F.
Woytkowski), 16 (AMNH); No speciﬁc locality: ‘Peru’, (W. Muller Vermacht), 1909, 16
(NMNH), 26 (SMTD).

Distribution. Bolivia, Peru (Junin).

Hosts. Unknown.

72

Biology. Unknown.

Camptocerus orientalis Eggers
(Figs. 1.63, 1.64)
Camptocerus orientalis Eggers 1943:244. Holotype 6: Brazil: Bahia: Salobro, 6.vii.1885,
L. Gounelle; Strohmeyer Collection, DEI.
Diagnosis. The male is diagnosed by an unarmed median apical epistomal margin; a
transverse moderately developed arcuate carina on the basal epistomal margin, and the
declivity with spatulate setae to apex.
The female is diagnosed from C. auricomus by the declivity with spatulate setae
to apex.

Description. Male 3.2 - 3.7 mm long (i = 3.6 m; n = 4); 1.9 times as long as wide.

Color uniform, head, pronotum, thorax, abdomen, elytra, antennae and legs red-brown.
Elytra interstriae covered with variegated tan and dark brown spatulate setae forming
highly variable patterns.

Apical epistomal margin unarmed; lateral margins of epistoma angled inward,
lined with 1-3 rows of setae; a transverse moderately developed arcuate carina on the
basal epistomal margin, ventrally lined with two rows of setae; epistoma weakly
excavated; expanded to less than quarter length of head; from strongly excavated with
excavation bordering the ocular margin and the posterior margin of the carina; lateral and
dorsal margins are sub-acute; densely covered with two thick bands of long pale yellow
setae; single row of lining lateral margins; ﬁons shagreen; antennal scape elongate and

narrowly rounded distally; scape with a row of setae on distal half of ventral margin;

73

funicular segments 2-7 lined with brushes of setae two times the length of scape on
ventral margin; dorsal margin glabrous; anterior face of the club setose with a partial
septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; medial
area of anterior third of pronotum covered with long bristles on the anterolateral areas;
short bristles form three rows above lateral carinae; base and disc covered with ﬁne, short
hair-like setae; pronotum strigulate on apical third and punctate basally; punctures and
vestiture on median line; densely and minutely punctured; punctures deep; carina on
lateral margin type F; base strongly bisinuate.

Declivity occupying apical half of elytra; each interstria densely covered with
three rows of semi-erect spatulate setae on disc and one to two rows on declivity; ﬁrst
interstria with uniseriate spatulate setae on apical third; interstriae three times width of
striae; interstriae punctures strongly confused; striae weakly impressed; each puncture
with a hair-like seta arising from basal margin, covering puncture; scutellum type C; no
carina originating at the base of the tenth interstria; metepistemum with setae palmately
divided into four or more ﬁlaments; metepistemum type G; mesostemum type C; apex
smooth.

Female similar to male except epistoma unarmed, weakly impressed; covered by
ﬁve rows of setae; ﬁ'ons ﬂat, faintly impressed between eyes, surface shagreen; 3-4 rows
of setae lateral to median line; antennal funicle segments scattered with setae less than
length of six funicular segments; gena with triangular mycangium; anterior third of
pronotum covered with long, dense hair-like setae on anterolateral areas; long bristles

absent; base and disc covered with ﬁne, short hair-like setae.

74

Specimens examined. (46, 13?)

Type material: Holotype Camptocerus orientalis, 6 (DEI). Paratypes Camptocerus
orientalis, BRAZIL: Bahia: Salobro, 6.vii.1885, (L. Gounelle), 26, 1S? (DEI), 16
(NMNH).

Distribution. Bolivia, Brazil (Amapa, Bahia).

Hosts. Protium sp.

Biology. Unknown.

Camptocerus rectus Wood
(Figs. 1.74, 1.75)

Camptocerus rectus Wood, 1972: 245. Holotype 6: VENEZUELA: Barinas: 40 km E
Canton, 70 m, ex. Protium tenuifolium, 8.iii.1970, (S.L. Wood) (NMNH).
Diagnosis. The male is diagnosed by the transverse, moderately developed and straight,
sub-acute carina highest at middle, nearly tuberculate.

The female is diagnosed by a faint triangular impression ﬁ'om posterior margin of
epistoma to with vertex just below upper level of eyes.
Description. Male 3.5 — 4.3 mm long (x = 4.0 mm; 11 = 20); 1.8 - 2.1 times as long as
wide. Color uniform, head, pronotum, thorax, abdomen, elytra, antennae and legs red-
brown to dark brown. Elytra interstriae covered with variegated tan and dark brown
spatulate setae forming highly variable patterns.

Apical epistomal margin unarmed; lateral margins of epistoma angled inward,
lined with 1-3 rows of setae; a straight, sub-acute strongly developed straight carina on

the basal epistomal margin highest at middle, nearly tuberculate; carina lined ventrally

75

with three rows of setae; epistoma weakly excavated; expanded to less than quarter
length of head; ﬁons strongly excavated with excavation bordering the ocular margin and
the posterior margin of the carina; lateral and dorsal margins are rounded; ﬁ'ons densely
covered with two thick bands of long pale yellow setae; single row of spatulate setae
lining lateral margins; frons shagreen; antennal scape elongate and narrowly rounded
distally; scape with a row of setae on ventral margin; ventral margin of funicular
segments 2-7 lined with brushes of setae two times the length of scape; posterior margin
of segments 5-7 with setae as thick as ventral margin and less than four funicle segments;
anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; medial
area of anterior third of pronotum covered with long bristles on the anterolateral areas;
short bristles form two rows above lateral carinae; base and disc covered with ﬁne, short
hair-like setae pronotum strigulate on apical third and punctate basally; punctures and
vestiture on median line; densely and minutely punctured; punctures deep; carina on
lateral margin type F; base strongly bisinuate.

Declivity occupying apical half of elytra; each interstria densely covered with
three to four rows of semi-erect stout spatulate setae on disc, and one to three rows on
declivity; ﬁrst interstria with uniseriate spatulate setae on apical third; interstriae three to
four times width of striae; interstriae punctures strongly confused; striae not impressed;
each puncture with a hair-like seta arising from basal margin, covering puncture;
scutellum type C; no carina originating at the base of the tenth interstria; metepistemum
with setae pahnately divided into four or more ﬁlaments; metepistemum type G;

mesostemum type C; apex smooth.

76

Female similar to male except epistoma unarmed, weakly impressed; covered by
four rows of setae; frons ﬂat, with a faint triangular impression from posterior margin of
epistoma to with vertex just below upper level of eyes; surface shagreen; 5 rows of setae
lateral to median line; antennal funicle segments 5-7 with setae on dorsal margin less than
length of three funicular segments; gena with triangular mycangium; anterior third of
pronotum covered with long, dense hair-like setae on anterolateral areas; long bristles
absent; base and disc covered with ﬁne, short hair-like setae.

Specimens examined. (256, 292)

Type Material. Holotype Camptocerus rectus, 6 (NMNH); Allotype Camptocerus rectus
S2 (NMNH); Paratypes VENEZUELA: Barinas, 40 km E Canton, 70 m, ex. Protium
tenuifolium, 8.iii.l970, (S.L. Wood), 36, 59(NMNH); 9 km S Barrancas, 150 m, ex.
Protium sp., 2.xii.1969, (S.L. Wood), 26, 292 (NMNH); 10 km SE Miri, 150 m, ex.
Protium tenuifolium, 8.ii.1970, (S.L. Wood), 36, 192 (NMNH); 40 km SE Socopo, 150 m,
ex. Protium sp., 25.i.l970 (S.L. Wood), 46, 89 (NMNH).

Other material. ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare
Camp, -0.652778, -76.433333, 220 m, ex. canopy fogging, (T.L. Erwin et al.), lot 1464,
i.1996, 16 (NMNH); ex. canopy fogging lot 1496, i.1996, l6 (NMNH). VENEZUELA:
Barinas: 40 km E Canton, 70 m, ex. Protium tenuifolium, 8.iii.l970, (S.L. Wood), 16, 3S?
(MSUC); 10 km SE Miri, 150 or, ex. Protium tenuifolium, 9.ii.l970, (S.L. Wood), 26, 49
(MSUC); 40 km SE Socopo, 150 m, ex. Protium sp., 22.i.1970, (S.L. Wood), 26
(MSUC), 56, 55.2 (NMNH).

Distribution. Ecuador (N apo), Venezuela (Barinas).

Hosts. Protium spp., Protium tenuifolium [=Protium tenufolium (Engl.) Engl.].

77

 

Biology. Wood (2007) describes the biology as similar to that of C. aeneipennis, although
the gallery pattern likely resembles that of C. auricomus and C. niger based on their close
relationship to C. rectus. All three species excavate a short circumferential gallery below
the bark on both sides of the entrance tunnel. Colonization occurs in tree limbs with a

diameter less than 10 cm (Wood, 2007).

“Latipilis” Clade

Species in the “Latipilis” clade are diagnosed by the male lateral epistomal
margin with vertical costa; basal epistomal margin tumid above scape insertion; carina

originating at base of tenth interstria encompassing elytra; lateral margin of pronotrnn

type C.

Camptocerus charpentierae Schedl status 11.

(Figs. 1.29, 1.30)
Camptocerus charpentierae Schedl, 1970: 582. Holotype 6: French Guiana: Massikiri-

Haut Oyapock, 18.xi.1969, Balachowsky-Gruner Guyane Mission; Schedl Collection,

(NHMW).
Diagnosis. Both sexes of C. charpentierae are distinguished by the black abdomen, red

the normal second visible abdominal sternite and short pale scales on the interstriae.
Description. Male 3.8 — 4.5 mm long (.7 = 4.3 mm; n = 6); 1.7 — 2.0 times as long as

wide. Head, antennae dark red brown; abdomen, elytra, and legs black. Pronotrrm and

legs may be red to dark red brown.

78

Apical epistomal margin armed with a short rectangular carina; lateral margins of
epistoma with vertical costae; epistoma impressed; six rows of setae from lateral
epistoma margins to median line; basal epistomal margin tumid above scape insertion;
epistoma expanded to greater than quarter length of head; male frons is strongly
excavated, sparsely ornamented with one erect golden setae lateral to medial line; surface
clearly visible; 20-30 on lateral and dorsal margin; surface shagreen. Antennal scape
elongate and wider distally, dense setae length of scape on distal three-quarters; funicle
segments 2-7 with setae equal to 1.5 times funicle length on ventral margin; setae on
dorsal margin less than length of ﬁve funicle segments; setae on ventral margin twice as
thick as those on dorsal margin; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales;
anterior edge with 2-3 rows of yellow-brown setae; surface minutely granulate- punctate;
punctures minute, shallow; lateral margin type C; base weakly recurved.

Scutellum type B; base never with a tumescence from seventh to ninth interstriae;
disc shallow rugosites occupying ﬁrst to fourth striae and interstriae just beyond the
scutellum, remaining elytral surface minutely granulate; metepistemum type C, with
setae palmately divided into four or more ﬁlaments; mesostemum type B; carina
originating at base of tenth interstria encompassing elytra; un- and diseriate rows of
yellow-brown scales on interstriae; second visible abdominal sternite normal. Apex
smooth.

Female similar to male except epistoma not expanded, less impressed and
unarmed; epistoma with a four rows of hairs on apical margin; frons ﬂat, with medial

area of from slightly impressed, joining with impression of epistoma; punctures small,

79

shallow, larger and deeper laterally; antennal frmicle segments 1-7 with scattered setae on
less than length of three funicular segments; gena with circular mycangium. Pronotum
margin between the eyes lined with scales.

Specimens examined. (96, 8?)

Type Material. Holotype Camptocerus charpentierae, 6 (NHMW).

Other material. BRAZIL: Amazonas: Hwy ZF 2, km 20.7, ca 60km N. Manaus, 2°30’S,
60°15 ’W, ex. canopy fogging, l6.viii.l979, (Adis, Erwin, Montgomery et al.), 26, 1S?
(NMNH); 18.viii.1979, 16 (NMNH). ECUADOR: Napo: Tiputini Biodiversity Station, -
0.631944, -76.l44167, 220-250 m, x.1998, ex. canopy fogging (T.L. Erwin et al.), lot
1947, 16 (NMNH); ii.l999, lot 2030, 19 (NMNH); ii.l999, lot 2047, 132 (NMNH);
Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778,-76.433333, 220 m, ex.

canopy fogging, i.1994, (T.L. Erwin et al.), lot 644, 16 (NMNH); i.1995, lot 1000, 16
(NMNH); vi.l996, lot 1550, I? (NMNH); vi.l996, lot 1586, I? (NMNH); x.1996, lot
1753, 25, 352 (NMNH).

Discussion. This species was considered a synonym of Camptocerus cinctus (Wood,
2007). Camptocerus cinctus is a synonym of C. suturalis. Camptocerus charpentierae is
here resurrected based on comparison of the holotype with that of C. suturalis.
Phylogenetic analysis revealed the C. charpentierae (“latipilis” clade) to be in a separate
clade from C. suturalis (“Aeneipennis” clade) (Fig. 1.16).

Distribution. Brazil (Amazonas), Ecuador (N apo), French Guiana.

Hosts. Unknown.

Biology. Unknown.

80

Camptocerus coccoformus Smith & Cognato, sp. n.
(Figs. 1.31, 1.32)

Type Material. Holotype, 6, BRAZIL: Amazonas: Hwy ZF 2, km 20.7, ca 60 km N.
Manaus, 2°30’S, 60°15’W, ex. canopy fogging, l6.viii.l979, (Adis, Erwin, Montgomery
et al.) (NMNH); Paratypes, BRAZIL: 12, Amazonas: Hwy ZF 2, km 20.7, ca 60 km N.
Manaus, 2°30’S, 60°15’W, ex. canopy fogging, l6.viii.l979, (Adis, Erwin, Montgomery
et al.) (MSUC); 192, 69 km N Manaus, 7.xii. 1979, (G. Stevens) (NMNH); ECUADOR:
192, Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778,-76.433333,
220 m, ex. canopy fogging, i. 1994, (T.L. Erwin et al.), lot 626, (NMNH).
Diagnosis. The male is diagnosed by a tuft of setae on distal third of lateral epistoma
margins, red brown head, red pronotum black body, second visible abdominal sternite
less convex, scarcely protuberant below level of sternite 1.

Female can be distinguished from C. latipilis by the more impressed medial frons.
Description. Male 2.7 mm long (n = 1); 1.6 times as long as wide. Frons dark red brown;
gena, antennae and pronotum reddish. Abdomen, elytra and coxa black; and legs dark red
brown.

Apical epistomal margin armed with a short rectangular carina; lateral margins of
epistoma with vertical costae; epistoma impressed tuft of setae on distal third of lateral
epistoma margins; basal epistomal margin tumid above scape insertion; epistoma
expanded to greater than quarter length of head; male frons is strongly excavated,
ornamented with two golden setae lateral to medial line; surface shagreen; setose lateral
edges; frons sparsely covered by erect setae, surface clearly visible. Antennal scape

elongate and wider distally, four rows of setae half-length of scape on distal two-thirds;

81

funicle segments 2-7 with setae equal to 1.5 times funicle length on ventral margin; setae
on dorsal margin less than length of ﬁve funicle segments; setae on ventral margin twice
as thick as those on dorsal margin; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales;
anterior edge with 2-3 rows of yellow-brown setae; surface minutely granulate- punctate;
punctures minute, shallow; lateral margin type C; base weakly recurved.

Scutellum type B; base never with a tumescence ﬁ'om seventh to ninth interstriae;
disc shallow rugosites occupying ﬁrst to fourth striae and interstriae just beyond the
scutellum, remaining elytral surface minutely granulate; metepistemum type C, with
setae palmately divided into four or more ﬁlaments; mesostemum type D; carina
originating at base of tenth interstria encompassing elytra; uniseriate rows of yellow-
brown yellow scales on interstria; second visible abdominal sternite convex, scarcely
protuberant below level of sternite 1. Apex smooth.

Female similar to male except epistoma not expanded, less impressed and
unarmed; epistoma with a row of hairs on apical margin; frons ﬂat, with medial area of
ﬁ'ons slightly impressed, joining with impression of epistoma; antennal frmicle segments
3-7 with setae on dorsal margin less than length of three frmicular segments; gena with
circular mycangium; second visible abdominal sternite ventrally convex and distally
distended, noticeably smaller than male.

Etymology. Cocc- (G) = berry, -form (L) = shape. In reference to the species globular
and reddish appearance.

Specimens examined. (16, 3 S?)

82

 

Discussion: This species was identiﬁed as Camptocerus latipilis by Wood (2007).
Distribution. Brazil (Amazonas).
Hosts. Unknown.

Biology. Unknown.

Camptocerus dolei Smith & Cognato, sp. n.
(Figs. 1.38, 1.39)

Type Material. Holotype, 6, ECUADOR: Napo: Tiputini Biodiversity Station, -
0.631944, -76.144167, 220-250 m, ex. canopy fogging, ii.l999, (T.L. Erwin et al.), lot
2044, (NMNH held in trust for MECN). Paratypes, 2Q, ECUADOR: Napo: Tiputini
Biodiversity Station, -0.631944, -76. 144167, 220-250 m, ex. canopy fogging, ii.l999,
(T.L. Erwin et al.), lot 2044, (MSUC) (NMNH); l6, x.1998, lot 1944, (MSUC); 16,
i.1999, lot 2091 (NMNH).
Diagnosis. Both sexes are identiﬁed by an elongate body, elytra interstria with uni- and
diseriate golden setae on elytra; unique color pattern described below, the carina that
encompasses the elytra on the tenth interstria, The male is identiﬁed by its moderately
excavated ﬁ'ons with the excavation never bordering the eye.
Description. Male 3.1 — 3.5 mm long (Tc = 3.3 mm; 11 = 3); 2.1 — 2.3 times as long as
wide. Head, antennae and thorax orange; legs dark red brown; abdomen black; elytra
black with an orange patch covering basal third of elytra ﬁ'om the suture to eighth
interstriae.

Apical epistomal margin armed with a short rounded tubercle; lateral margins of

epistoma with vertical costa; epistoma weakly impressed; nine rows of setae from lateral

83

epistoma margins to median line; basal epistomal margin tumid above scape insertion;
epistoma expanded to greater than quarter length of head; male ﬁons is weakly
excavated, excavation not reaching ocular margin; moderately ornamented with four
rows of erect golden setae lateral to medial line; surface mostly visible; setose lateral
margins; surface shagreen. Antennal scape elongate and expanded distally, dense setae
two-thirds length of scape on distal half; funicle segments 2-7 with setae equal to 1.5
times ﬁmicle length of funicle on ventral margin; dorsal margin of 1-5 glabrous, setae on
segment 6 less than length of two funicle segments, setae on segment 7 length of funicle;
setae on ventral margin twice as thick as those on dorsal margin; anterior face of the club
setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; lateral
margins a row of yellow setae; anterolateral margins with 2-3 rows of yellow setae;
surface smooth, minutely punctate; lateral margin type C; base weakly recurved.

Elytra base ﬂat; scutellum type D; base never with a tumescence ﬁ'om seventh to
ninth interstriae; disc shallow rugosites occupying ﬁrst to fourth striae and interstriae just
beyond the scutellum, remaining elytral surface minutely granulate; metepistemum type
R, with setae palmately divided into four or more ﬁlaments; mesostemum type B; carina
originating at base of tenth interstria encompassing elytra; un- and diseriate rows of pale
yellow setae on interstriae; second visible abdominal sternite normal. Apex smooth.

Female similar to male except epistoma not expanded, less impressed and
unarmed; epistoma with a four rows of hairs on apical margin; frons ﬂat; nine rows of

setae lateral to medial line; punctures small, shallow, larger and deeper laterally; antennal

84

funicle segments 5-7 with scattered setae on less than length of three frmicular segments;
gena with triangular mycangium.

