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5/08 KthroleccspresIClRC/DaIeDue,Indd

BIODIVERSITY OF TROPICAL SCOLYTINAE (COLEOPTERA:
CURCULIONIDAE)

By

Stephanie Alexandra Dole

A DISSERTATION

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

DOCTOR OF PHILOSPHY
Entomology

2008

ABSTRACT

BIODIVERSITY OF TROPICAL SCOLYTINAE (COLEOPTERA:
CURCULIONIDAE)

By

Stephanie Alexandra Dole

A phylogenetic revision of the Xyleborina genus Xylosandrus Reitter based on
morphological and molecular data sets is presented. The monophyly of the genus was
tested using a 43 character morphological data set analyzed separately and in
combination with a molecular data set comprised of five independent gene loci: 28S
rDNA; the mitochondrial gene cytochrome oxidase I (C01); and the nuclear genes
arginine kinase (ArgK), rudimentary (CAD), and Elongation Factor l-a (EF-l a). The
nuclear protein-coding genes CAD and ArgK were used for the first time in
phylogenetics of Scolytinae. Both genes demonstrated phylogenetic utility across
varying nodal depths. Phylogenetic analyses were performed using Parsimony and
Bayesian optimality criteria. Xylosandrus was recovered as polyphyletic with the present
classification containing species from four different genera: Amasa Lea, Anisandrus
Ferrari, Cnestus Sampson, and Xylosandrus. Several species of Xylosandrus were
consistently placed in clades with the genera Anisandrus and C nestus with high support
values (100% bootstrap support). Among these, was the economically important invasive
species X. mutilatus, which was consistently recovered as part of the “C nestus” clade. A
taxonomic revision of X ylosandrus is presented based on these results. The following
new combinations are given: Amasa cylindrotomicus (Schedl), A. omissus (Schedl), A.

oralis (Schedl), Anisandrus butamali (Beeson), A. ursa (Eggers), A. ursinus (Hagedom).

A. ursulus (Eggers), Cnestus ater (Eggers), C. fijianus (Schedl), C. gravidus (Blandford),
C. improcerus (Sampson), C. laticeps (Wood), C. mutilatus (Blandford), C. orbiculatus
(Schedl), C. peruanus (Wood), C. retifer (Wood), C. retusus (Eichhoff), C. testudo
(Eggers), C yclorhipidion squamulatus (Beaver and Loyttyniemi), X ylosandrus amputalus
(Blandford), X. mixtus (Schedl), and X. rotundicollis (Browne). Two new species of
Xylosandrus are described: X. borneensis and X. hulcri. An illustrated key to worldwide
species of Xylosandrus is provided. Biogeography, host plants, diagnosis, and images are
presented for each species.

Canopy fogging was used to sample the diversity of bark and ambrosia beetles
(Coleoptera: Curculionidae: Scolytinae) at two Western Amazonian rain forest sites in
Ecuador. Sampling spanning from 1994-2006 yielded 1,158 samples containing 2,500
scolytine specimens representing more than 400 morphospecies. Here, we analyze a
subset of this data representing two ecological groups: true bark beetles (52
morphospecies) and ambrosia beetles (69 morphospecies). A high percentage of these
taxa occurred as singletons and doubletons and their species accumulation curves did not
reach an asymptote. Diversity estimates placed the total scolytine species richness for
this taxon subset present at the two sites between 260 and 323 species. High levels of Ot-
and B-diversity were discovered. However, high B-diversity was found to be an artifact
of undersampling and does not appear to be biologically significant. This study
demonstrates the utility of canopy fogging methods for the sampling of scolytine richness

and for the discovery of new scolytine taxa.