Etymology. This species is named after our colleague, Stephanie Dole for her gracious
mentoring and help throughout my graduate study.

Specimens examined. (36, 29)

Distribution. Ecuador (N apo).

Hosts. Unknown.

Biology. Unknown.

Camptocerus igniculus Smith & Cognato, sp. 11.
(Figs. 1.42, 1.43)

Type Material. Holotype, 6, BRAZIL: Amazonas: Hwy ZF 2, km 20.7, ca 60km N.
Manaus, 2°30’S, 60°15’W, ex. canopy fogging, l6.viii.l979, (Adis, Erwin, Montgomery
et al.) (NMNH); Parames, BRAZIL: 36, 132, Amazonas: Hwy ZF 2, km 20.7, ca 60 km
N. Manaus, 2°30’S, 60° 15’W, ex. canopy fogging, l6.viii.l979, (Adis, Erwin,
Montgomery et al.) (MSUC) 66, 12, (NMNH).
Diagnosis. Both sexes of C. igniculus have 1- 2 rows of golden scales on the elytral
interstrial spaces, a reddish-brown hue and a normal second visible abdominal sternite.

Its habitus is distinct ﬁom all other Camptocerus species.
Description. Holotype, Male 2.5 — 2.8 mm long (.7? = 2.7 m; n = 10); 1.7 — 2.0 times as

long as wide. Head, antennae, thorax, abdomen, elytra, and legs reddish- brown.
Apical epistomal margin armed by a short, transverse rectangular carina; lateral

epistomal margin armed with a straight carinae ornamented with cluster of 8-12 setae;

85

 

epistoma excavated; expanded to greater than quarter length of head; basal epistomal
margin tumid above insertion of scape ; frons strongly excavated ﬁom epistoma to
anterodorsal margin of the eyes with the excavation bordering the ocular margin, ﬁ'ons
shagreen; 2-3 golden setae lateral to medial line; setose lateral edges. Antennal scape
elongate and wider distally, four rows of setae half-length of scape on distal two-thirds;
frmicle segments 1-7 with setae equal to funicle length on ventral margin; setae on dorsal
margin less than length of four funicle segments; setae on ventral margin twice as thick
as those on dorsal margin; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes glabrous; pronotum with
2-3 rows of golden scales on anterior edge; surface minutely granulate- punctate;
punctures with ﬁne golden setae; lateral margin type C; base weakly recuved.

Elytral base margined by a weak carina; with shallow rugosites occupying ﬁrst to
fourth striae and interstriae just beyond the scutellum, remaining elytral surface minutely
granulate; strial punctures shallow with a ﬁne golden seta arising from the anterior
margin; interstrial golden scales mostly uniseriate but also di- seriate on disk; uniserial on
declivity; strial punctures 1.5- twice the size of interstriae. Carinae arising on the basal
margin of tenth interstriae extending around elytra; scutellum type B; apex of the
declivital margin of each elytron smooth. Metepistemum type C, with setae palmately
divided into four or more ﬁlaments; process of the mesostemum shape type B. Visible
abdominal stemites 2-4 each margined with a row of golden scales, sternite 5 with two

rows; second visible abdominal sternite inﬂated. Apex smooth.

86

Female similar to male in most features, except epistoma not expanded; frons ﬂat,
unarmed; antennal funicle segments scattered with setae less than length of three
funicular segments; gena mycangia oval shaped; second abdominal sternite normal.
Etymology. Igniculus (L) = little ﬂame, spark. Name for its “ﬁery” appearance (golden
setae and reddish-brown color) and in honor of a traditional Cognato family name
“Ignatius”.

Specimens examined. (106, 282)
Distribution. Brazil (Amazonas).
Hosts. Unknown.

Biology. Unknown.

Camptocerus latipilis Schedl
(Figs. 1.46, 1.47, 1.48)
Camptocerus latipilis Schedl, 1973:166. Holotype 6: Brazil: Para: Benﬁca: Ananindeua;
(MZUSP).
Diagnosis. The male is diagnosed from C. petrovi by its smaller size, black color; second
visible abdominal sternite strongly convex and protuberant, height of declivity behind
much less than length of sternite 3; dense brush of setae extending halfway on the lateral
margin from the epistoma to the level of scape insertion; ﬁontal excavation devoid of
setae; 10-12 setae on lateral and dorsal margins.
Female can be distinguished from C. coccoformus by the less impressed medial

frons.

87

Description. Male 2.7 - 3.3 mm long (3 = 3.2 mm; 11 = 6); 1.7- 1.9 times as long as wide.

Head, antennae, thorax, abdomen, elytra, and legs black.

Apical epistomal margin armed with a short rectangular carina; lateral margins of
epistoma with vertical costae; epistoma impressed; dense brush of setae extending
halfway on the lateral margin from the epistoma to the level of scape insertion; single
seta lateral to median line; basal epistomal margin tumid above scape insertion epistoma
expanded to greater than quarter length of head; male frons is strongly excavated, setae
absent from excavation; surface shagreen; 10-12 setae on lateral and dorsal margins.
Antennal scape elongate and wider distally, ﬁve rows of setae length of scape on distal
two-thirds; funicle segments 2-7 with setae equal to 1.5 times funicle length on ventral
margin; setae on dorsal margin less than length of ﬁve frrnicle segments; setae on ventral
margin twice as thick as those on dorsal margin; anterior face of the club setose with a
partial septum.

Anterior pronotal margin rounded, margin between the eyes glabrous; anterior
edge with 2-3 rows of yellow-brown setae; surface minutely granulate— punctate;
punctures minute, shallow; lateral margin type C; base wealdy recurved.

Scutellum type B; base never with a tumescence from seventh to ninth interstriae;
disc with shallow rugosites occupying ﬁrst to fourth striae and interstriae just beyond the
scutellum, remaining elytral surface minutely granulate; metepistemum type C, with
setae palmately divided into four or more ﬁlaments; mesostemum type B; carina
originating at base of tenth interstria encompassing elytra; uniseriate rows of yellow-
brown scales on interstria; second visible abdominal sternite strongly convex and

protuberant, height of declivity behind much less than length of sternite 3. Apex smooth.

88

Female similar to male except epistoma not expanded, less impressed and
unarmed; epistoma with a row of hairs on apical margin; ﬁ'ons ﬂat, with medial area of
ﬁ'ons above dorsal margin of eyes slightly impressed; antennal funicle segments 4-7 with
scattered setae less than length of three ﬁrnicular segments; gena with circular
mycangium; pronotum anterior margin between the eyes lined with scales; second visible
abdominal sternite ventrally convex and distally distended, noticeably smaller than male.
Specimens examined. (86, 5 9)

Type Material. Paratype Camptocerus latipilis, BRAZIL: Para: Belem, 18.x.1962, (J.
Bechyne), 6 (NHMW).

Other material. ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare
Camp, -0.652778,-76.433333, 220 m, ex. canopy fogging, x.1994, (T.L. Erwin et al.), lot
936, 182 (NMNH); vii.1996, lot 1546, 19 (NMNH); Tiputini Biodiversity Station, -
0.631944, -76.144167, 220-250 m, ex. canopy fogging, x.1995, (T.L. Erwin et al.), lot
1257, 16 (NMNH); x.1996, lot 1727, 16 (NMNH); vi.l998, lot 1895, 16, IS? (NMNH);
x.1998, lot 1930, 192 (NMNH); x.1998, lot 1947, 16 (NMNH); x.1998, lot 1953, 16
(NMNH); x.1998, lot 1987, 16 (NMNH); ii.l999, lot 2044, 16 (NMNH); ii.l998, lot
2068, 19 (NMNH).

Distribution. Brazil (Para), Ecuador (N apo).

Hosts. Unknown.

Biology. Unknown.

Camptocerus mallopterus Smith & Cognato, sp. 11.

(Figs. 1.51, 1.52)

89

 

Type Material. Holotype, 6, ECUADOR: Napo: Tiputini Biodiversity Station,-
0.631944, -76.144167, 220-250 m, ex. canopy fogging, ii.l999, (T.L. Erwin et al.), lot
2087, (NMNH held in trust for MECN). Paratypes, 192, ECUADOR: Napo: Tiputini
Biodiversity Station, -0.631944, -76.144167, 220-250 m, ex. canopy fogging, ii.l999,
(T.L. Erwin et al.), lot 2087 (NMNH), 16, 182, (MSUC).

Diagnosis. Both sexes of C. mallopterus are diagnosed by the distinctive light orange
head, pronotum and black abdomen and elytra, with the elytra densely covered in pale

recumbent setae.
Description. Male 3.8 — 3.9 mm long ()7 = 3.85 mm; 11 = 2); 1.8 — 1.9 times as long as

wide. Head, thorax, prolegs and coxae orange; antennae black and orange; meso and
metathoracic femora black, orange on distal ﬁfth; tibiae orange; abdomen and elytra,
black. Elytra striae and interstriae densely covered with pale recumbent setae.

Apical epistomal margin armed with a short rectangular carina; lateral margins of
epistoma with vertical costae; epistoma impressed; ﬁve rows of setae from lateral
epistoma margins to median line; basal epistomal margin tumid above scape insertion;
epistoma expanded to greater than quarter length of head; male frons is strongly
excavated, ornamented with eight rows of golden setae lateral to medial line; surface
shagreen; setose lateral edges; ﬁ'ons moderately covered by erect setae, surface
moderately covered. Antennal scape elongate and wider distally, dense setae length of
scape on distal three-quarters; ﬁmicle segments 2-7 with setae equal to length on ventral
margin; setae on dorsal margin less than length of ﬁve frmicle segments; setae on ventral
margin twice as thick as those on dorsal margin; anterior face of the club setose with a

partial septum.

90

Anterior pronotal margin rounded, margin between the eyes glabrous; anterior
edge with 4-5 rows of yellow-brown setae; surface densely covered with short, ﬁne hairs;
surface minutely granulate- punctate; punctures minute, shallow; lateral margin type C;
base weakly recurved.

Scutellum type B; base never with a tumescence ﬁom seventh to ninth interstriae;
disk with shallow rugosites occupying ﬁrst to fourth striae and interstriae just beyond the
scutellum, remaining elytral surface minutely granulate; metepistemum type C, with
setae palmately divided into four or more ﬁlaments; mesostemum type D; carina
originating at base of tenth interstria encompassing elytra; punctures not confused; striae
indistinguishable from interstriae; 3-4 rows of uniseriate pale yellow setae on interstria;
uniseriate on striae; setae equal size on both striae and interstriae; second visible sternite
weakly convex and strongly protuberant, height of declivity behind greater than the
length of sternite 3. Apex smooth.

- Female similar to male except epistoma not expanded; lateral two-thirds to
median line impressed; epistoma unarmed; epistoma with ﬁve transverse rows of setae;
frons ﬂat, with medial area of ﬁ'ons weakly impressed; ﬁ'ons with four setae lateral to
medial line; antennal ﬁrnicle segments 1-7 with scattered setae less than length of four
funicular segments; gena with circular mycangium; second visible abdominal sternite
ventrally convex and distally distended, noticeably smaller than male.

Etymology. Mallo- (G) wool, -pterus (G) = wing. In reference to the woolly appearance
due to the dense pale setae covering the elytra.
Specimens examined. (26, 282)

Distribution. Ecuador (N apo).

91

Hosts. Unknown.

Biology. Unknown.

Camptocerus petrovi Smith & Cognato, sp. 11.
(Figs. 1.65, 1.66)

Type Material. Holotype, 6, ECUADOR: Napo: Tiputini Biodiversity Station,-
0.631944, -76.144167, 220-250 m, ex. canopy fogging, ii.l999, (T.L. Erwin et al.), lot
1930, (NMNH held in trust for MECN). Paratype, 6, ECUADOR: Napo: Tiputini
Biodiversity Station, -0.631944, -76.144167, 220-250 m, ex. canopy fogging, ii.l999,
(T.L. Erwin et al.), lot 1930 (NMNH).
Diagnosis. The male is diagnosed by its large size, black color; strongly convex and
protuberant second visible sternite the height of declivity behind greater than length of
sternite 3; dense brush of setae extending halfway on the lateral margin from the
epistoma to the level of scape insertion and frontal excavation with 1-2 setae lateral to

median line, 15-20 setae on lateral and dorsal margins.
Description. Male 4.0 mm long (it = 4.0 mm; 11 = 2); 1.6 times as long as wide. Head,

antennae, thorax, abdomen, elytra, and legs black; apical margin of pronotum dark
brown. Teneral individuals with a red pronotum.

Apical epistomal margin armed with a short rectangular carina; lateral margins of
epistoma with vertical costae; epistoma impressed; dense bnrsh of setae extending
halfway on the lateral margin from the epistoma to the level of scape insertion and ﬁontal
excavation; single seta lateral to median line; basal epistomal margin tumid above scape

insertion; epistoma expanded to greater than quarter length of head; male frons is

92

strongly excavated, ﬁ'ontal excavation with 1-2 setae lateral to median line, 15-20 setae
on lateral and dorsal margins; surface shagreen. Antennal scape elongate and wider
distally, ﬁve rows of setae length of scape on distal two-thirds; funicle segments 3-7 with
setae equal to 1.5 times ﬁrnicle length on ventral margin; setae on dorsal margin less than
length of ﬁve funicle segments; setae on ventral margin twice as thick as those on dorsal
margin; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes lined with scales; lateral
margin type C; anterior edge with 2-3 rows of yellow-brown setae; surface minutely
granulate- punctate; punctures minute, shallow; lateral margin type C; base weakly
recurved.

Scutellum type B; base never with a tumescence from seventh to ninth interstriae;
disc with shallow rugosites occupying ﬁrst to fourth striae and interstriae just beyond the
scutellum, remaining elytral surface minutely granulate; metepistemum type C, with
setae palmately divided into four or more ﬁlaments; mesostemum type E; carina
originating at base of tenth interstria encompassing elytra; uniseriate rows of yellow-
brown yellow scales on interstria; strongly convex and protuberant second visible sternite
the height of declivity behind greater than length of sternite 3. Apex smooth.

Female unknown.

Etymology. This species is named in honor of our colleague, Alex Petrov who also
shares a love of Camptocerus and has generously contributed important material for this
study.

Specimens examined. (26, 09)

Distribution. Ecuador (N apo).

93

Hosts. Unknown.

Biology. Unknown.

“Costatus” Clade
Species in the “Costatus” clade are diagnosed by the pronotum lateral margin type
D, recurved pronotum base, scutellum shape type I and basal margin of elytra with a

weak subbasal carina extending to scutellar apex.

Camptocerus costatus Chapuis
(Figs. 1.33, 1.34, 1.35)

Camptocerus costatus Chapuis, 1869: 259. Type locality: ‘Brazil, Dejean’. Holotype 6,
(IRSNB).
Camptocerus seriatus Eggers 1933: 12. Type locality: French Guiana: Les roches de
Kourou; Holotype $2, (MNHN). Synonymy: Wood, 2007: 205.
Diagnosis. The male of C. costatus is diagnosed by the elytral declivity strongly
impressed below level of disk, occupying apical four-ﬁfths of elytra; carinate costa on
eighth interstriae ﬁ'om basal ﬁfth and extending around apex to sutural margin; striae
deeply impressed; bristling brown setae originating ﬁom recumbent asperities on the
interstriae.

The female is distinguished from C. piliﬁons by color, larger size, and more

impressed striae.

94

Description. Male 3.5 - 3.9 mm long (7: =37 mm; 11 = 6); 1.8 — 2.0 times as long as

wide. Head, thorax, abdomen, elytra, antennae and legs red-brown to black. Teneral
individuals are orange red.

Apical epistomal margin unarmed; lateral margin with a sub-acute arcuate costa
pointing proximally, lined entirely with setae; short, rectangular carina on basal epistomal
margin; epistoma excavated; expanded to greater than quarter length of head; male frons
strongly excavated and covered by long, dense, erect, dark yellow hairs; two rows on
lateral and caudal margin twice the length of other setae on the margins; surface
shagreen. Scape is elongate and broadly rounded distally, with a dense brush of setae as
long as scape on distal half; funicular segments 2-7 dense brushes of setae 2 times longer
than funicle on ventral margin; setae equal width on ventral and dorsal margin; setae on
dorsal margin of segments 1-7 less than four segments long; anterior face of the club
setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes glabrous; lateral margin
type D; pronotum base procurved; pronotum smooth, densely and minutely punctured,
medial line devoid of punctures, 5-6 setae transverse on anterior fourth of pronotum.

Elytra disk short, occupying basal ﬁfth of elytra; interstriae 1-4 rugose, remaining
disk smooth. Declivity strongly impressed below level of disk, occupying apical four-
ﬁfths of elytra; striae moderately impressed; interstriae on basal ﬁfth with recumbent
asperities above punctures, each puncture with a dark yellow, short bristle setae arising
from the posterior margin; asperities progressively become smaller in size forming
crenulations becoming small tubercles at apical ﬁfth and are obsolete at elytra apex;

scutellum type I; basal margin of elytra with a weak carina extending to scutellar apex;

95

base never with a tumescence ﬁom seventh to ninth interstriae; carina originating at base
of tenth interstria shorter than metepistemum; metepistemum shape type M;
metepistemum setae pahnately divided into more than four ﬁlaments; mesostemum shape
type B; declivity interstriae twice width of striae, setae uniseriate; apex of declivity
margin smooth.

Female similar to male except frons ﬂat and unarmed; shagreen; short, sparse,
setae present only on lateral margin of frons; setae present on epistoma; antennal funicle
segments 1-7 with setae on dorsal margin less than length of three ﬁrnicular segments;
gena with circular mycangium. Elytra ﬁnely sculptured, without indication of spines or
carinate costa; disk covered with shallow rugosities to interstria 8; disk covered with
shallow rugosities; striae weakly impressed; declivital interstriae with short, bristles of
equal length; declivital interstria 1-5 on apical half of declivity with a crenulation at the
base of each puncture with short bristle originating; setae of equal length.

Specimens examined. (66, 592)

Type Material. Holotype Camptocerus costatus, 6 (IRSNB). Holotype Camptocerus
seriatus, Q (MNHN).

Other material. BRAZIL: Mato Grosso: Villa Vera, 12°46’S 55°30’W, x.1970, (M.
Alvarenga), 16 (CAS). ECUADOR: Napo: Tiputini Biodiversity Station, -0.63l944, -
76.144167, 220-250 m, vi.l998, ex. canopy fogging (T.L. Erwin et al.), lot 1817, 16
(NMNH). GUYANA: [=Region 8]: Iwokrama Forest, 4.671822, -58.6841 14, 66 m, ex.
Ocotea oblonga, 4-9.iii.2007, (Cognato, Hulcr, Smith, Dole, McCall), 26, 432 (MSUC).

PERU: Loreto: 1.5km N Teniente Lopez, 2°35’S 76°6.92’W, 210-240 m, ex. ﬂight

96

intercept trap #166, 22.vii.1970, (Richard Leschen), 192 (SMEC). TRINIDAD: Morne
Bleu, 2700 ft, 21.viii.l969, (H & A Howden), 16 (NMNH).

Distribution. Brazil (Mato Grosso, Rio de Janeiro, Rondonia), Ecuador (Napo), French
Guiana, Guyana, Peru (Loreto), Trinidad.

Hosts. Ocotea oblonga (Meisn.) Mez.

Biology. This species creates galleries in 1 — 2 cm diameter branches.

Discussion. Identity of the female and synonymy of C. seriatus conﬁrmed when ﬁve

mated pairs of C. costatus were collected from galleries by Cognato et al. in 2007.

Camptocerus piliﬁpns Smith & Cognato, sp. 11.
(Figs. 1.67, 1.68)

Type Material. Holotype, 6, ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S
Onkone Gare Camp, -0.652778, -76.433333, 220 m, ex. canopy fogging, vii.1995, (T.L.
Erwin et al.) (NMNH held in trust for MECN). Paratypes, ECUADOR: l6, Napo:
Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778, -76.433333, 220 m,
ex. canopy fogging, vii.1994, (T.L. Erwin et al.) lot 714 (NMNH); 16, vii.1995, lot 1114
(NMNH); l6, viii.1995, lot 1115 (MSUC) 16 (NMNH); 16, 182, x.1995, lot 1254
(NMNH); 192, x.1995, lot 1256 (NMNH); 1S2, x.1995, lot 1260 (MSUC); l6, x.1996, lot
1669 (NMNH).
Diagnosis. Both sexes of C. pilifrons are diagnosed by its small size and distinctive color
pattern described below. The male is distinguished by an elytral disk with setae on
interstriae; the absence of a carinate costa on the eighth interstria and by the unimpressed

declivity and striae.

97

The female is distinguished from C. costatus by the color, smaller size, and

unimpressed striae.
Description. Male 2.6 — 3.1 mm long (i = 2.8 m; n = 7); 1.9 - 2.2 times as long as

wide. Head, thorax, abdomen, elytra, antennae and legs red-brown. Pronotum with a
black patch extending from the anterior margin of the pronotum and reaching a point to
the posterior third of the pronotum along the median line. An oval-shaped black patch is
present extending from the elytral base to the posterior margin of the ﬁrst visible sternite
and from the lateral margin to interstria 7.

Apical epistomal margin unarmed; lateral margin with an arcuate costa pointing
proximally, lined entirely with setae short rectangular carina on the basal epistomal
margin; epistoma excavated; expanded to less than quarter length of head; frons strongly
excavated; excavation borders ocular margin; densely covered with white bristles; lateral
two rows on frons margin slightly longer than other setae. Scape is elongate and broadly
rounded distally, with a dense brush of setae as long as scape on distal half; dorsal margin
of ﬁrnicular segments 2-7 with dense brushes of setae two times longer than funicle;
anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes glabrous; lateral margin
type D; pronotum base recurved; surface punctate, anterior and posterior margins more
densely punctured; medial line devoid of punctures; row of setae along lateral margins, 5-
6 setae transverse on anterior fourth of pronotum

Basal margin of elytra with a weak subbasal carina extending to scutellar apex;
scutellum shape type 1; base never with a tumescence from seventh to ninth interstriae;

carina originating at base of tenth interstria shorter than metepistemum; metepistemum

98

shape type F; metepistemum setae palmately divided into more than four ﬁlaments;
mesostemum shape type B; elytra smooth; declivity interstriae setae uniseriate, uniform
in size; interstriae twice width of striae; strial punctures twice size of interstriae; strial
punctures with a setae the length of puncture originating from anterior margin; apex of
declivity margin smooth.

Female similar to male except frons ﬂat and unarmed; short, sparse, setae present
only on lateral margin of ﬁons; ﬁons shagreen; setae present on epistoma; antennal
funicle segments 5-7 with setae on dorsal and ventral margins less than length of three
funicular segments; gena with circular mycangium. Elytra ﬁnely sculptured; disk covered
with shallow rugosities to interstria 8; disk covered with shallow rugosities; declivital
interstriae with short, bristles of equal length; declivital interstria 1-5 on anterior half of
declivity with a crenulation at the base of each puncture with short bristle originating;
setae of equal length.

Etymology. Pili- (L) = hairy, -frons (L) = forehead, brow. In reference to the setose
excavation of the ﬁons.

Specimens examined. (76, 3 9)

Distribution. Ecuador (N apo).

Hosts. Unknown.

Biology. Unknown.

Camptocerus quadridens Blackman

(Figs. 1.70, 1.71, 1.72, 1.73)

99

Camptocerus quadridens Blackman, 1943: 379. Holotype 6: Panama: Canal Zone:
Cooper’s, near source of Rio Aejeta l9.vii.1923, J. Zetek, fallen tree; (NMNH).
Diagnosis. The male is diagnosed by the short transverse carina on the basal epistomal
margin, each elytron with two discal spines arising from the ﬁrst to ﬁfth striae, with a
sharp spine on the ﬁrst striae and a blunt spine on the second extending over the
declivity, and the red-brown color.

The female is distinguished ﬁ'om C. costatus by the color, smaller size, and less

impressed striae.
Description. Male 3.4 — 3.7 mm long (i = 3.6 mm; 11 = 4); 1.8 — 1.9 times as long as

wide. Head, thorax, abdomen, elytra, antennae and legs orange to red brown.

Apical epistomal margin is unarmed; lateral margin with an arcuate costa pointing
proximally, entirely lined with setae; basal epistomal margin with a short, rectangular,
transverse carina; epistoma excavated; expanded to less than quarter length of head; male
frons strongly excavated and covered by long, dense, erect, pale yellow setae of equal
length; excavation borders ocular margin; scape is elongate and broadly rounded distally;
distal half of scape covered by hair-like setae the length of scape; ventral margin of
funicle segments 2-7 with brushes of setae 1.5 times the length of funicle; dorsal margin
of segments 1-7 with hair-like setae half the thickness of ventral margin and three-
quarters length of funicle; anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between eyes glabrous; lateral margin
type D; pronotum base recurved; pronotum smooth, densely and minutely punctured,

medial line devoid of punctures, 5-6 setae transverse on anterior fourth of pronotum.

100

Elytra disk with two spines arising from the ﬁrst to ﬁfth striae; a sharp spine on
the ﬁrst striae and a blunt rounded spine on the second, spines extend over the declivity,
declivity originates in the recess formed by the spines. The declivity under the margin of
the spines is moderately depressed until interstria 8, where a carinate costa originates the
middle of visible sternite 1 and terminates above the midpoint of visible sternite 5.
Scutellum shape type I; basal margin of elytra with a weak subbasal carina extending to
scutellar apex; base never with a tumescence from seventh to ninth interstriae; carina
originating at base of tenth interstria shorter than metepistemum; interstriae uniseriate,
twice width of striae; interstriae punctures indistinct; interstriae 1-4 with thin bristles 2-4
times the length of other bristles arising in the concavity under the spines to the posterior
third of declivity; apex smooth.

Female similar to male except ﬁons ﬂat and unarmed; short, sparse, setae present
only on lateral margins; frons shagreen; setae present on epistoma; antennal funicle
segments 5-7 with setae on dorsal margin less than length of three funicular segments;
gena with circular mycangium. Elytra ﬁnely sculptured, no indication of spines or
carinate costa; disk covered with shallow rugosities from sutural space to interstria 8;
declivital interstriae with short bristle of equal length; declivital intrerstria 1-5 on anterior
half of declivity with a crenulation at the base of each puncture with short, bristles; setae
of equal length.

. Specimens examined. (66, 192)
Type Material. Holotype Camptocerus quadridens, 6 (NMNH).
Other material. PANAMA: Canal Zone: Barro Colorado Island, ex. trap, l3.ii. 1980,

(Henk Wolda), 16 (UCDC); 21, 24, 26.vii.1986, 16 (MSUC); 21, 23, 25.vii.1986, 192,

101

(MSUC), 1-3.vii.l987, l6 (UCDC); Panama: Cerro Jefe, 19°12’N, 79°21 ’W, 24.ii.1973,
(H. Stockwell), 16 (NMNH).

Distribution. Panama.

Hosts. The holotype was collected on the same date and in the same locality as C.
aeneipennis. These C. aeneipennis were collected from Protium sp. and since multiple
species often colonize the same host, a Protium sp. is the probable host (Wood, 1982).
Biology. Unknown.

Discussion. This is the ﬁrst time that a female has been identiﬁed for this species. It was
found in a museum rather than collected in the ﬁeld, so its identity is hypothesized.
Camptocerus quadridens displays marked sexual dimorphism similar to that of its sister

taxa, C. costatus and C. piliﬁ'ons.

Camptocerus zucca Smith & Cognato, sp. 11.
(Figs. 1.80, 1.81)

Type Material. Holotype, 6, ECUADOR: Napo: Huahua Sumaco, Km 44 on Hollin-
Loreto Rd. 16.xii. 1989, (MS & JS Wasbauer, H. Real) (CSCA). Paratypes, ECUADOR:
132, Napo: Huahua Sumaco, Km 44 on Hollin-Loreto Rd. 16.xii.l989, (MS & JS
Wasbauer, H. Real) (CSCA); 16 (MSUC).
Diagnosis. This species is closely related to C. quadridens and C. costatus. It is readily
distinguished by its large size, unarmed apical epistomal margin, a distinctive orange and
black color pattern of the head, antenna, thorax, abdomen and elytra as described below,

absence of a carinate costa on the eighth interstriae and sparse setae on the declivity.

102

The female is distinguished from other females in the clade by the distinctive
color pattern, larger size, and unimpressed striae and the short, ﬁne setae on the
interstriae.

Description. Male 4.0 — 4.1 mm long ()7 = 4.05 mm; 11 = 2); 1.9 times as long as wide.
Antennal scape and club black, funicle orange, epistoma and frons orange, gena black,
protibia orange with black margins, profemur and meso- and metalegs black with orange
patches, coxae brownish-orange, pronotum orange with a black triangle extending from
the apex to the basal third of the pronotum, elytra black with each elytron displaying an
orange oval encompassing the area ﬁom the basal margin of the disk and to the basal ﬁfth
of the elytra and from the sutural interspace to the eighth interstriae; ventral surface of
thorax and abdomen black. Teneral adults are pale orange with dark spots.

Apical epistomal margin unarmed; lateral margin with an arcuate costa pointing
inward, entirely lined with setae; basal epistomal margin unarmed; epistoma excavated;
expanded to greater than quarter length of head; male frons strongly excavated and
covered by long, dense, erect, yellow-white hairs; excavation borders ocular margin;
male scape is elongate and broadly rounded distally; distal half with patch of setae length
of scape; ventral margin of funicle segments 2-7 lined with a brush of hair-like setae 1.5
times the length of funicle; dorsal margin of segments 1-7 lined with hair-like setae half
length of ventral margin, anterior face of the club setose with a partial septum.

Anterior pronotal margin rounded, margin between the eyes glabrous; lateral
margin type D; pronotum base recurved; pronotum smooth, sparsely covered by very

ﬁne, almost indistinguishable punctures.

103

Elytra disk with shallow rugosities to third interstria; remaining disk smooth,
striae not impressed; scutellum type I; basal margin of elytra with a weak subbasal carina
extending to scutellar apex; base never with a tumescence from seventh to ninth
interstriae; carina originating at base of tenth interstria shorter than metepistemum;
metepistemum shape type C; metepistemum setae biﬁd; mesostemum shape type 0;
declivity interstriae four times width of striae, setae uniseriate; setae sparse; setae on
declivital interstriae 1-5 two to three times as long as those on interstria 6; 2-3 long setae
present on each interstria; three lateral rows of short setae present on declivity margin
from visible sternite l to suture; apex of declivity margin smooth.

Female similar to male except ﬁons ﬂat and unarmed; short, sparse, setae present
only on lateral margin of frons; frons shagreen; setae present on epistoma; antennal
funicle segments 3-7 with setae on dorsal and ventral margins less than length of four
funicular segments; gena with circular mycangium. Elytra ﬁnely sculptured, no indication
of spines or carinate costa; disk covered with shallow rugosities to interstria 8; disk
covered with shallow rugosities; declivital interstriae with short, ﬁne pointed setae of
equal length.

Etymology. Zucca (Italian) = pumpkin (noun in apposition). In reference to this species
jack-o-lantem appearance.

Specimens examined. (26, 192)

Distribution. Ecuador (N apo).

Hosts. Unknown.

Biology. Unknown.

104

“Aeneipennis” Clade
Species in the “Aeneipennis” clade are distinguished by elongate and narrow male

scape; pronotum lateral margin type B; metepistemum type B; and mesostemum type A.

Camptocerus aeneipennis (F abricius)
(Figs. 1.19, 1.20)
Camptocerus aeneipennis (F abricius) 1801: 392 (Hylesinus). Lectotype 6: Essequibo
[River] [=Guyana], (UZMC). Lectotype designated Wood, 1982: 412
Camptocerus gibbus (Fabricius) 1801: 392 (Hylesinus). Type locality: ‘Essequibo’
[River] [=Guyana]; Lectotype SB (here designated), (UZMC). Synonymy: Eggers 1933:
1 7.
Notes. Synonymy of C. gibbus conﬁrmed by Wood (1972: 243).
Diagnosis. This species always has an elytra colored metallic bronze or black with a
metallic sheen and is smaller in size than its sister taxa, C. noel. The male can and is
distinguished by the impressed ﬁrst discal striae, deeper strial punctures and a moderate
pronotum sulcus, encompassing lateral two-thirds of anterior margin.
Females are identiﬁed from C. noel by their smaller size and deeper strial
punctures.

Description. Male 4.4 — 5.9 mm long (J? = 5.5 mm; 11 = 20); 1.8 — 2.1 times as long as

wide. Head, thorax, antennae and legs red-brown; pronotum dark red-brown; abdomen
black; elytra metallic bronze.
Apical epistomal margin is unarmed; lateral margin of epistoma with a sub-acute

arcuate costa pointing distally; epistoma strongly excavated, expanded to greater than

105

quarter length of head; basal epistomal margin armed with a transverse, concave carina;
frons strongly excavated and densely covered with long white to yellow hair-like setae;
margin of excavation lined with a single row of long hairs; male scape elongate and
narrow; glabrous on proximal quarter, setae on ventral half, one third length of scape;
frons and epistoma smooth, shining; funicle segments 2-7 with brushes of setae equal of
length of scape; dorsal margin with setae less than three funicle segments; anterior face of
the club setose with a partial septum.

Lateral declivities on anterior margin of pronotum extending two-thirds of
distance to midline; margin between eyes lined with ﬁne biﬁd setae; pronotum glabrous,
smooth, shining; minutely punctate; lateral margin type B; basal half deeply rugose
creating a wrinkled appearance; base straight.

Elytra glabrous; the ﬁrst discal striae impressed; interstriae punctures strongly
confused; elytra smooth; base never with a tumescence from seventh to ninth interstriae;
scutellum type A; carina at base of interstria ten shorter than metepistemum;
metepistemum with biﬁd setae; metepistemum type B; mesostemum type A; apex
smooth.

Female similar to male except epistoma with a small median tubercle; ﬁ'ons ﬂat
with a triangular impressed area originating between the eyes and terminating at the distal
and lateral epistoma margins, the triangle is bisected by a shallow carina on the median
line; small tubercle at dorsal end; frons ornamented with few short setae; ﬁ'ons shagreen,
dorsal and lateral margin reticulate, covered with moderate, deep punctures; antennal

funicle segments 5-7 with setae on dorsal margins less than length of two funicular

106

segments; gena with crescent shaped mycangium. Pronotum anterior margin weakly
sulcate, margin between the eyes lined with scales; pronotum, smooth, shining.
Specimens examined. (1636, 1092)

Type Material. Lectotype, 6, Camptocerus aeneipennis (MZMC); Paralectotypes,
Camptocerus aeneipennis, NO LOCALITY, 26, 4Q, (MZMC); Lectotype,
Camptocerus gibbus, Q (MZMC).

Other specimens: BOLIVIA: Bent: Cosincho region, (G.L. Harrington), l6 (NMNH);
Romansos, 1 km N. Junction Rio Itenez & Rio Paragua, 30.Vii.1964, (J.K. Bousemen, J.
Lussenhop), 16, 132 (AMNH); La Paz: Chuma [dubious locality due to high elevation
and biome]: xii.1936 (Y. Mexia), 16 (CAS). BRAZIL: Amazonas: km 60 N~Manaus,
18.1 km E Campinas ﬁeld station, 2° 30’S, 60°15’W, ex. terra ﬁrme forest fogged with
pyrethrum (Montgomery, Erwin, Schimmel, Krischik, Date, Bacon), sample 6, 22.ii.1979,
16 (NMNH); Manaus, 1 km W Taruma Falls, 100m, 12-17.ii.1981, (Ekis, Young), IS?
(RJRC); Bahia: (G. Bondar), 46, 12 (FMNH); Salobro: 6.vii.1885, (E. Gounelle), 26
(DEI); Parana: Rolaudia, x.1947, (A. Maller), l6 (AMNH); Pemambuco: Pery-Pery,
5.vi.1892, (Gounelle), 36, IS? (DEI); 11.xii.1892, 16 (DEI); Rio de Janeiro: (Kirsch)
l6, (SMTD); Santa Catharina: Corupa (Hansa Humbolt), xii.1945, (A. Maller), 16
(AMNH); ‘Sao Paulo ’ (Kratz), 16 (DEI); No speciﬁc locality: ‘Brazil’ (Mearkel), 192
(SMTD); ‘Brazilien’, 16 (MZMC); ‘Rio Madeira’ [river runs through states of Rondonia
and Amazonas], 1954, (Mann, Baker), 26 (NMNH). COLOMBIA: Cauca: 10 km SE
Tambo, 30 m, ex. ‘Anime’ [local name of Protium], 9.vii.1970, (S.L. Wood), 382
(MSUC); Choco: Quebrada Docordo, Rio San Juan, (B. Malkin), 4—8.vi.1969, 16

(FMNH); Putumayo: Santa Rosa (Kofan Indian village), headwaters of Rio San Miguel,

107

ex. at light, (B. Malkin, P. Buchard), 9-12.x.1970, l6 (FMNH); Santa Fe de Bogotd
[=Bogotd D. C.], 46 (DEI). COSTA RICA: Peralta: Cartago, ex. 4” limb, 10.iii.1964,
(S.L. Wood), 66, 4Q (MSUC); Santa Clara: Hamburg Farm, ex. on dry bark, (F.
Nevermann), 16 (NMNH). ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S
Onkone Gare Camp, -0.652778, -76.433333, 220 m, ex. canopy fogging, i.1994, lot 614,
19 (NMNH); vii i.1995, lot 1028, 16 (NMNH); vii.1995, lot 1110, 16 (NMNH); i.1996,
(T.L. Erwin et al.), lot 1479, 16, (NMNH); i.1996, lot 1498, 56, 3S? (NMNH); vii.1996,
lot 1530, 16‘ (NMNH); vii.1996, lot 1618, 12 (NMNH); Tiputini Biodiversity Station, -
0.631944, -76.144167, 220-250 m, ii.l999, ex. canopy fogging (T.L. Erwin et al.), lot
2056, 16 (NMNH); ii.l999, lot 2075, 16 (NMNH). FRENCH GUIANA: No speciﬁc
locality: ‘Cayenne’ 26 (DEI); 26 (MZMC). GUYANA: [=Region 8]: Iwokrama Forest,
4.671822, -58.684114, 66m, ex. Ocotea oblonga, (Cognato, Hulcr, Smith, Dole, McCall),
4-9.iii.2007, 686, 4252 (MSUC). PANAMA: Bocas del T oro: Corriente Grande, 100 m,
9°17’30”N, 82°32’41”W, 30.iv-5.v.l980, (H. Wolda), 1S2; Canal Zone: Barro Colorado
Island, ex. trap, 9-13.xi.1987, (H. Wolda), 3S? (UCDC); 24.vi.l986, (H. Wolda), 192
(UCDC); Cooper’s nr. source of Rio Ajeta [= along Pipeline Road], l9.viii.l928, (J.
Zetek), 16, 29 (NMNH); Panama, Parque Nacional Soberania, Pipeline Rd, 9.166667, -
79.75, 95 m, 3.ix.2008, (SM. Smith, A.D. Smith, A.R. Gillogly), 16 (MSUC). PERU:
Madre de Dios: Los Amigos Biological Station, -12.569l6, -70.100114, 250 m, ex.
Protium amazonicum (Smith, Hulcr) 26.iv.-27.v.2008, 16, 2Q (MSUC). SURINAME:
No speciﬁc locality: ‘Surinam’ 26 (MZMC). VENEZUELA: Barinas: 9 km S
Barrancas, ex. Protium sp., 6.xii.1970, (S.L. Wood), 256 289 (MSUC); 40 km E Canton,

100 m, ex. Protium sp. 11.iii.1970, (S.L. Wood), 26, 49 (MSUC); 10 km SE Miri, 150

108

 

m, ex. Protium sp., 9.ii.l970, (S.L. Wood), 46, 29 (MSUC). N0 SPECIFIC
LOCALITY: ‘Amazons’, 16 (MZMC); ‘Cujam’[?] 76, 29 (DEI).