This work is dedicated to my husband, Krishna Peter Dole, who offered constant support
during the Ph.D. process and made great sacrifices so that I could achieve my
entomological dreams.

iv

ACKNOWLEDGMENTS

This work would not have been possible without the generous support and
continued guidance of my major professor, Dr. Anthony Cognato. I would also like to
thank my Ph.D. committee members Dr. Richard Merritt, Dr. Therese Poland, Dr. Alan
Prather, and Dr. Richard Triemer for their guidance and review. This work was also
made possible by the support and advice given by members of the Holistic Insect
Systematics Laboratory: Jiri Hulcr, Aaron Smith, and Sarah Smith. Dr. Roger Beaver
proved to be an exceptional colleague through his dedication to advising me on this work
and his valuable comments on the revision of X ylosandrus. I would like to thank Dr. Don
Bright, Dr. Bjarte Jordal, Dr. Robert Rabaglia, Dr. Saowapa Sonthichai, and Dr. Stephen
Wood for their assistance with this research. I thank Dr. Terry Erwin and Dr. Warren
Steiner of the Smithsonian Institute for enabling me to collaborate on their canopy
fogging research. I would also like to thank the California Academy of Sciences
Department of Entomology for the use of their research facilities and the generous
support of their researchers and staff. With tremendous gratitude, I thank my parents. F.
Clark and Gabrielle Stephens, for the opportunities they provided me throughout my life
that made this achievement possible. This research was supported by NSF-PEET grant

(DEB-0328920) to Anthony 1. Cognato.

TABLE OF CONTENTS

LIST OF TABLES ................................................................................. vii
LIST OF FIGURES ................................................................................ ix
CHAPTER 1

POLYPHYLY OF XYLOSANDR US REITTER INFERRED FROM NUCLEAR AND
MITOCHONDRIAL GENES (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE:

XYLEBORINA) ..................................................................................... I
Abstract ...................................................................................... 2
Introduction .................................................................................. 3
Materials and Methods ..................................................................... 8
Results ....................................................................................... 14
Discussion .................................................................................. 16

CHAPTER 2

PHYLOGENETIC REVISION OF XYLOSANDRUS REITTER (COLEOPTERA:

CURCULIONIDAE: SCOLYTINAE: XYLEBORINA) ..................................... 45
Abstract ..................................................................................... 46
Introduction ................................................................................. 47
Materials and Methods .................................................................... 55
Results ....................................................................................... 66
Discussion .................................................................................. 68

CHAPTER 3

DIVERSITY OF SCOLYTINAE (COLEOPTERA: CURCULIONIDAE) IN THE

ECUADORIAN RAIN FOREST CANOPY ................................................. 254
Abstract ................................................................................... 255
Introduction ............................................................................... 256
Materials and Methods .................................................................. 260
Results and Discussion ................................................................. 266

LITERATURE CITED .......................................................................... 286

vi

LIST OF TABLES

Table 1.1: Specimens sequenced, collecting data, and hosts. .............................. 25

Table 1.2: PCR primers and annealing temperatures used for the amplification of gene
sequences. ........................................................................................... 28

Table 1.3: Support found for selected clades found by different analyses of the five gene
dataset. Numbers represent bootstrap/Bremer supports or posterior probabilities for
Bayesian analyses. NA = not applicable or not found in the resulting tree(s). ............ 29

Table 1.4: Contribution of data partitions to the character matrix and to the resolution of
the most parsimonious tree found in analysis 1. ............................................... 30

Table 1.5: Partition branch support based on tree found by phylogenetic analysis 1. (see
Materials and Methods). Nodes refer to Figure 1.1. .......................................... 31

Table 1.6: Partition branch support based on tree found by phylogenetic analysis 2 (see
Materials and Methods). Nodes refer to Figure 1.2. .......................................... 32

Table 1.7: Partition branch support based on tree found by phylogenetic analysis 3 (see
Materials and Methods). Nodes refer to Figure 1.3. .......................................... 33

Table 1.8: Partition branch support based on tree found by phylogenetic analysis 4 (see
Materials and Methods). Nodes refer to Figure 1.4. .......................................... 34

Table 1.9: COI intra- and interspecific distances expressed as the proportion of sites
differing between sequences, with interspecific distances given as the average observed

between species. .................................................................................... 35