Distribution. Argentina (Chaco), Bolivia (Beni, La Paz), Brazil (Amazonas, Bahia,
Brasilia, Mato Grosso, Para, Parana, Pemambuco, Rio de Janeiro, Santa Catharina, Sﬁo
Paulo), Colombia (Bogota, Cauca, Choco, Putumayo), Costa Rica, Ecuador (N apo),
French Guiana, Guyana, Panama, Peru (Loreto, Madre de Dios), Suriname, Venezuela
(Barinas).

Hosts. Euphorbiaceae, Ocotea oblonga (Meisn.) Mez, Myrocarpusﬁ'ondosus Fr. Allem,
Myrtaceae, Protium sp. (“Almesca branca” [Brazil], “Almesca vermelha” [Brazil], “Breu
branco” [Brazil] (Beaver, 1972)), Protium amazonicum Swart, Protium decandrum
(Aubl.) Marchand, Protium glabrum (Rose) Engl. Protium sp., Protium tenufolium
(Engl.) Engl., T etragastris altissima (Aubl.) Swart.

Biology. C. aeneipennis colonize large branches ranging ﬁ'om 5 cm diameter to more
than 30 cm. Wood (2007) offers a detailed description of the gallery system. Beaver
(1972) provides a detailed life history. The galleries closely resemble those of C. noel. C.
aeneipennis aggregate in leaves around host trees and perform maturation feeding on
leaves (von Winning, 1929; chapter 2).

Discussion. Camptocerus aeneipennis from Costa Rica and Bolivia exhibited slight
variation in the degree of separation was observed in the sensory claspers of the

aedeagus.

Camptocerus angustior Eggers

(Figs. 1.21, 1.22)

109

Camptocerus angustior Eggers, 1928: 91. Lectotype, 6, BOLIVIA: Yungas: 1000 m
(NMNH). Lectotype designated by Anderson & Anderson (1971: 4).
Diagnosis. This species has metallic green elytra. The male of the species is diagnosed by
a median tubercle on the apical epistomal margin, constricted proximal epistomal margin,
nearly sulcate anterior pronotum margin, less rugose basal half of prontoum, larger,
deeper punctures on the pronotum and by shallow strial punctures.

The female is diagnosed from C. aeneipennis by the unimpressed striae and
cannot be distinguished from C. noel. This species occurs at higher elevations (1200-

3000m) as compared to C. aeneipennis or C. noel.
Description. Male 5.0 - 5.8 mm long ()7: = 5.5 mm; 11 = 4); 1.9 — 2.0 times as long as

wide. Head, antennae, thorax, abdomen and legs black; elytra metallic bronze to black.

Apical epistomal margin armed with a median tubercle; lateral margin of
epistoma with a sub-acute arcuate costa pointing distally; epistoma strongly excavated,
expanded to greater than quarter length of head; basal epistomal margin armed with a
transverse, concave carina; proximal margin distinctly constricted; ﬁ'ons strongly
excavated and densely covered with long white to yellow hair-like setae; margin of
excavation lined with a single row of long hairs; frons and epistoma smooth, shining;
male scape elongate and narrow; glabrous on proximal quarter, setae on ventral half, one
third length of scape; funicle segments 2-7 with brushes of setae equal of length of scape;
dorsal margin with setae less than three funicle segments; anterior face of the club setose
with a partial septum.

Pronotum anterior margin declivious, margin between eyes lined with ﬁne biﬁd

setae; pronotum glabrous, smooth, shining; minutely punctate; lateral margin type B;

110

basal half deeply rugose creating a wrinkled appearance; transverse medial groove on
base; base straight.

Elytra glabrous; minutely punctate; elytra smooth; striae not impressed; interstriae
punctures strongly confused; base never with a tumescence from seventh to ninth
interstriae; scutellum type A; carina at base of interstria ten shorter than metepistemum;
metepistemum with biﬁd setae; metepistemum type B; mesostemum type A; apex
smooth.

Female specimen missing majority of ﬁ'ons. Female similar to male except
epistoma with a small median tubercle; antennal funicle segments 4-7 with setae on
dorsal margin less than length of two funicular segments; gena with crescent shaped
mycangium. Pronotum anterior margin weakly sulcate; margin between the eyes lined
with scales; pronotum smooth, shining; transverse medial groove on base absent.
Specimens examined. (56, 19)

Type material. Lectotype, Camptocerus angustior, 6 (NMNH); Paratypes, BOLIVIA:
No speciﬁc locality: ‘Bolivia’, 16 (BMNH) 19 (NMNH); COLOMBIA: T olima:
(Natagaima), E. Pehlke S., 1915, (M. Hagedom), 16 (NMNH).

Other material. PERU: Junin: Utcuyacu, Tarms, iii. 1948, (F. Woytkowski), 16
(AMNH); 1600—3000 m, 12.iii.1948, (F. Woytkowski), 16 (AMNH).

Discussion. Female is badly damaged and is missing most of head. Camptocerus
pseudoangustior is commonly identiﬁed as C. angustior in collections and keys (Wood
2007)

Distribution. Bolivia (La Paz), Colombia (Tolima), Peru (Junin).

Hosts. Unknown.

111

Biology. Unknown.

Camptocerus aterrimus Eggers
(Figs. 1.25, 1.26)

Camptocerus aterrimus Eggers, 1933: 12. Holotype, 9, French Guiana: Passoura
[=River] (MNHN).
Diagnosis. This species is black with dark brown gena and legs. The species is
distinguished by its black color; median tubercle on the apical epistomal margin; anterior
pronotum margin weakly sulcate (nearly round); groove on pronotum base, and mostly
uniseriate discal interstrial punctures.

Females are distinguished from C. pseudoangustior by the smaller size and by the

larger, deeper punctures on declivital striae.
Description. Male 3.1 — 4.5 mm long ()7 = 3.9 mm; 11 = 20); 1.7 - 2.2 times as long as

wide. Head, antennae, thorax, legs, abdomen and elytra and legs black; gena and coxa red
brown.

Apical epistomal margin with a median tubercle; lateral margin of epistoma sub-
acute arcuate costa pointing distally; epistoma strongly excavated, expanded to less than
quarter length of head; basal epistomal margin armed with a transverse, concave carina;
tuft of thick apically rounded setae above scape insertion; ﬁons strongly excavated and
densely covered with long white hair-like setae; margin of excavation lined with a single
row of long hairs; hairs near ocular margin with their apex ﬂattened; male scape elongate
and narrow; glabrous on proximal quarter, setae on ventral half, one third length of scape;

frons and epistoma smooth, shining; funicle segments 2-7 with brushes of setae equal of

112

length of scape; dorsal margin with setae less than three ﬁmicle segments; anterior face of
the club setose with a partial septum.

Anterior pronotal declivity weakly sulcate (nearly round), margin between eyes
lined with ﬁne biﬁd setae; pronotum glabrous, smooth, shining; minutely punctate; lateral
margin type B; basal half deeply rugose creating a wrinkled appearance; transverse
medial groove on base; base straight.

Elytra glabrous; interstriae punctures strongly confused on disk; elytra smooth;
striae not impressed; base never with a tumescence from seventh to ninth interstriae;
scutellum type A; carina at base of interstria ten longer than metepistemum;
metepistemum with biﬁd setae; metepistemum type B; mesostemum type A; apex
smooth.

Female similar to male except epistoma with a small median tubercle; from ﬂat
with a triangular impressed area originating between the eyes and terminating at the distal
and lateral epistoma margins, the triangle is bisected by a shallow carina on the median
line; frons ornamented with few short setae; frons shagreen, covered with small, deep
punctures; dorsal and lateral margins reticulate; antennal funicle segments 4-7 with setae
on dorsal margin less than length of two funicular segments; gena with crescent shaped
mycangium. Pronotum anterior margin between the eyes lined with scales; pronotum
smooth, shining; transverse medial groove on base absent.

Specimens examined. (326, 939)

Type Material. FRENCH GUIANA: Passoura [River], vi.l907, (E. Le Moult), 9

(MNHN).

113

Other material. BRAZIL: Amazonas: 69 km N Manaus, 7.xii. 1979, (G. Stevens), 86, 79
(NMNH); Manaus, ‘Corcovado, N.P.’ [error?- this park is in Costa Rica], 26.xi.1979, (G.
Stevens), 19, (NMNH); Mata Grassa: 12° 31’S, 51° 46’W, R.S. & R.G.S. Exped, 12-
18.xi.l968, ex. dry forest, (0. W. Richards) 16 (BMNH; [Mata Grasso]: 12° 31 ’S, 51°
46’W, RS-RGS, 12-18.xi.l968, (R.A. Beaver), 66, 89 (NMNH); Randénia: BR: 364, km
48, 27.ix.l979, armadilha de malaise [=malaise trap], (Cecli Blancardi), 16 (NMNH).
ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778, -

76.433333, 220 m, ex. canopy fogging, vii.1994, (T.L. Erwin et al.), lot 744, 26, 19,

(NMNH); x.1994, lot 936, 19, (NMNH); i.1995, lot 961, 16‘, (NMNH); x.1995, lot 1265,

36, 49 (NMNH); x.1995, lot 1495, 16 (NMNH); x.1996, lot 1755, 16, 29 (NMNH).
GUYANA: [=Region 8]: Iwokrama Forest, Pakalau hills, 4.748333, -59.026667, 70 m,

ex. beating treefall litter, 27.v.2001, (R. Brooks, Z. Falin), 16, (SMEC); ex. on bark,
downed tree, 27.v.2001, (R. Brooks, Z. Falin), l6, (SMEC). PERU: Madre de Dios: Los
Amigos Biological Station, -12.569l6, -70.100114, 250 m, 10-16.v.2008, (Smith, Hulcr)
26, 459 (MSUC); l7-25.v.2008, (Smith, Hulcr) 36, 249 (MSUC); Tampopata, 15 km
NE Puerto Maldonado, Maldonado Reserva Cuzco Amazonico, Quebrada Mariposa, -
12.55, -69.55, 200 m, ﬂight intercept trap, 13-15.vi.l989, (A. Leschen), l6, (SMEC); No
speciﬁc locality: ‘Ibaria’ (?), Rio Pachitea, 120 m, 10.xii.l967, (R. Garcia), 16,

(MU SM).

Distribution. Brazil (Amazonas, Mato Grosso, Para, Rondonia), Ecuador (N apo), French
Guiana, Guyana, Peru (Madre de Dios).

Hosts. Protium amazonicum, Protium sp. (“Almesca branca” [Brazil], “Almesca

vermelha” [Brazil], “Breu branco” [Brazil]) (Beaver, 1972).

114

Biology. C. aterrimus infest 2 - 5 cm diameter branches and create a vertical maternal
gallery extending above and below the entrance tunnel in the pith‘of small branches and
twigs. Two short radial maternal galleries may also be created in larger stem. Females lay
single eggs along the maternal gallery in wide niches cut at right angles to the gallery and
plugged with boring dust. Larvae enlarge their cradles parallel to the maternal gallery
and parallel to the grain of wood. Beaver (1972) provides a detailed description of the life

history.

Camptocerus distinctus Smith & Cognato, sp. n.
(Figs. 1.36, 1.37)

Type Material. Holotype, 6, ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S
Onkone Gare Camp, -0.652778, -76.433333, 220 m, ex. canopy fogging, i.1994, (T.L.
Erwin et al.), lot 613 (NMNH held in trust for MECN). Paratypes, ECUADOR: 29,
Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778, -76.433333,
220 m, ex. canopy fogging, i.1994, (T.L. Erwin et al.), lot 613 (NMNH) 16, 19 (MSUC).
Diagnosis. This species is solid black. The male of the species is diagnosed by the
shining, black elytra, each elytron with eight rugae on the discal sutural interspace to
second interstria; each rugae with a single setae arising from the center; large median
tubercle on the apical epistomal margin, the vertical carinae on the lateral margins of the
epistoma; basal epistomal margin tumid above scape insertion; erect frons setae, and the
absence of a groove on the basal margin of pronotum.

The female is diagnosed by the eight rugae on the discal sutural interspace of each

elytron, each rugae with a single setae arising from the center and reticulate ﬁ'ons.

115

Description. Male 5.3 mm long (J? = 5.3 mm; 11 = 2); 1.9 — 2.1 times as long as wide.

The head, antennae, thorax, legs abdomen and elytra are shining black.

Apical epistomal margin with a median tubercle; lateral margin of epistoma with
a vertical sub-acute costa; epistoma strongly excavated; expanded to greater than quarter
length of head; epistoma covered with a few short setae; basal epistomal margin tumid
above scape insertion; frons strongly excavated and moderately covered with erect, ﬁne,
long pale hair-like setae; excavation borders the ocular margin; margins of excavation
rounded and lined with a single row of long hairs; frons and epistoma reticulate; male
scape elongate and expanded distally; glabrous on proximal quarter; setae on ventral half,
one third length of scape; funicle segments 2-7 with brushes of setae equal length of
scape; dorsal margin with setae less than three funicle segments; anterior face of the club
setose with a partial septum.

Anterior pronotal margin rounded; margin between the eyes glabrous; surface
smooth, shining, glabrous; ﬁnely punctate; lateral margin type B; base recurved.

Elytra glabrous; eight rugae on the discal sutural interspace to second interstria of
each elytron, each rugae with a single setae arising from the center, striae not impressed,
punctures small, minute; interstriae punctures strongly confused, minute; base never with
a tumescence from seventh to ninth interstriae; scutellum type H; carina at base of
interstria ten shorter than metepistemum; metepistemum with biﬁd setae; metepistemum
type B; mesostemum type A; apex smooth.

Female similar to male except epistoma with a small median tubercle; epistoma
moderately covered with setae; ﬁ'ons ﬂat with a triangular impressed area originating

near the vertex and terminating at the distal and lateral epistoma margins, the triangle is

116

bisected by a shallow carina on the median line; frons ornamented with few short setae;
basal epistomal margin tumid above scape insertion; frons sparsely covered with short,
ﬁne setae; ﬁ'ons reticulate, covered with moderate, deep punctures; antennal funicle
segments 5-7 with setae on ventral and dorsal margins less than the length of two
funicular segments; gena with crescent shaped mycangium. Rugae and setae on sutural
interspace smaller, nearly indistinct.

Etymology. Distinctus (L) = distinct. This species possesses numerous distinct
apomorphies and is very different ﬁ'orn other Camptocerus species.

Specimens examined. (26, 49)

Distribution. Ecuador (N apo).

Hosts. Unknown.

Biology. Unknown.

Camptocerus hirtipennis Schedl status 11.
(Figs. 1.40, 1.41)

Camptocerus hirtipennis Schedl, 1973:165. Holotype, 6, BRAZIL: Amazonas: Tefe;
(MZUSP).
Diagnosis. This species has a dark brown pronotum and black elytra. The elytra is
densely covered with short, recumbent golden hairs. This species is closely related to C.
suturalis and the male is distinguished by the larger size, solid black elytra, arcuate costa
pointing out on the lateral margin of the epistoma, partially sulcate anterior pronotum

margin, rugose basal medial area of pronotum.

117

The female is diagnosed from C. suturalis by the solid black elytra and vestiture
as mentioned above, scales on the pronotum anterior margin, groove mycangium on gena

and enlarged punctures on vertex.
Description. Male 4.9 — 5.6 mm long (3 = 5.0 mm; 11 = 9); 1.9 — 2.2 times as long as

wide. Head, antennae, thorax, legs, abdomen and elytra and legs dark black or brown.
The body, except pronotum disc, is entirely covered by dense recumbent, hair-like golden
setae. Setae are often abraded.

Apical epistomal margin unarmed; lateral margin of epistoma with a sub-acute
arcuate costa pointing distally; epistoma strongly excavated; expanded to greater than
quarter length of head; epistoma covered with a few short setae; basal epistomal margin
armed with a glabrous, transverse, concave carina; frons strongly excavated and densely
covered with long golden yellow hair-like setae; margin of excavation lined with a single
row of long hairs; hens and epistoma shagreen; male scape elongate and narrow;
glabrous on proximal quarter, setae on ventral half, one third length of scape; funicle
segments 2-7 with brushes of setae 1.5 times the length of scape; dorsal margin with setae
less than three funicle segments; anterior face of the club setose with a partial septum.

Anterior pronotal declivity partially sulcate, margin between eyes lined with ﬁne
biﬁd setae; pronotum glabrous, minutely punctate; lateral margin type B; long yellow
setae on anterior ﬁfth, vestitrrre emarginate at median line; setae diagonally extend along
the lateral margin from the anterolateral margin; basal margin with a row of short setae,
setae twice as long on media] third; basal half deeply rugose creating a wrinkled

appearance; base straight.

118

Elytra densely covered with long recumbent golden hairs on both striae and
interstriae; striae not impressed, punctures wide, deep; interstriae punctures strongly
confused, minute half as deep at striae; interstriae with 24 rows of setae on disc
interstriae; base never with a tumescencet from seventh to ninth interstriae; scutellum
type A; carina at base of interstria ten shorter than metepistemum; metepistemum with
biﬁd setae; metepistemum type B; mesostemum type A; apex smooth.

Female similar to male except epistoma with a small median tubercle; frons ﬂat;
with a triangular impressed area originating between the eyes at the ventral end of a weak
impression and terminating at the distal and lateral epistoma margins, the triangle is
bisected by a shallow carina on the median line; frons ornamented with short, moderately
dense setae; ﬁons shagreen, covered with moderate, deep punctures; antennal funicle
segments 5-7 with setae on dorsal margin less than length of two funicular segments;
gena with crescent shaped mycangium. Anterior pronotal margin rounded, margin
between eyes lined with scales; pronotum smooth, shining.

Specimens examined. (86, 13 9)

Type material. Allotype, BRAZIL: Amazonas: Tapurucuara am Rio Negro, 29.xi.1962,
(C. Lindemann), 9 (NHMW).

Other material. BRAZIL: Amazonas: Hwy ZF 2, km 20.7, ca 60km N. Manaus, 2°30’S,
60°15’W, l6.viii.l979, (Adis, Erwin, Montgomery et al.), ex. canopy fogging l9
(NMNH); 69 km N Manaus, 7.xii.1979, (G. Stevens), 76, 79 (NMNH). ECUADOR:
Napo: Tiputini Biodiversity Station, -0.63l944, -76.144167, 220-250 m, ex. canopy

fogging, vi.l998, (T.L. Erwin et al.), lot 1853, 192 (NMNH); vi.l998, lot 1867, 192

119

(NMNH); vi.1998, lot 1892, 16 (NMNH); x.1998, lot 1955, 19 (NMNH); x.1998, lot
1988 19 (NMNH).

Discussion. This species was previously considered a synonym of Camptocerus suturalis
by Wood (1992). Holotype comparison indicated the resurrection of C. hirtipennis based
on the solid black color of the elytra, the presence of mycangia on the gena, dense
vestiture restricted to the anterior margin of the pronotum, the larger size, and by male
genitalia.

Distribution. Brazil (Amazonas), Ecuador (N apo).

Hosts. Unknown.

Biology. Unknown.

Camptocerus mandelshtami Petrov
(Figs. 1.53, 1.54)

Camptocerus mandelshtami Petrov, 2007:101. Holotype 6: Peru: Loreto: 68km SW from
Iquitos to Itaya river, 4°11’S 73°26’W, 120 m, A. Petrov (ZMMU).
Diagnosis. This species is distinguished by a bicolored pronotum (posterior ﬁfth - two
ﬁfths orange, anterior black) and other color pattern described below. Elytra glabrous.
Males of this species are distinguished a median tubercle on the apical epistoma margin,
strongly rugose basal medial region of pronotum, prontoum anterior margin sulcate, and
medial groove on pronotum base.

Females are distinguished by the color pattern and by a black frons with an orange

area from the vertex to epistoma.

120

Description. Male 5.9 — 6.5 mm long (3? = 6.3 mm; 11 = 6); 1.8 — 2.0 times as long as

wide. Head and antennae, dark red brown; pronotum bicolored (anterior ﬁfth to two-
ﬁfth’s orange remainder black), profemur and protibia red orange, remaining legs black;
thorax, abdomen and elytra black.

Apical epistomal margin with a median tubercle; lateral margin of epistoma with
a sub-acute arcuate costa pointing distally; epistoma strongly excavated, expanded to less
than quarter length of head; basal epistomal margin armed with a transverse, concave
carina; tuft of thick apically rounded setae above scape insertion; frons strongly
excavated and densely covered with long gold hair-like setae; margin of excavation lined
with a single row of long hairs; hairs near ocular margin with their apex ﬂattened; male
scape elongate and narrow; glabrous on proximal quarter, setae on ventral half, one third
length of scape; frons and epistoma smooth, shining; funicle segments 2-7 with brushes
of setae equal of length of scape; dorsal margin with setae less than three funicle
segments; anterior face of the club setose with a partial septum.

Anterior pronotal declivity entirely sulcate, margin between eyes lined with ﬁne
biﬁd setae; pronotum glabrous; smooth, shining; minutely punctate; lateral margin type
B; basal three-quarters deeply rugose creating a wrinkled appearance; transverse medial
groove on base; base straight.

Elytra glabrous, shining; interstriae punctures strongly confused on disk;
punctures shallow, minute; elytra smooth; striae not impressed; base never with a
tumescence from seventh to ninth interstriae; scutellum type A; carina at base of interstria
ten shorter than metepistemum; metepistemum with biﬁd setae; metepistemum type B;

mesostemum type A; apex smooth.

121

Female similar to male except epistoma with a small median tubercle; frons ﬂat;
with a triangular impressed area originating between the eyes and terminating at the distal
and lateral epistoma margins, the triangle is bisected by a shallow carina on the median
line; medial impressed triangular region orange; frons sparsely covered with short setae;
frons shagreen, covered with large, deep punctures; antennal funicle segments 1-7 with
setae on dorsal and ventral margins less than length of two funicular segments; gena with
crescent shaped mycangium. Anterior pronotal declivity partially sulcate, margin
between the eyes lined with scales; pronotum smooth, shining; transverse medial groove
on base absent.

Specimens examined. (116, 13 9)

Type Material. Unable to examine type material.

Other material. ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare
Camp, -0.652778,-76.433333, 220 m, ex. canopy fogging, x.1994, (T.L. Erwin et al.), lot
952, 36 (NMNH); vii.1995, lot 1082, 26, 49 (NMNH); x.1995, lot 1222, 19 (NWH);
x.1995, lot 1254, 16 (NMNH); vii.1996, lot 1604, 16, 19 (NMNH); x.1996, lot 1712,
29 (NMNH); vi.l998, lot 1856, 29 (NMNH). PERU: Loreto: 7010n SSW from Iquitos
to Nauta, left bank of Amazon River, 140 m, 23.ii.2008, (A. Petrov), 36, 29 (MSUC);
29.ii.2008, 16, 19 (MSUC).

Distribution. Ecuador (N apo), French Guiana, Peru (Loreto).

Hosts. Protium spp.

Biology. Petrov (2007) provides a detailed account of the gallery system, which appears

to resemble that of C. aeneipennis.

122

Camptocerus noel Smith & Cognato, sp. 11.
(Figs. 1.57, 1.58)

Type Material. Holotype, 6, PERU: Loreto: 1.5km N Teniente Lopez, 2°35’S
76°6.92’W, 21.0-240m, ex. ﬂight intercept trap #134, 20.vii. 1993, (Richard Leschen)
(SMEC); BOLIVIA: La Paz: Chuma [dubious locality due to high elevation and cold
climate]: xii.l936 (Y. Mexia), 66, 49 (CAS). BRAZIL: Amazonas: AM 010, km 26,
Reserva Ducke, ex. malaise trap, 27 .ix.1978, (J. Arias), 19 (NMNH); Manaus, ii.l944,
(Praetorius), 19 (AMNH); Mata Grasa: 12° 50’S, 51° 45 ’W, Roy. Soc.-Roy. Georg Soc.
Aavantina-Cachimbo Exped. 1967-1969, coll. 1968 (BE. Freeman) 16 (BMNH); [Minas
Gerais]: Sete Lagoas, 19 (MZMC); Para: Benevides, ii.1895, (Gounelle), 29 (DEI); Rio
de Janeiro: (Kirsch), 19 (SMTD); Santa Catharina: Joinville, ?.ii.l920, (Melzer), 19
(DEI); No speciﬁc locality: ‘Sdo Paulo ’, 139 (DEI); ‘Brasil’, (W. Miiller), 29 (SMTD);
‘Brasilia’, 19 (MZMC); ‘Brazilien’, (Kratz), 19 (DEI); ‘Rio Madeira’ [river runs
through states of Rondonia and Amazonas], Madeira — Mamore R.R. Co Camp 35,
(Mann, Baker) 26 (NMNH). COLOMBIA: Santa Fe de Bagatd [=Bagata' D. C.], 16
(DEI). ECUADOR: [Esmeralda]s: Cachabé: low e., xii[l8]96 (Rosenberg) 16 (BMNH);
Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -0.652778, -76.433333,
220 m, ex. canopy fogging, i. 1994, (T.L. Erwin et al.), lot 599, 19, (NMNH); i.1994, lot
602, 19 (MSUC);.1994, lot 751, 16 (MSUC); vii.1994, lot 763, 16 (NMNH); vii.1994,
lot 774, 19 (NMNH); vii.1994, lot 776, 16 (NMNH); vii.1994, lot 777, 19 (NMNH);
vii.1994, lot 778, 26, 19 (NMNH); i.1995, lot 952, 36, 19 (NMNH); i.1995, lot 1030,
19 (NMNH); vii.1995, lot 1082, 16 (NMNH); x.1995, lot 1179, 29 (NMNH); x.1995,
lot 1217, 16 (NMNH); i.1996, lot 1452, l6, l9 (NMNH), 19 (MSUC); vii.1996, lot

123

1572, 16 (MSUC); vii.1996, lot 1604, 26 (MSUC), 29 (NMNH); vii.1996, lot 1618, 16
(NMNH); vi.l998, lot 1853, 16 (NMNH); vi.l998, lot 1856, 16‘, 19 (NMNH); x.1998,
lot 1949, 16 (NMNH); ii.l999, lot 1856, 16 (NMNH); Pastaza: Cusuimi, Rio Cusuimi,
150 km SE of Puyo, 320 m, 15-31.v.1971, (B. Malkin), 19 (FMNH). FRENCH
GUIANA: No speciﬁc locality: ‘Cayenne’, 19 (MZMC); ‘Cayenne’, 1915, (Felsche
Geschenk), 16, 19 (SMTD); ‘French Guiana’, l9 (NMNH). GUYANA: [=Region 8]:
Iwokrama Forest, 4.671822,-58.684114, 66 m, ex. Ocotea oblonga, 4-9.iii.2007,
(Cognato, Hulcr, Smith, Dale, McCall), 66, 49 (MSUC). PERU: [Hudnuco]: 15 mi NE
of Tingo Maria, 700 m, 11.xi.1954, (E.I. Schlinger, E.S. Ross), 49 (CAS); Junin: Satipo,
xii.l943, (Paprzycki), 16 (NMNH); Loreto: Iquitos [province not speciﬁed, Iquitos is in
Loreto], iii-iv.193l, (R. C. Shannon), 19 (NMNH); Madre de Dias: Los Amigos
Biological Station, -12.56916, -70.100114, 250 m, ex. Protium amazonicum, 26.iv.-
27.v.2008, (Smith, Hulcr) 366, 449. SURINAME: Saramacca: W Suriname Rd, 108
km WSW Zanderij Airport, 5°13’35N, 55°52’54”W, 30 m, ex. ﬂight intercept trap, 8-
10.vi.1999, (Z.H. Falin, B. DeDijn), l6 (SMEC). VENEZUELA: Amazonas: Cerro de la
Neblina, 1 km S Basecamp, 0°50’N, 66°10’W, 140 m, ex. along a small whitewater
stream; pools of dead leaves & sticks, 11.ii.1985, (P.J. & PM. Spangler, R. Faitoute, W.
Steiner), 16 (NMNH).
Diagnosis. This species has metallic elytra and is closely related to C. aeneipennis. The
male is distinguished by larger size, smooth elytral disk, shallow strial punctures and a
deep sulcus on the anterior pronotum extending to median line.

The female is diagnosed ﬁ'om C. aeneipennis by the larger size and shallow strial

punctures.

124

Description. Male 6.0 — 7.1 mm long (J? = 6.6 mm; 11 = 20); 1.8 — 2.0 times as long as

wide. Head, thorax, antennae and legs red-brown; pronotum commonly crimson, but may
also be black; abdomen black; elytra versicolor, ranging in color from metallic bronze to
metallic green; rarely metallic blue in Peru and Bolivia.

Apical epistomal margin is unarmed; lateral margins of epistoma with a sub-acute
arcuate costa pointing distally; epistoma strongly excavated, expanded to greater than
quarter length of head; basal epistomal margin armed with a transverse, concave carina;
frons strongly excavated and densely covered with long white to yellow hair-like setae;
margin of excavation lined with a single row of long hairs; hens and epistoma smooth,
shining; male scape elongate and narrow; glabrous on proximal quarter, setae on ventral
half, one third length of scape; funicle segments 2-7 with brushes of setae equal of length
of scape; dorsal margin with setae less than three frmicle segments; anterior face of the
club setose with a partial septum.

Anterior pronotal margin with a lateral declivity extending to median line, margin
between eyes lined with ﬁne biﬁd setae; pronotum glabrous; ﬁnely, minutely punctate;
lateral margin type B; basal half minutely punctate; elytra smooth; striae not impressed;
interstriae punctures strongly confused; base never with a tumescence from seventh to
ninth interstriae; scutellum type A; carina at base of interstria ten shorter than
metepistemum; metepistemum with biﬁd setae; metepistemum type B; mesostemum
type A; apex smooth.

Female similar to male except epistoma with a small median tubercle; ﬁ'ons ﬂat
with a triangular impressed area originating between the eyes and terminating at the distal

and lateral epistoma margins, the triangle is bisected by a shallow carina on the median

125

line. Frons ornamented short sparse setae; frons rugose, covered with moderate, deep
punctures; antennal funicle segments 1-7 with setae on dorsal and ventral margins less
than length of two funicular segments; gena with crescent shaped mycangium. Anterior
pronotal declivity weakly sulcate, margin between eyes lined with scales; pronotum
smooth, shining.

Etymology. noel (English) = Christmas (used as a noun in apposition). This species
commonly display ‘Christmas’ colors: crimson pronotum and brilliant metallic green
elytra.

Specimens examined. (77 6, 999)

Distribution. Bolivia (La Paz), Brazil (Amazonas, Mato Grosso, Minas Gerais, Para,
Rondonia?, Rio de Janeiro, Santa Catharina, S50 Paulo), Colombia (Bogota), Ecuador
(Esmeraldas, Napo, Pastaza), Guyana, Peru (Huanuco, Junin, Loreto, Madre de Dios),
Suriname, Venezuela (Amazonas).

Hosts. Ocotea oblonga (Meisn.) Mez, Protium amazonicum Swart.

Biology. C. noel colonize large branches ranging from 5 cm diameter to more than 20
cm. Females create an entrance tunnel running in a transverse plane toward the center of
the stem. From this tunnel, one or two circumferential branches are added creating
maternal galleries. The galleries closely resemble those of C. aeneipennis. C. noel
aggregate in leaves around host trees (Chapter 2) and perform maturation feeding on
leaves as (von Winning, 1929; chapter 2).

Discussion. Camptocerus noel has been considered C. aeneipennis for many years and is
identiﬁed as C. aeneipennis in collections. However, the male genitalis clearly diagnoses

the species. The apex of the median lobe is rounded in C. noel (mushroom shaped in C.

126

aeneipennis); the sensory claspers are spatulate in C. noel (scythe shaped in C.
aeneipennis). In addition to the morphological characters the validity of this species is
supported by molecular characters. The species is differentiated from C. aeneipennis by a
12.5% sequence divergence of mitochondrial cytochrome oxidase l and has a 5bp
insertion in the D2Fl-D3R2 region of nuclear ribosomal 28S. Little nucleotide variation
is seen within each species (Smith & Cognato, unpublished).

Geographic variation was found in several species, but most notable is the size

differences and slight color variation of C. noel. The elytra of C. noel is a brilliant

I'tiém

metallic green in northern South America and the color changes to a metallic bronze in
southern Peru and to metallic blue in southern Peru and Bolivia. Color also varies

between teneral and mature adults; mature adults are darker colored.

Camptocerus pseudaangustior Smith & Cognato, sp. n.
(Figs. 1.69, 1.70)

Type Material. Holotype 6, ECUADOR: Pastaza: Cononaco, ex. malaise trap,
29.iv. 1976, (J. Cohen) (NMNH). Paratypes, BRAZIL: Amazonas: km 60 N Manaus,
18.1 km E Campinas ﬁeld station, 2° 30’S, 60°15’W, ex. terra ﬁrme forest fogged with
pyrethrum, (Montgomery, Erwin, Schimmel, Krischik, Date, Bacon), sample 20,
22.ii.1979, 19 (NMNH); Manaus, 1 km W Taruma Falls, 100 m, 2.iii.l981, (C. Young),
19 (RJRC); 69 km N Manaus, 7.xii.1979, (G. Stevens) 16, 19 (NMNH); Teffé (Ega)
‘7bre-8bre.1879’ (M. de Mathan), 19 (NMHW); Pard: Santarém 16 (BMNH); ‘Par 8’
l9 (BMNH). ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare

Camp, -0.65277 8, -76.433333, 220 m, ex. canopy fogging, vii.1994, (T.L. Erwin et al.),

127

lot 684, 16 (NMNH); vii.1995, lot 1125, 19 (NMNH); x.1995, lot 1185, 16 (NMNH);
x.1995, lot 1265, 16 (NMNH); vii.1996, lot 1615, 26, 19 (MSUC), 19 (NMNH);
x.1996, lot 1755, 16 (NWH); Tiputini Biodiversity Station, -0.631944, -76.144167,
220—250 m, ex. canopy fogging, vi.l998, (T.L. Erwin et al.), lot 1876, 16 (NMNH);
x.1998, lot 1927, 19 (NMNH); vi.l998, lot 1950, 19 (MSUC); Pastaza: Cononaco, ex.
malaise trap, 29.iv.1976, (J. Cohen), 16, 29 (NMNH); PERU: [=Amazanas]: ‘Peru, Rio
Santiago’ xi.12. 1924, 19 (AMNH); [Huanuco]: 15 mi NE of Tingo Maria, 700 m
11.xi.1954, (EJ. Schlinger, E.S. Ross), 19 (NMNH); Loreto: 58 km SSW from Iquitos to.
Nauta, Itaya river 120m, 6.ii.2007, (A. Petrov) 16 (MSUC); Madre de Dios: Los Amigos
Biological Station, -12.56916, -70.100114, 250 m, ex. Protium amazonicum, 10-
l6.v.2008, (Smith, Hulcr) 56, 279 (MSUC); 7-25.v.2008, (Smith, Hulcr) 36, 49
(MSUC); Z.R. Tampopata, 300 m, iv-v.1986, (I. Bohérquez), 19 (MU SM); NO
SPECIFIC LOCALITY: ‘Bassin de l’Amazone’, [Amazon Basin] 1913, (Guilherma
Haﬂ'mann) [Eggers cotype Camptocerus aterrimus] 16 (NHMW).
Diagnosis. This species is black with dark brown gena and legs. Males are distinguished
by the generally larger body size, median tubercle on the apical epistoma margin,
transversely impressed, groove on pronotum base, basal medial pronotum region strongly
rugose disk, elytral interstrial punctures mostly confused.

Females are distinguished from C. aterrimus larger size and smaller, shallower

punctures on declivital striae.
Description. Male 4.5 — 5.4 mm long (J? = 5.1 mm; 11 = 18); 1.7 — 2.0 times as long as

wide. Head, antennae, thorax, abdomen and elytra and legs black; gena and legs dark red

brown.

128

 

Apical epistomal margin with a median tubercle; lateral margin of epistoma with
a sub-acute arcuate costa pointing distally; epistoma strongly excavated, expanded to less
than quarter length of head; basal epistomal margin armed with a transverse, concave
carina; tuft of thick apically rounded setae above scape insertion; frons strongly
excavated and densely covered with long white hair-like setae; margin of excavation
lined with a single row of long hairs; hairs near ocular margin with their apex ﬂattened;
male scape elongate and narrow; glabrous on proximal quarter, setae on ventral half, one
third length of scape; ﬁ'ons and epistoma smooth, shining; funicle segments 2-7 with
brushes of setae equal of length of scape; dorsal margin with setae less than three funicle
segments; anterior face of the club setose with a partial septum.

Anterior pronotal declivity entirely sulcate, margin between the eyes lined with
ﬁne biﬁd setae; pronotum glabrous; smooth, shining; minutely punctate; lateral margin
type B; basal half deeply rugose creating a wrinkled appearance; transverse medial
groove on base; base straight.

Elytra glabrous; interstriae punctures strongly confused on disk; punctures
shallow, minute; elytra smooth; elytra smooth; base never with a tumescence from
seventh to ninth interstriae; scutellum type A; carina at base of interstria ten longer than
metepistemum; metepistemum with biﬁd setae; metepistemum type B; mesostemum
type A; apex smooth.

Female similar to male except epistoma with a small median tubercle; frons ﬂat; a
medial impressed triangular region; a shallow carina originating on the median line
between the eyes and terminating at the epistoma; frons ornamented short, sparse setae;

ﬁ'ons shagreen, covered with large, deep punctures; antennal funicle segments 1-7 with

129

setae on dorsal margin less than length of two funicular segments; gena with crescent
shaped mycangium. Anterior pronotal declivity partially sulcate, margin between eyes
lined with scales; pronotum smooth, shining; transverse medial groove on base absent.
Etymology. Pseuda- (G) = false, - angustior = a Camptocerus species.

Specimens examined. (206, 469)

Distribution. Brazil (Amazonas; Para), Ecuador (N apo, Pastaza), Peru (Amazonas,
Huanuco, Junin, Loreto, Madre de Dios).

Hosts. Protium amazonicum Swart, Protium sp.

Biology. C. pseudoangustior infest 2 - 5 cm diameter branches. The maternal gallery
extends transversely through the stem to near the opposite side. Females create a vertical
maternal gallery extending above and below the entrance tunnel in the pith of small
branches and twigs. Two short radial maternal galleries are created in larger stem.
Females lay single eggs along the maternal gallery in wide niches cut at right angles to
the gallery and plugged with boring dust. Larvae enlarge their cradles parallel to the
maternal gallery and parallel to the grain of wood. Galleries strongly resemble those of C.
aterrimus. Beetles aggregate in leaves around host trees and perform maturation feeding
on leaves (Chapter 2).

Discussion. Specimens of this species have been commonly identiﬁed as C. angustior in

collections and keys (Wood, 2007).

Camptocerus suturalis (F abricius)

(Figs. 1.76,1.77)

130

Camptocerus suturalis (F abricius) 1801: 393 (Hylesinus). Type locality: ‘Essequibo’
[River] [=Guyana]; Lectotype (3‘ (here designated), UZMC.
Camptocerus cinctus Chapuis 1869: 51. Type locality ‘Amerique meridionale’ [=South
America]; Holotype 92, IRSNB. syn. n.
Camptocerusfasciatus (F abricius) 1801: 392 (Hylesinus). Type locality: ‘Essequibo’
[River] [=Guyana]; Lectotype 6 (here designated), UZMC. Synonymy: Eggers 192%:
45.
Notes. Synonymy of C. fasciatus conﬁrmed by Wood (1972: 243).
Diagnosis. Both sexes display a unique color pattern described below. The elytra is
moderately covered by short recumbent golden hairs on the striae and interstriae. The
male is distinguished from other species in the clade by the vertical costae on the lateral
margins of the epistoma, and transverse weakly developed carina on the proximal
epistomal margin.

The female is recognized by the anterior pronotal margin with setae palmately
divided into 34 ﬁlaments on between the eyes and by the absence of mycangia on the

gena or vertex.
Description. Male 3.6 — 5.0 mm (3 = 4.3 mm; 11 = 14); 1.8 — 2.1 times as long as wide.

The head, antennae, thorax, pronotum, legs are red-orange to red. Each elytron black with
an orange oval spot extending from the lateral margin of the elytral interspace to the tenth
interstria and from the basal ﬁfth of elytra to the of apical four-ﬁﬁhs. Body moderately
covered by recumbent, hair-like setae. Setae are often abraded.

Apical epistomal margin unarmed; lateral margin of epistoma with a vertical sub-

acute costa pointing distally; epistoma strongly excavated; expanded to greater than

131

quarter length of head; epistoma covered with a few short setae; basal epistomal margin
armed with a glabrous, transverse, concave carina; frons strongly excavated and densely
covered with long golden yellow hair-like setae; margin of excavation lined with a single
row of long hairs; frons and epistoma shagreen; male scape elongate and narrow;
glabrous on proximal quarter, setae on ventral half, one third length of scape; funicle
segments 2-7 with brushes of setae 1.5 times the length of scape; dorsal margin with setae
less than three funicle segments; anterior face of the club setose with a partial septum.

Anterior pronotal declivity partially sulcate, margin between the eyes lined with
ﬁne setae palmately divided into three or four ﬁlaments; pronotum smooth, covered with
recumbent hairs (often abraded); ﬁnely punctate, punctures moderately deep; lateral
margin type B; base straight.

Elytra densely covered with long recumbent golden hairs on both striae and
interstriae (often abraded); striae weakly impressed, punctures small, shallow; interstriae
punctures minute, strongly confused; base never with a tumescence from seventh to ninth
interstriae; scutellum type A; carina at base of interstria ten shorter than metepistemum;
metepistemum with biﬁd setae; metepistemum type B; mesostemum type A; apex
smooth.

Female similar to male except epistoma with a small median tubercle; frons ﬂat
with a shallow carina on the median line originating between the eyes and terminating at
the epistoma; ﬁons ornamented short, moderately dense setae; frons shagreen, covered
with moderate, deep punctures; antennal funicle segments 1-7 with setae on dorsal
margin less than length of two funicular segments; mycangia on the gena and vertex

absent. Pronotum anterior margin rounded.

132

Specimens examined. (326, 31 $2, 3 unknown sex)

Type Material. Lectotype Camptocerus suturalis, 6‘ (UZMC); paralectotypes
Camptocerus suturalis, NO LOCALITY: 16‘, 3Q (MZMC); Holotype Camptocerus
cinctus, 92 (IRSNB); Lectotype Camptocerusfasciatus, NO LOCALITY: 6‘ (MZMC);
Paralectotypes Camptocerusfasciatus, N O LOCALITY: 16‘, 3 Q, 1 unknown sex
(MZMC).

Other material: BRAZIL: Amazonas: 69 km N Manaus, (G. Stevens), 7.xii.1979, 16‘
(NMNH). ECUADOR: Napo: Reserva Ethnica Waorani, 1 km S Onkone Gare Camp, -
0.652778,—76.433333, 220 m, ex. canopy fogging, (T.L. Erwin et al.), lot 859, x.1994, IS?
(NMNH); lot 3265, vii.2006, 1S2 (NMNH). GUYANA: [=Region 8]: Iwokrama Forest,
4.671822, -58.6841 14, 66 m, ex. Ocotea oblonga, 4-9.iii.2007, (Cognato, Hulcr, Smith,
Dole, McCall), 116‘, 1092 (MSUC). PERU: Loreto: 68km SW from Iquitos to Nauta,
Nauta River, 120 m, 7.ii.2005, (A. Petrov), 126‘, 79, 1 unknown sex (MSUC); Madre de
Dios: Los Amigos Biological Station, -12.56916, -70.1001 14, 250 m, ex. Protium
amazonicum 26.iv.-27.v.2008, (Smith, Hulcr), 26‘, 3S? (MSUC). SURINAME: No
specific locality: ‘Surinam’, (H. Luna), 16‘ (UZMC). VENEZUELA: Amazonas: ‘Mt.
Duida’, 4.xi.1928, 16‘ (AMNH). N0 SPECIFIC LOCALITY: (292, 1 unknown sex;
UZMC).

Distribution. Brazil (Amazonas, Para), Ecuador (N apo), French Guiana, Guyana, Peru
(Loreto, Madre de Dios), Suriname, Venezuela (Amazonas).

Hosts. Ocotea oblonga (Meisn.) Mez, Protium amazonicum Swart, Protium sp.
Biology. In Guyana, Camptocerus suturalis adults create entrance trmnels within 1 mm

of C. aeneipennis entrance tunnels. The gallery systems of each species are separated by

133

 

a thin septum (less than 1mm), allowing fungal mycelium in C. aeneipennis galleries to
grow into those of C. suturalis indicating that C. suturalis is likely mycocleptic (Hulcr,
2009). Females create an entrance tunnel running in a transverse plane toward the center
of the stem. From this tunnel, two circumferential branches are added creating maternal
galleries. Galleries follow those of the ambrosia fungus host and the thin septum is
maintained. Females lay single eggs along the maternal gallery in wide niches cut at right
angles to the gallery and plugged with boring dust. Niches are only created on the surface f
opposite to the host gallery. Larvae enlarge their cradles with the grain of the wood. 1
Mycocleptism was also observed in Peru with Camptocerus suturalis parasitizing fungus

from galleries of C. noel, C. pseudoangustior, C. aeneipennis (Smith, per. ob.) and C.

mandelshtami (A. Petrov, per. com). This species has also been observed aggregating

in leaves around host trees (A. Petrov, per. com.) in the same manner as described in

chapter 2.

Discussion. Due to its mycocleptic habit and position within Camptocerus, C. suturalis

has likely lost its mycangia.

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. Characters and states are described in Materials

[CS

Morphological character matrix of 66 characters for 41 spec

Table 1.2

and Methods

Characters

 

19 20 21 22

9101112131415161718

8

Species

 

1

Camptocerus aeneipennis

C. angustior
C. annectens

C. aterrimus

C. auricomus

C. charpentierae
C. coccoformus

C. costatus

C. distinctus
C. dolei

C. hirtipennis
C. igniculus

C. inoblitus
C. latipilis
C. major

C. mallopterus

C. mandelshtami

C niger
C. noeli

C. occidentalis
C. opacicollis

Table 1.2 (cont): Morphological character matrix of 66 characters for 41 species. Characters and states are described in

Materials and Methods

Characters

 

19 2O 21 22

9101112131415161718

8

 

Species

C. orientalis

C. petrovi

C. pilifrons

C. pseudoangustior
C. quadridens

C. rectus

C. suturalis

C. unicomus

C. zuccus

?

‘?

S. multism'atus

S. propinquus

Scolytopsis peruanus
Scolytus excavatus

Cnemonyx boliviae
C. errens

C. insignis

C. rugulosus

C. vestitus

S. schevyrewi

S. ventralis

m

f 66 characters for 41 species. Characters and states are described

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Characters

 

44

42 43

32 33 34 35 36 37 38 39 40 41

23 24 25 26 27 28 29 30 31

l

pecies

Camptocerus aeneipennis

C. angustior
C. annectens

C. aterrimus

 

‘)

'.’

'.’

‘I

'2

C. auricomus

C. charpentierae

C. coccoformus
C. costatus

C. distinctus
C. dolei

139

C. hirtipennis
C. igniculus

C. inoblitus
C. latipilis
C. major

C. mallopterus

C. mandelshtami

C. niger
C. noeli

C. occidentalis
C. opacicollis

1n

. Characters and states are described '

ICS

Morphological character matrix of 66 characters for 41 spec

Materials and Methods

Table 1.2 (cont)

Characters

 

42 4344

32 33 34 35 36 37 38 39 40 41

23 24 25 26 27 28 29 30 31

C. orientalis

C pseudoangustior
C. quadridens

C. rectus

C. petrovi
C. pilifrons
C. suturalis

Species

 

C. unicomus
C. zuccus

140

Cnemonyx boliviae
C. errens

C. insignis

C. rugulosus

C. vestitus
Scolytopsis peruanus

DDO

2

Scolytus excavatus
S. multistriatus

S. pmpinquus

S. scheryrewi

S. ventralis

. Characters and states are described in

168

1 character matrix of 66 characters for 41 spec'

1C3

Morpholog

Materials and Methods

Table 1.2 (cont)

Characters

 

525354555657585960616263646566

45 46 47 48 49 50 51

0

 

Species

Camptocerus aeneipennis

C. angustior
C. annectens

C. aterrimus

C. auricomus

C. charpentierae

C. coccoformus
C. costatus

C. distinctus
C. dolei

141

C. hirtipennis
C. igniculus

C. inoblitus
C. latipilis
C. major

C. mallopterus

C. mandelshtami

C. niger
C. noeli

C. occidentalis
C. opacicollis

m

Morphological character matrix of 66 characters for 41 species. Characters and states are described

Materials and Methods

Table 1.2 (cont)

Characters

52 53 54 55 56 57 58 59 60 6162 63 64 65 66

45 46 47 48 49 50 5]

Species

 

C. orientalis

C. pseudoangustior
C. quadridens

C. rectus

C. petrovi
C. pilifrons

 

C. s'utura It's

142

B A

C. unicomus

C. zuccus
Cnemonyx boliviae
C. errens

C. insignis

C. rugulosus

C. vestitus

3

O

Scolytopsis peruanus

Scolytus excavatus
S. multistriatus

S. pmpinquus

S. schevyrewi

S. ventralis

Egg Niches

Maternal Galleries

Entrance Tunnel

 

Larvae

Larval Cradles

 

FIGURE 1.1: Horizontal cross section (A); vertical cross section (B) of Camptocerus
aeneipennis galleries.

143

    
   

TRINIDAD
2

ARGENTINA
3

FIGURE 1.2: Distribution map illustrating the number of Camptocerus species reported
for each country in the Neotropics. Countries without numbers lack records.

Camptocerus species range from Veracruz, Mexico (Atkinson & Equihua- Martinez,
1986) to Chaco, Argentina (Bruch, 1914).

144

Frons

  
 

Basal Epistomal
Margin

Epistoma

Apical Epistomal
Margin

FIGURE 1.3: Camptocerus suturalis male head illustrating terminology used for the
epistoma and frons.

145

 

FIGURE 1.4: Character 7: Male proximal epistomal margin (A) transverse, concave
carina; (B) tumid above scape insertion; (C) strongly developed transverse carina; (D)
wealdy developed transverse carina; (E) no demarcation; (F) transverse carina with
strongly developed lateral areas; (G) short transverse carina.

146

 

FIGURE 1.5: Character 14: Male frons vestiture on impressed area (A) absent; (B) few
setae, less than 20 on impressed area, frons clearly visible; (C) moderate setae; frons
largely covered by setae, but surface visible; (D) dense, frons surface not visible.

147

 

FIGURE 1.6: Character 21: Mycangia on female gena: (A) absent; (B) crescent shaped;
(C) oval; (D) triangular.

148

 

FIGURE 1.7: Character 25: Male anterior pronotal margin (A) rounded; (B) partially
sulcate; (C) entirely sulcate; (D) transversely impressed.

149

 

FIGURE 1.8: Character 26: Female anterior pronotum margin (A) rounded; (B) partially
sulcate; (C) transversely impressed.

150

 

FIGURE 1.9: Character 30: Lateral margin of pronotum (A) type A; (B) type B; (C) type
C; (D) type D; (E) type E; (F) type F; (G) type G; (H) type H; (I) type I-

151

 

FIGURE 1.10: Character 31: Pronotum base (A) straight; (B) recurved; (C) bisinuate;
(D) broadly emarginated along medial half.

152

 

FIGURE 1.11: Character 35: Metepistemum shape (A) type A; (B) type B; (C) type C;
(D) type D; (E) type B; (1“)13'1)e F; (G) type G; (H) type H; (I) type I; (J) W6 J; (K) type
K; (L) type L; (M) type M; (N) type N; (0)1ype 0; (P) type P; (Q) type Q; (R) type R-

153

 

FIGURE 1.12: Character 36: Mesosternum (A) type A; (B) type B; (C) type C; (D) type
D; (E) type B; (F) type F; (G) type G; (H) type H; (I) type I; (J) type J; (K) type K; (L)
type L; (M) type M; (N) type N; (0) type 0; (P) type P-

154

 

FIGURE 1.13: Character 37: Basal margin of elytra (A) ﬂat; (B) faint carina; (C) faint
carina extending to scutellar apex (D) crenulations.

155

 

FIGURE 1.14: Character 38: Scutellum shape (A) type A; (B) type B; (C) type C; (D)
type D; (E) type B; (F) type F; (G) type G; (H) type H; (1) type I; (I) type J; (K) type K;
(L) type L; (M) type M; (N) type N; (0) type 0; (P) type P-

156

 

FIGURE 1.15: Character 52: Density of elytral setae (A) glabrous; (B) light; (C)
moderate; (D) dense, elytral surface not visible.

157

FIGURE 1.16: Phylogeny of Camptocerus. Phylogram of one of 26 most parsimonious
trees (432 steps; consistency index, 0.488; retention index, 0.694) generated for 39 taxa
with 66 characters ﬁ'om a heuristic search of 1000 stepwise random additions with TBR
in PAUP“. Numbers above the node indicate bootstrap values (>50) based on 1000
pseudoreplicates. Bremer support values are generated from a heuristic search of 100
random addition sequence replicates.

158

Camptocerus aeneipennis
Camptocerus angustior
Camptocerus aterrimus
0 Camptocerus pseudoangustior
Camptocems mandelshtami
Camptocerus noel
Camptocerus hirtipennis
Camptocerus suturalis
Camptocems distinctus
Camptocerus charpentierae
Camptocerus coccoformus
Camptocerus latipilis
Camptocems petrovi
Camptocerus dolei
Camptooems mallopterus
Camptocerus igniculus
Camptocerus costatus
Camptocerus quadridens
Camptocerus zucca
Camptocerus pilifmns
75 Camptocerus auricomus
78 Camptocerus oﬁentalis
2 Camptocerus niger
Camptocerus tectus
Camptocerus occidentalis
Camptocerus unicomus
Camptocerus lnoblitus
Camptocerus opacicollis
Cnemonyx rugulosus
Cnemonyx vesﬁtus
Cnemonyx boliviae
Cnemonyx insignis
Cnemonyx enens
Soolytopsls peruanus
Scolytus excavatus
Scolytus pmpinquus
Scolytus ventralis
Scolytus multisb'iatus
Scolytus schevyrewi
_ 5 changes

 
 
 
 
 
  
  
  
 
 
 
 
 
  
  
  
  
 

 

  
 
  
  
  
 
 
   
  

159

 

FIGURE 1.17: Pit mycangia on female C. aeneipennis vertex. Pits are 3-5 times the
diameter of other punctures and 3-10 times as deep.

160

 

FIGURE 1.18: Camptocerus suturalis stealing fungus from a C. aeneipennis gallery: (A)
close proximity of entrance holes, with the smaller C. suturalis on top; (B) entrance
tunnel and two maternal galleries, C. suturalis created the bottom gallery;(C) maternal
galleries separated by a thin septum, within the region of fungal staining, C. suturalis is
in the bottom gallery.

161

 

FIGURE 1.19: Lateral (A) and dorsal (B) views of Camptocerus aeneipennis, 4.4 — 5.9
mm, male.

162

 

FIGURE 1.20: Camptocerus aeneipennis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

163

 

FIGURE 1.21: Lateral (A) and dorsal (B) views of Camptocerus angustior, 5.0 — 5.8
mm, male.

    

A

FIGURE 1.22: Camptocerus angustior male head anterior (A), anterior oblique (B).

 

165

 

FIGURE 1.23; Lateral (A) and dorsal (B) views of Camptocerus annectens, 3.5 mm,
female holotype.

166

 

A

FIGURE 1.24: Camptocerus annectens female holotype head anterior (A), anterior
oblique (B).

167

 

  

FIGURE 1.25: Lateral (A) and dorsal (B) views of Camptocerus aterrimus, 3.1 — 4.5
mm, male.

168

 

FIGURE 1.26: Camptocerus aterrimus male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

169

 

FIGURE 1.27: Lateral (A) and dorsal (B) views of Camptocerus auricomus, 3.5 — 4.1
mm, male.

170

 

FIGURE 1.28: Camptocerus auricomus male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

171

 

FIGURE 1.29: Lateral (A) and dorsal (B) views of Camptocerus charpentierae, 3.8 —
4.5 mm, holotype male.

172

 

FIGURE 1.30: Camptocerus charpentierae holotype male head anterior (A), anterior
oblique (B); female head anterior (C), anterior oblique (D).

173

 

. .1 - (1.

FIGURE 1.31: Lateral (A) and dorsal (B) views of Camptocerus coccoformus, 2.7 mm,
holotype male.

174

 

FIGURE 1.32: Camptocerus coccoformus holotype male head anterior (A), anterior
oblique (B); allotype female head anterior (C), anterior oblique (D).

175

 

FIGURE 1.33: Lateral (A) and dorsal (B) views of Camptocerus costatus, 3.5 - 3.9 mm,
male.

176

 

FIGURE 1.34; Male declivity (A) and female lateral (B) views of Camptocerus costatus.

177

 

C

FIGURE 1.35: Camptocerus costatus male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

178

 

FIGURE 1.36: Lateral (A) and dorsal (B) views of Camptocerus distinctus, 5.3 mm,
holotype male.

179

 

FIGURE 1.37: Camptocerus distinctus holotype male head anterior (A), anterior oblique
(B); allotype female head anterior (C), anterior oblique (D).

180

 

FIGURE 1.38: Lateral (A) and dorsal (B) views of Camptocerus dolei, 3.1 — 3.5 mm,
holotype male.

181

 

FIGURE 1.39: Camptocerus dolei holotype male head anterior (A), anterior oblique (B);
allotype female head anterior (C), anterior oblique (D).

182

 

FIGURE 1.40: Lateral (A) and dorsal (B) views of Camptocerus hirtipennis, 4.9 — 5.6
mm, male.

183

 

FIGURE 1.41: Camptocerus hirtipennis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

184

 

FIGURE 1.42: Lateral (A) and dorsal (B) views of Camptocerus igniculus, 2.5 — 2.8
mm, male.

185

 

FIGURE 1.43: Camptocerus igniculus male head anterior (A), anterior oblique (B);
allotype female head anterior (C), anterior oblique (D).

186

 

FIGURE 1.44: Lateral (A) and dorsal (B) views of Camptocerus inoblitus, 2.4 — 3.4 mm,
male.

187

 

FIGURE 1.45: Camptocerus inoblitus male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

188

 

FIGURE 1.46: Lateral (A) and dorsal (B) views of Camptocerus latipilis, 2.7 - 3.3 mm,
male.

189

 

v‘ﬁ".

T

was”:

 

FIGURE 1.47: Lateral (A) and dorsal (B) views of Camptocerus latipilis, 2.7 - 3.3 mm,
female.

190

  

C 'LD,

FIGURE 1.48: Camptocerus latipilis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

191

 

FIGURE 1.49: Lateral (A) and dorsal (B) views of Camptocerus major, 3.4 mm,
holotype female.

192

 

FIGURE 1.50: Camptocerus major holotype female head anterior (A), anterior oblique
(B)-

193

 

FIGURE 1.51: Lateral (A) and dorsal (B) views of Camptocerus mallopterus, 3.8 — 3.9
mm, holotype male.

194

 

      

C

FIGURE 1.52: Camptocerus mallopterus holotype male head anterior (A), anterior
oblique (B); allotype female head anterior (C), anterior oblique (D).

D

195

   

r111.
LBH'

FIGURE 1.53: Lateral (A) and dorsal (B) views of Camptocerus mandelshtami, 5.9 —
6.5 mm, male.

196

 

FIGURE 1.54: Camptocerus mandelshtami male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

197

 

B ‘

FIGURE 1.55: Lateral (A) and dorsal (B) views of Camptocerus niger, 2.8 — 4.1 mm,
male.

198

 

FIGURE 1.56: Camptocerus niger male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D).

199

 

FIGURE 1.57 : Lateral (A) and dorsal (B) views of Camptocerus noel, 6.0 — 7.1 mm,
male.

200

 

FIGURE 1.58: Camptocerus noel male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D).

201

 

FIGURE 1.59: Lateral (A) and dorsal (B) views of Camptocerus occidentalis, 4.0 - 4.5
mm, male.

202

 

FIGURE 1.60: Camptocerus occidentalis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

203

 

    

FIGURE 1.61: Lateral (A) and dorsal (B) views of Camptocerus opacicollis, 2.2 — 3.6
mm, male.

204

   

FIGURE 1.62: Camptocerus opacicollis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

205

 

FIGURE 1.63: Lateral (A) and dorsal (B) views of Camptocerus orientalis, 3.2 — 3.7
mm, holotype male.

206

 

FIGURE 1.64: Camptocerus orientalis holotype male head anterior (A), anterior oblique
(B); paratype female head anterior (C), anterior oblique (D).

207

 

FIGURE 1.65: Lateral (A) and dorsal (B) views of Camptocerus petrovi, 4.0 mm,
holotype male.

208

 

FIGURE 1.66: Camptocerus petrovi holotype male head anterior (A), anterior oblique
(B); abdomen posterior oblique (C).

209

 

FIGURE 1.67: Lateral (A) and dorsal (B) views of Camptocerus piliﬁ'ons, 2.6 - 3.1 mm,
holotype male.

210

 

FIGURE 1.68: Camptocerus pilifrons holotype male head anterior (A), anterior oblique
(B); allotype female head anterior (C), anterior oblique (D).

211

 

FIGURE 1.69: Lateral (A) and dorsal (B) views of Camptocerus pseudoangustior, 4.5 —
5.4 mm, male.

212

   

FIGURE 1.70: Camptocerus pseudoangustior male head anterior (A), anterior oblique
(B); female head anterior (C), anterior oblique (D).

213

   

FIGURE 1.71: Lateral (A) and dorsal (B) views of Camptocerus quadridens, 3.4 - 3.7
mm, male.

214

 

FIGURE 1.72: Posterior oblique (A) view of Camptocerus quadridens male. Lateral (B)
view of female.

215

   

FIGURE 1.73: Camptocerus quadridens male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

216

   

FIGURE 1.74: Lateral (A) and dorsal (B) views of Camptocerus rectus, 3.5 — 4.3 mm,
male.

217

   

C

FIGURE 1.75: Camptocerus rectus male head anterior (A), anterior oblique (B); female
head anterior (C), anterior oblique (D).

218

   

FIGURE 1.76: Lateral (A) and dorsal (B) views of Camptocerus suturalis, 3.6 — 5.0 mm,
male.

219

 

FIGURE 1.77 : Camptocerus suturalis male head anterior (A), anterior oblique (B);
female head anterior (C), anterior oblique (D).

220

 

FIGURE 1.78: Lateral (A) and dorsal (B) views of Camptocerus unicomus, 2.5 mm,
holotype male.

221

    

A

FIGURE 1.79: Camptocerus unicomus holotype male head anterior (A).

 

222

 

FIGURE 1.80: Lateral (A) and dorsal (B) views of Camptocerus zucca, 4.0 — 4.1 mm,
holotype male.

223

 

   

FIGURE 1.81: Camptocerus zucca holotype male head anterior (A), anterior oblique
(B); allotype female head anterior (C), anterior oblique (D).

224

 

CHAPTER TWO

Observations of Camptocerus Dejean (Coleoptera: Curculionidae: Scolytini) in Peru.

225

ABSTRACT

I report observations of Camptocerus attracted to felled Protium amazonicum
Swart in southeastern Peru and the behavior of four species. An unusual behavior for
scolytines was observed as three species aggregated to leaves surrounding potential hosts,
bored into leaves and fed, a rare behavior for Scolytines. Males competed for mates with

physical contact. Ant predation killed ~50% of colonizing individuals for one species.

226

INTRODUCTION

Neotropical scolytines represent a relatively unknown fauna. It is been estimated
that only two-thirds of South American species have been described (Wood, 2007) and
very little is known about their biology (Beaver, 1972; Wood, 2007). One poorly known
xylomycetophagous genus is Camptocerus Dejean (Coleoptera: Curculionidae:
Scolytinae), despite being among the ﬁrst described American scolytines (Blandford,
1896; Illiger, 1807). However, certain aspects of its biology have been documented. Like
other ambrosia beetles, Camptocerus have specialized mycangia that carry fungal spores.
Females bore into the xylem and sapwood and create ‘galleries.’ The symbiotic fungus
grows in the galleries where eggs are laid. Larvae enlarge their own galleries and feed on
both xylem and fungal mycelia; Some species demonstrate some level of host speciﬁcity,
but, less than half of Camptocerus species have been associated with a host tree (Wood &
Bright, 1992). Camptocerus aeneipennis (F abricius) and Cnemonyx vismiae Eggers have
been observed to chew holes as wide as their pronotum in leaves and lodge themselves in
them. Feeding behavior around this hole was reported from the Panama Canal Zone for
both species (von Winning, 1930). Camptocerus aeneipennis were observed to simply
rest or pass through these holes in Venezuela (Wood, 2007). It has been presumed that
this behavior is maturation feeding (Wood, 2007). Beaver (1972) presented the ﬁrst study
of Camptocerus biology and provided a detailed account of host use and life-history
information for four species in Brazil. Beaver cut trees in the Brazilian Amazon, north of
Manaus and allowed them to rest in situ for 1 to 2 months before dissecting beetles out of

infested trees. Camptocerus aeneipennis, C. aterrimus Eggers C. niger (F abricius), C.

227

 

aquilus Wood (=C. opacicollis (Eggers)) were collected and studied. He also provided
the ﬁrst summary of hosts for the genus.

An ecological study to assess ambrosia beetle (Coleoptera: Curculionidae:
Scolytine and Platypodine) diversity undertaken in Peru provided an unexpected
opportunity to further describe the biology of Camptocerus. Four species were found and
monitored during the study: C. opacicollis (Eggers), C. suturalis (Fabricius) and two new
to science Camptocerus noel Smith & Cognato and Camptocerus pseudoangustior Smith
& Cognato. Observations of life-history aggregation, leaf-feeding, courtship and mating

were observed and are reported herein.

MATERIALS AND METHODS

The study was performed at Centro de Investigacién y Capacitacion Rio Los
Amigos in Madre de Dios, Peru (S12°34’9” W70°6’0.04”, 248 m elevation). Species of
Protium (Burseraceae) are the most ﬁ'equently documented hosts for Camptocerus
species (Beaver, 1972; Wood, 2007). The species Protium amazonicum Swart was
chosen as a bait tree because of its wide range throughout Pem (Paul Fine, per. com.).
Preparation of the trees followed the protocol of Hulcr et a1. (2008). Six 10-15cm DBH
trees were used in the study. Three trees were killed standing by girdling at 1.5m
removing 300m circumferential strips of bark with multiple cuts on 27 April, and then
burned at the base on 29 April 2008. These trees (“A”-“C”), a fourth “D” which was
girdled but not burned and a ﬁfth ungirdled tree (“E”) were felled on 29 April. A sixth P.

amazonicum tree (“F”) was felled and then girdled on 11 May. All trees were felled in

228

 

secondary forest and left in situ to attract ambrosia beetles. Trees were examined
approximately every other day throughout the duration of the ecological study and

observations were recorded.

RESULTS

Colonization

Initial galleries were created 1-2 days after girdling (28-29 April) by Camptocerus
noel females, however additional galleries were not observed until seven days after
girdling (4 May). The ﬁve initial attacks were restricted to the girdle and occurred before
the trees were felled. Attacks rapidly became more frequent with 40 after nine days (6
May), 80 after 11 days (8 May) and continued until 15 days after girdling (12 May),
when they started to diminish but continued on all trees until 26 days after girdling (23
May) when the study was terminated. Camptocerus noel was the ﬁrst species to arrive on
28 April, followed by Camptocerus pseudoangustior on 4 May, C. suturalis on 10 May
and C. opacicollis on 16 May. Camptocerus suturalis galleries were constructed next to
larger galleries created by Camptocerus noel and Camptocerus pseudoangustior that
were 3-6 days old. Protium specimen “D” was not girdled prior to cutting and
colonization did not begin until 4 May and Protium specimen “E” received its ﬁrst
attacks on 6 May. There did not appear to be a difference between girdled and ungirdled

trees for the start of colonization. Each tree was colonized by all four species.

Aggregation

229

 

Camptocerus were ﬁrst observed 2 May, three days after the trees were cut and
ﬁve days aﬁer girdling. Camptocerus noel and Camptocerus pseudoangustior were
initially found within 1m of two of the P. amazonicum trees. Camptocerus opacicollis
were observed aggregating 19 days after the trees were cut and corresponded to the start
of gallery creation by the species. All species except C. suturalis aggregated in leaves of
nearly every plant located within 2 m of the felled P. amazonicum trees. Host leaves were
utilized approximately 10% of total aggregation observations. Leaves with aggregating
beetles ranged ﬁom thick and waxy to new and thin, living and dead leaves, and leaves ,
with and without latex. Leaves represented 18 families and encompassed trees, shrubs,
grasses, climbers and ferns with the exception of leaves with large trichomes and
included the following taxa: Aspidosperma parvifolium (Apocynaceae), Bactris spp.
(Arecacae), Geonoma sp. (Arecaceae), Matisia ochrocalyx (Bombaceae), Protium
amazonicum (Burseraceae), Chrysobalanaceae, Symphonia globulifera (Clusiaceae),
Bauhinia sp. (F abaceae), Inga spp. (F abaceae), Eschweilera coriacea (Lecythidaceae),
Carpotroche longifolia (F lacourtiaceae), Lomariopsis japurensis (Lomariopsidaceae),
Calathea spp. (Marantaceae), Marantaceae, OIyra sp. (Poaceae), Petrea maynensis
(Polygonaceae), Polygonaceae, Polybotrya sp. (Pteriodophyta), Rubiaceae,Leonia
glycicatpa (V erbenaceae), Costus sp. (V iolaceae), Zingiberaceae. Leaves were selected
regardless of their spatial orientation to the forest ﬂoor. Holes always touched at least one
vein and ﬁ'equently touched the midrib and a secondary vein. This was not true in grasses
where veins are abundant. No differences were observed between the sexes. There was

always a pile of dried leaf pieces near the beetles on the leaves (Fig. 2.1) and frass was

230

never observed. Leaves on three occasions contained two beetles at a time, once in the
case of Camptocerus noel and twice by Camptocerus pseudoangustior.

Beetles were observed with their abdomen above the leaf surface and the head
facing the ground in 3 of 113 observations. The pronotum was divided between the two
surfaces and the head was pushed tightly against the pronotum. The meso- and
metathoracic legs rested against one leaf surface and the prothoracic legs were held
tightly against the other leaf surface. Beetles were not observed to move and maintained
their grip on the leaf in observed situations, such as movements caused by wind and when
the leaf was brushed by an animal. The beetles tucked their legs in, dropped to the ground
and hid in the leaf litter when covered by shadows or perturbed by a predator. Empty
holes were ﬁ'equently observed and the beetles did not seem to return to the same hole
alter a disturbance.

Camptocerus beetles were also observed in leaves around an unidentiﬁed fallen
tree (15 cm DBH) that produced thick, white latex from cuts in the bark. More than 20
Camptocerus noel were found in leaves around this fallen tree, but none had attacked it.

There were over 100 holes in leaves around this tree.

Leaf Feeding
Feeding on the leaf surface near the hole was observed. Holes were identical to
those used for aggregation and were always placed adjacent to a rib. Leaf feeding was
observed for all species except C. suturalis. Feeding was restricted to leaves of the host
(P. amazonicum) and Carpotroche Iongifolia (Poepp.) Benth. (F lacourtiaceae). All

species were observed in leaves of P. amazonicum, but Camptocerus noel was found

231

 

feeding on Catpotroche longtfolia approximately 90% of the time. F rass was always
present on the leaf surface near a beetle after 6 May. When beetles were present, the
leaves had evidence of window feeding on the upper side of the leaf in close proximity to
the hole (Fig. 2.2). Over time, feeding damage progressed to skeletonizing. Four
Camptocerus noel beetles were each observed once performing window feeding on the
upper leaf surface within 2 cm of their hole. When disturbed, they quickly retreated to
their hole with their abdomens on the upper leaf surface. Observation of leaf damage by
feeding was observed in one Camptocerus noel ﬁve days after girdling (2 May) and
became more common as time progressed with twenty Camptocerus noel observed after
nine days (6 May). Three Camptocerus pseudoangustior were ﬁrst observed after 11 days
(8 May) and three C. opacicollis after 19 days (16 May). Aggregation was observed in
equal frequency by both sexes.

Protium amzaonicum “F” was closely observed for the presence of beetles
beginning the day after felling. F orty-eight hours after felling, six beetles were observed
in leaves and one leaf had fresh signs of feeding. Feeding created skeletonizing damage
and both leaf surfaces were eaten with veins left intact. Eight days after felling, leaves
commonly showed window-feeding on the upper leaf surface. Dissections of aggregating
beetles revealed an empty gut, while dissections of feeding beetles yielded a full digestive
tract, conﬁrming the observations. Leaf and beetle vouchers are stored in the AJ. Cook

Arthropod Research Collection, Michigan State University (MSUC).

Courtship and Mating

232

Courtship was observed ten times and mating was observed four times in
Camptocerus noel. Mating was observed three times in Camptocerus pseudoangustior.
Camptocerus noel females stayed in the entrance tunnel with their abdomens above the
bark surface. Males walked the bark in presumed search of females. Two to three males
gathered around a female and males fought for access to the female. Fighting consisted of
charging at the opponent and pushing him away from the female’s vicinity by placing his
frons against the abdomen of the opposing male. The male brushed his antennal funicle
setae over the female abdomen in a ﬂuttering motion producing a whirling sound. He
then repeatedly placed his ﬁ'ons against her abdomen, which corresponded to the
concavity of his ﬁ'ons. The male periodically produced chirps using his elytra-abdominal
stridulatory mechanism, rather than the gula-prostemum stridulatory mechanism on his
head. While the female constructed the gallery, he waited outside and used his brush-like
antennae to sweep away boring-dust and ﬁ'ass. In both species, females stayed in the
enhance tunnel and copulated with the male as he stood on the bark surface. The beetles
mated at a 90-degree angle with the male perpendicular to the female elytra. Two digital
movies of these behaviors can be found at www.scolytid.msu.edu/

Camptocerus_biology.html.

Predation
Three bullet ants, Paraponera clavata (F .), were observed trying to pry
Camptocerus noel and pseudoangustior out of their entrance tunnels with their
mandibles. Ants were also seen twice carrying Camptocerus pseudoangustior females in

their mandibles on the trunk of one tree.

233

 

A Dolichoderus sp. worker was observed to hunt Camptocerus on leaves. A
Camptocerus noel adult was standing on a leaf and the ant lunged for it. The beetle
tucked its legs in and rolled off the leaf on to the forest ﬂoor where it buried itself in leaf
litter.

Workers of a Crematogaster species were observed on two P. amazonicum trees
performing engaging in behaviors that at ﬁrst did not resemble predation. Ants were
clustered around gallery entrances in excess of twenty individuals. As the male swept

frass and sawdust from the gallery and outside the tree, the ants came forward, grabbed

 

any residual frass or sawdust, removed it from the gallery, and dropped it to the ground.
Beetles were observed beating their abdomens against the sides of the gallery to block the
ant’s entry and attempted to push them out of their galleries. However predation became
evident when Crematogaster were also observed clustering around a Camptocerus noel
male on a branch. The ants ﬁequently attacked and engulfed beetles on the tree surface.
These beetles were likely paralyzed by the ants and were moribund or dead.
Crematogaster were observed to bite Camptocerus and kill 54 of 110 females creating
their galleries as well as numerous males blocking gallery entrances. The ants separated
the 7m sternite from the 8th tergite and hollowed out the beetles through the genital
opening. Ant predation was a signiﬁcant problem for Camptocerus pseudoangustior
(50% predation in one tree), but was observed to a lesser degree on Camptocerus noel
and C. opacicollis. No predation was observed on C. suturalis. A digital movie of
Crematogaster predation is available at www.scolytid.msu.edu/

Camptocerus_biology.html.

234

DISCUSSION

Colonization duration was longer than previously documented. This minimal 25-
day period was substantially longer than the 3-10 days that had been reported ﬁ'om Brazil
(Beaver, 1972). Observations of maturation feeding were identical to those reported by
von Winning (1930). Maturation feeding, ﬁrst observed by von Winning to account for
the behavior of C. aeneipennis, has now been documented for Camptocerus noel,
Camptocerus pseudoangustior and C. opacicollis. Only nine C. suturalis individuals
were collected during this study compared to a minimum of 200 for other species. C.
suturalis has also been observed perform aggregation and maturation feeding behaviors
in Loreto province, Peru (A. Petrov, per. com.). Similar observations were made with C.
aeneipennis and C. auricomus Blandford in Panama (Smith, per. obs.). However,
observations at Los Amigos suggest this behavior is actually two different behaviors in
Camptocerus: aggregation and maturation feeding. The pile of dried, unconsumed leaf
material next to aggregating beetles suggests the beetles were creating a place in which to
aggregate around the potential host. Beetles were also observed aggregating around a tree
that produced thick latex when bark was removed. It was apparent that the beetles had
been there for some time due to the large number of holes and maturation feeding
observed. It is possible the beetles were waiting for the latex to subside before
colonization.

The biology of Camptocerus is similar to the other genera in the tribe Scolytini
(Cnemonyx Eichhoff, Scolytus Geoffroy and Scolytopsis Blandford). Species of the

Scolytini are known to colonize moribund material in masse (Wood, 1982). Multiple

235

 

Camptocerus species were observed to colonize individual hosts in Brazil (Beaver,

1972), Colombia and Venezuela (Wood, 2007), Guyana and Panama (Smith, per. 0b.),
and Peru. When multiple Camptocerus species colonized a single host, the species
segregated by size, as previously reported (Beaver, 1972; Wood 2007). Large numbers of

beetles of a single species or of several species of Cnemonyx are common in the same

 

host (Wood, 2007; Smith per. 0b.). Camptocerus mating behavior differs from that of

Scolytus multistriatus (Marsham). Male S. multistriatus also place their frons against the

female abdomen, but copulation occurs end-to-end inside the gallery (Collins et al.,

1936). I
Prior to this study, limited information existed on predation of Camptocerus.

Beaver (1972) reported larval parasitism by an unidentiﬁed species of Braconidae and

predation by an unidentiﬁed Coleoptera larva. Ants such as species of Crematogaster, are

commonly known to prey upon ambrosia beetles (Browne, 1961), bark beetles (Moeck &

Safranyik, 1984; Gonzalez & Campos, 1990) and ﬁ'uit boring scolytines (Philpott &

Armbrecht, 2006). Thus, ant predation on Camptocerus is unsurprising given the

abundance of ants in the tropics. Beetles attempted to defend themselves by thrashing

their abdomens against the sides of their galleries and pushing ants out of their holes as

observed in Beaver (1972).

236

 

FIGURE 2.1: (A) Males of Camptocerus noel aggregating and resting in a leaf near a
felled Protium amazonicum tree. They made the holes in which they rest. (B) Female
Camptocerus noel resting in a leaf and (C) boring through the leaf with undigested leaf
cuttings.

237

 

FIGURE 2.2: Damage caused by feeding male of Camptocerus noel to a P. amazonicum
leaﬂet.

238

LITERATURE CITED

Anderson, W.H. & Anderson, D.M. (1971). Type specimens in the Hans Eggers
collection of scolytid beetles (Coleoptera). Smithsonian Contributions to Zoology,
94, 1-38.

Atkinson, T.H. & Equihua-Martinez, A. (1986) Biology of bark and ambrosia beetles
(Coleoptera: Scolytidae and Platypodidae) of a tropical rain forest in southeastern

Mexico with an annotated checklist of species. Annals of the Entomological
Society of America, 79, 414-423.

Beaver, RA. (1972) Biological studies of Brazilian Scolytidae and Platypodidae
(Coleoptera). I. Camptocerus Dej ean. Bulletin of Entomological Research, 62,
247-256.

Beaver, RA. (1989) Insect-fungus relationships in the bark and ambrosia beetles. Insect —
Fungus Interactions (ed. By N. Wilding, N.M. Collins, P.M. Hammond and IF
Webber), pp. 121-143. Academic Press, London.

Blackman, M.W. (1943) New species of American scolytoid beetles, mostly Neotropical.
United States National Museum, Proceedings, 94, 371-399.

Blackwelder, RE. (1947) Checklist of the Coelopterous insects of Mexico, Central
America, The West Indies and South America. United States National Museum,
Bulletin, 185, 765-925.

Blandford, W. F. H. (1893) The Scolyto-platypini, a new subfamily of Scolytidae.
Entomological Society of London, Transactions, 425-442.

Blandford, W.F.H. (1896) Scolytidae. Biologia Centrali-Americana, Coleoptera, 4,
97-144.

Blatchley, W.S. & Leng, C .W. (1916) Rhynchophora or weevils of north eastern
America. Nature Publishing Co., Indianapolis.

Bondar, G. (1950) Notas entomologicas da Baia, XXII. Revista de Entomologia, 21,
449-480.

Bright, DE. (1993) Systematics of bark beetles. Beetle Pathogen Interactions in Conifer
Forests (ed. T. Schowalter and G. Flip), pp. 23-36. Academic Press, New
York.

Bright, D.E. & Skidmore, RE. (1997) A Catalog of Scolytidae and Platypodidae

239

(Coleoptera), Suppl. 1 (1990-94). NRC Research Press, Ottawa.

Bright, D.E. & Skidmore, RE. (2002) A Catalog of Scolytidae and Platypodidae
(Coleoptera), Suppl. 2 (1995-99). NRC Research Press, Ottawa.

Browne, PG. (1961) The work of the West Aﬁican Timber Borer Research Unit for the
year 1960. Report of the West African Timber Borer Research Unit 4, 5-14.

Bruch, C. (1914) Coleopteros de la Repr’rblica Argentina. Pars 7. Revista Del Museo de la
Plata, 19, 427-429.

LaPort de Castelneau, FL. (1840) Histoire naturelle des animaux articules,
annelids, crustaces, arachinides, myriapodes et insects. P. Dumenil. Paris, 2, 366-
372.

Chapuis, F. (1869) Synopsis des Scolytides. Liege, J. Desoer. 56p. [Reprinted as: Societe
Royale des Sciences de Liége, Memoires, 2, 213-269, 1873].

Chevrolat, Louis AA. (1838) Xylophages, In Guerin-Menville Iconographie du regne
animal. Paris, 111: 181-183.

Collins, C.W., Buchanan, W.D., Whitten, R.R. & Hoffman, CH. (1936) Bark beetles and
other possible insect vectors of the Dutch Elm disease Ceratostomella ulmi
(Schwarz) Buisman. Journal of Economic Entomology, 29,169-176.

Da Costa Lima, A.M. (1956) Insectos do Brazil. 10 Tomo, Coleopteros, 4 Ultima Parte.
Escola Nacional De Agronomia, Rio de Janeiro.

Dejean, P.F.M.A. (1821) Catalogue de la collection de coléopteres de M. Le Baron
Dejean, lieutenant-general des armées du roi, commandeur de l’ordre royal de la
Le’gion d’honneur, chevalier de l’ordere royal et militaire de Saint-Louis. Paris.
Crevot, 136pp.

Eggers, H. (1928) Ipidae (Coleoptera) da America do Sul. Archivos do Instituto
Biologico, 1, 83-99.

Eggers, H. (1929) Zehn neue Loganius-arten (Ipidae, Col.) aus Sildamerika. Wiener
Entomologische Zeitung, 46, 59-65.

Eggers, H. (1932) Borkenkafer (Ipidae, Col.) aus Sudamerika, VI. Material des Museum
Paris aus Franz. Guayana und Venezuela. T ravaux du Laboratoire d ’Entomologie,
Museum National d ’Histoire Naturelle, Memoires originaux, 1, 1-37.

Eggers, H. (1933) Zur Synonymie der Borkenkafer (Ipidae, Col). Entomologisches
Nachrichtenblatt, 7, 1 7-20.

240

Eggers, H. (1934) Zur Synonymie der Borkenkafer (Ipidae, Col.). Entomologisches
Nachrichtenblatt, 8, 25-29.

Eggers, H. (1943) Neue Borkenkafer aus dem Deutschen Entomologischen Institut.
Arbeiten uber Morphologische and T axonomische Entomologie, 10, 241-248.

Erichson, W.F. (1836) Systematische auseinandersetzung der familie der borkenkafer
(Bostrichidae). Archivﬁlr Naturgeschichte, 2, 45-65.

Evenhuis, N. L. (2007) Abbreviations for insect and spider collections of the world
[online] Available from http: //hbs. bishopmuseum. org/codens/codens-inst. htrnl
[accessed 10 February 2009].

Fabricius, J .C. (1801) Systema eleutheratorum, secundum ordines, genera, species,
adjectis synonymis, locis, observationibus, descriptionibus. Kiliae, Bibliopol.
Acad. Vol. 1, 24 + 506p., vol. 2, 687p.

Farrell, B.D., Sequeira, A.S., O’Meara, B.C., Normak, B.B., Chung, J.H. & Jordal, B.H.
(2001) The evolution of agriculture in beetles (Curculionidae: Scolytinae and
Platypodinae). Evolution, 55, 2011-2027.

Ferrari, J .A. (1867) Die Forst — und Baumzuchtschadlichen Borkenkfer (Tomicides Lac.)
aus der Familie der Holzverderber (Scolytides Lac.) Carl Gerold’s Sohn, Wien, 96

PP-

Girard, M.J.A (1873) Les Insects. Traité élémentaire d’entomologie comprenant
l’histoire des espéces utiles et de leurs produits, des espéces misables et des
moyens de les detuire, l’étude des métamorphoses et dans moeurs, les procédés de
chasse et de conservation. Paris. J .B. Bailliere et ﬁls.

Gonzalez, R., & Campos, M. (1990) Evaluation of natural enemies of the Phloeotribus
scarabaeoides (Bem.) (Col: Scolytidae) in Granada olive groves. Acta
Horticulturae, 286, 355-358.

Haack RA. (2001) Intercepted Scolytidae (Coleoptera) at US. ports of entry: 1985—
2000. Integrated Pest Management Reviews, 6, 253—282.

Hagedom, J .M. (1903) Enumeratio Scolytidarum e Guayana, Venezuela et Columbia
natarum Musei Historico-Naturalis Parisiorum, descriptionibus specierum
novarum adjectis. Museum d ’Histoire Naturelle, Bulletin, 1903, 545-550.

Hagedom, I.M. (1910) Coleoptera family Ipidae. Pars 111:1-17 8, in Wystrnan, Genera
Insectorium. Brussels. 178pp.

H0pkins, AD. (1909) Contributions toward a monograph of the Scolytid beetles. I. The
genus Dendroctonus. United States Department of Agriculture, Technical Series,

241

17, 1-164.

Hopkins, AD. (1914) List of generic names and their type-species in the Coleopterous
superfamily Scolytoidea. United States National Museum, Proceedings, 48, 115-
136.

Hopkins, AD. (1915) Contributions toward a monograph of the Scolytid beetles. H.
Preliminary classiﬁcation of the superfamily Scolytoidea. United States
Department of Agriculture, Technical Series, 17, 165-232.

Hulcr, J ., Mogia, M, Isua, B. & Novotny, V. (2007a) Host speciﬁcity of ambrosia and bark
beetles (Col., Curculionidae: Scolytinae and Platypodinae) in a New Guinea rainforest.
Ecological Entomology, 32, 762-772.

Hulcr, J., Dole, S.A., Beaver, R.A. & Cognato, A.I. (2007b) Cladistic review of generic
taxonomic characters in Xylorborina (Coleoptera: Curculionidae: Scolytinae). Systematic
Entomology, 32, 568-584.

Hulcr, J., Novotny, V., Maurer, B.A., & Cognato, A.I. (2008). Low beta diversity of
ambrosia beetles (Coleoptera: Curculionidae: Scolytinae and Platypodinae) in
lowland rainforests of Papua New Guinea. Oikos, 117, 214-222.

Hulcr, J. (2009). Diversity and systematics of ambrosia beetles (Curculionidae:
Scolytinae, Platypodinae) with emphasis on Xyleborina. Dissertation, Department
of Entomology, Michigan State University, East Lansing, Michigan, United
States.

Illiger, J .K.W. (1807) F abricischen Systeme fehlender Kafer-gattungen. Magazin fur
Insektenkunde, 6, 318-350.

Intemation Commission of Zoological Nomenclature. (1999) International Code of
Zoological Nomenclature (ed. by W.D.L. Ride, H.G. Cooper, C. Dupuis, O.
Kraus, A. Minelli, F.C. Thompson, P.K. Tubbs). 4th Edition. The International
Trust for Zoological Nomenclature, London.

Kelley, S.T. & Farrell, BB. (1998) Is specialization a dead end? The phylogeny of host
use in Dendroctonus bark beetles (Scolytidae). Evolution, 52, 1731-1743.

Kleine, R. (1913) Die geographische Verbreitung der Ipiden-Genera orbis terrarum.
(Col.). Berliner Entomologische Zeitschrift, 58, 1 13-176.

Kleine, R. (1914) Die geographische Verbreitung der Ipiden (Die auBereuropaischen
Gebiete.). Stettiner Entomologische Zeitung, 75, 243-410.

Kleine, R. (1934) Die Borkenkafer (Ipidae) und ihre Standpﬂanzen. Zeitschrtftﬁrr
Angewandte Entomologie, 21, 121-181.

242

Kniiek, M. & R. Beaver (2004). Taxonomy and systematics of bark and ambrosia beetles. Bark
and wood boring insects in living trees in Europe: a synthesis. (ed. By F. Lieutier, K. R.
Day, A. Battisti, J .-C. Gregoire and H. F. Evans), pp. 41-54. Kluwer Academic
Publishers, London.

Kuschel, G. (1995) A phylogenetic classiﬁcation of the Curculionoidea to families and
subfamilies. Memoirs of the Entomological Society of Washington, 14, 5-33.

Lacordaire, J .T. (1833) Coléoptéres de la Guyane F rancaise. Annales du Muséum d ’Hstoire
Naturelle, 12, 35-94.

Lacordaire, J .T. (1866) Histoire naturelle des insects genera des coléoptéres ou exposé
méthodique et critique de tous les genres proposes jusqu’ici dans cet ordre
d’insectes. Paris. Vol. 7, 620pp.

Latreille, PA. (1829) Les crustaces, le aracnides et les insects, distributes en famille
naturelle, ouvrage format des tomes 4 et 5 de celui de M. de Baron Cuvier sur le
regne animale (deuxieme edition). Deterville, Paris. vol. 2. xxiv + 556p.

Livingston, R. L. and A. A. Berryman (1972). Fungus transport structures in the Fir Engraver,
Scolytus ventralis (Coleoptera: Scolytidae). Canadian Entomologist, 104, 1793-1800.

Maddison, D.R. & Maddison, WP. (2000). MacClade 4: Analysis of Phylogeny and
Character Evolution. Version 4.0. Sunderland, MA: Sinauer Associates.

Maes, J .M. & Equihua Martinez, A. (1988a) Cataolgo des los scolytidae y platypodidae
(Coleoptera) de Nicaragua. Revista Nicaraguense de Entomologia, 3, 1-43.

Marvaldi, A. E. (1997) Higher level phylogeny of Curculionidae (Coleoptera:
Curculionoidea) based mainly on larval characters, with special reference to
broad-nosed weevils. Cladistics, 13, 285-312.

Marvaldi, A. E., A. S. Sequeira, C.W. O’Brien & Farrell, RD. (2002) Molecular and
morphological phylogenetics of weevils (Coleoptera, Curculionoidea): Do niche
shifts accompany diversiﬁcation? Systematic Biology, 51, 761-785.

Moeck H.A. & Safranyik, L. (1984) Assessment of predator and parasitoid control of
bark beetles. lst ed. Victoria (BC): Environment Canada Canadian Forestry
Service Paciﬁc Research Centre.

Nunburg, M. (1962) Zur kenntnis der Neotropischen borkenkaefer-fauna (Col.
Scolytidae). Papé is Avulsos do Departamento de Zoologia, Secretaria de
Agricultura, Si’ro Paulo, 15, 223-237.

Petrov, A.V. (2007) A new species of the genus Camptocerus Latreille (Coleoptera:

243

Scolytidae) ﬁ'om Peru. Russian Entomological Journal, 16, 101-102.

Philpott, S.M. & Armbrecht, I. (2006) Biodiversity in tropical agroforests and the
ecological role of ants and ant diversity in predatory function. Ecological
Entomology, 31, 369-377.

Schedl, K.E. (1931) Morphology of the Bark beetles of the genus Gnathotrichus Eichh.
Smithsonian Miscellaneous Collections, 82, 1-88.

Schedl, K.E. (1939) Scolytidae und Platypodidae (Col.). 68 Beitrag. Revista de
Entomologica, 10, 718-727.

Schedl, K.E. (1952) Neotropische Scolytoidea, 1]]. 110 Beitrag. Dusenia, 3, 343-366.

Schedl, K.E. (1962) Zur synonymie der borkenkafer VIII (Coleoptera). Beitra'ge zur
Entomologie, 12, 485-493.

Schedl, K.E. (1970) Scolytidae et Platypodidae (Coleoptera) récoltés en Guyane ﬁancaise
par la Mission du Museum National d’Hisroire Naturelle (1). 275 Contribution.
Societe Entomologique de France, Annales, 6, 581-584.

Schedl, K.E. (1972) Neotropische Scolytoidea, X1. 293 Beitrag. Koleopterologische
Rundschau, 50, 37-86.

Schedl, K.E. (1973) Neotropische Scolytoidea, XII. 295 Contribution. Papéis Avulsos de
Zoologia, Sao Paulo, 26, 149-172.

Schedl, K.E. (1976) Zur Synonymie der Borkenkafer XXVII. Zeitschriﬁ der
Arbeitsgemeinschaft 0sterrich Entomologen, 28, 68-72.

Six, D.L. (2003) Bark beetle-fungus symbioses. Insect Symbiosis (ed. K. Bourtzis and T.
A Miller), pp. 99-116. CRC Press. New York.

Sorenson, MD. (1999) TreeRot, version 2. Boston University, Boston, MA.

Swofford, D.L. (2001) PAUP“: Phylogenetic Analysis Using Parsimony (*and
Other Methods), 4.0b4a. Sinauer Associates. Sunderland, Massachusetts.

de la Torre-Bueno, J. R. The Torre-Bueno glossary of entomology. 1989. New York
Entomological Society and American Museum of Natural History New York, NY.

Viana, M.J. (1965) Datos ecologicaos de Scolytidae Argentinos (Coleoptera). Sociedad
Entomologica Argentina, Revista, 27, 119-130.

Wheeler, Q. D. & Platnick, NI. (2000) The phylogenetic species concept (sensu Wheeler and
Platnick). Species concepts and phylogenetic theory (ed. by Q. D. Wheeler and R.

244

Meier), pp. 55-69. Columbia University Press, New York.

von Winning, E. (1930) ReifeﬁaB von rindenbriitenden borkenkafern (Ipiden) an
blattern. Anzeigerﬁtr Schadlingskunde, 6, 42-45.

Wood, S.L. (1961) A key to the North American genera of Scolytidae. Coleopterists
Bulletin, 15, 4148.

Wood, S.L. (1969) New records and species of neotropical bark beetles (Scolytidae:
Coleoptera), Part IV. Brigham Young University Science Bulletin, Biological
Series, 10, 1-46.

Wood, S.L. (1972) Notes on the classiﬁcation of the tribe Scolytini (Coleoptera,
Scolytidae). Bulletin of Entomological Research, 62, 243-246.

Wood, S.L. (1978) A reclassiﬁcation of the subfamilies and tribes of Scolytidae
(Coleoptera). Annals of the Societe Entomologique de France, 14, 95-122.

Wood, S.L. (1982) The bark and ambrosia beetles of North and Central America
(Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist
Memoirs, 6, 1359 pp.

Wood, S.L. (1986) A reclassiﬁcation of the genera of Scolytidae (Coleoptera). Great
Basin Naturalist Memoirs, 10, 126 pp.

Wood, S.L. (2007) Bark and ambrosia beetles of South America (Coleoptera, Scolytidae)
Western North American Naturalist, 900 pp.

Wood, S.L. & Bright, DE. (1992) A catalog of Scolytidae and Platypodidae
(Coleoptera), Part 2: Taxonomic Index. Great Basin Naturalist Memoirs, 13,
1553 pp.

Yoder, M.J., Dole, K, Deans, A (2006). Introducing 'mx', a sharable digital workbench

for systematic biologists. Taxonomic Databases Working Group 2006 Annual
meeting. Poster.

245

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