Table 2.1: Biogeographic distribution of Xylosandrus species: 0 = present; A = present
as introduced species. ................................................................................................. 196

Table 2.2: Morphological character matrix of 43 characters for 73 species. Characters
are their scores described in Materials and Methods. ........................................ 197

Table 2.3: Species data for morphological and molecular characters martricies ( o = data
avaiable for species). ............................................................................. 205

Table 2.4: Contribution of data partitions to the data matrix and to the resolution of the
most parsimonious tree found in the analysis of the combined data set. .................. 209

Table 2.5: Partition branch support for the most parsimonious tree found by the analysis
of the combined data set. Node numbers refer to Figure 2.2. .............................. 210

vii

Table 3.1: Species data, Complementarity Indices (CI) and richness estimators for (A)
total data analyzed, (B) bark beetles (Hylesinini), and (C) ambrosia beetles (Xyleborina +
Premnobina). ...................................................................................... 273

Table 3.2: Morphospecies of bark beetles (Hylesinini) found in the canopy fogging
samples. ............................................................................................ 274

Table 3.3: Morphospecies of ambrosia beetles (Xyleborina + Premnobina) found in the
canopy fogging samples. ........................................................................ 276

Table 3.4: Previous South American generic and species records for various scolytine
subtribes compared with taxa collected by this study. ....................................... 278

Table 3.5: Species numbers and Complementarity Indicies (CI) for analyzed subtribes of
Scolytinae. ......................................................................................... 279

viii

LIST OF FIGURES

Figure 1.1: Most parsimonious tree found by parsimony analysis of data aligned using
28S secondary structure (Jordal et al. 2008), with gaps treated as missing data. Numbers
above nodes are Bremer support values. Numbers below nodes are bootstrap support
values. ................................................................................................ 36

Figure 1.2: One of three most parsimonious trees found by parsimony analysis of data
aligned using 28S secondary structure (Jordal et al. 2008), with gaps treated as a 51"
character state. Clade numbers are given above nodes. Numbers below nodes are
bootstrap support values. .......................................................................... 37

Figure 1.3: Single most parsimonious tree found by parsimony analysis of data aligned
in MUSCLE, with gaps treated as missing data. C lade numbers are given above nodes.
Numbers below nodes are bootstrap support values. .......................................... 38

Figure 1.4: One of two most parsimonious trees found by parsimony analysis of data
aligned in MUSCLE, with gaps treated as a 5"1 character state. Clade numbers are given
above nodes. Numbers below nodes are bootstrap support values. ........................ 39

Figure 1.5: One of two post parsimonious trees found by dynamic alignment and
analysis using POY, with a gap cost = 2. Numbers above nodes are Bremer support
values. ............................................................................................... 40

Figure 1.6: Bayesian tree found by analysis of data aligned using 288 secondary
structure (Jordal et al. 2008). Numbers above nodes are Bayesian posterior probabilities.
......................................................................................................... 41

Figure 1.7: Bayesian tree found by analysis of data aligned in MUSCLE. Numbers
above nodes are Bayesian posterior probabilities. ............................................. 42

Figure 1.8: Relationships between PBS values and nodal distance (maximum likelihood
branch length from a given node to the tip of the tree) for the gene partitions: 28S, C01,
and EF-1_ (A), CAD and ArgK (B). ............................................................. 43

Figure 2.1: Strict consensus of 10,000 most parsimonious trees found in parsimony
analysis of the morphological data martrix. Bootstrap values are given below the nodes
for clades with support 2 90%. ................................................................. 211

Figure 2.2: Most parsimonious tree found in the analysis of the combined data matrix.
Node numbers given above branches and bootstrap support/Bremer support values given
below. .............................................................................................. 212

Figure 2.3: Lateral habitus of Amasa cylindrotomicus n. comb., female (A) and Amasa
bicostatus, female (B). ........................................................................... 213

ix

Figure 2.4: Lateral habitus of Anisandrus ursulus n. comb., female (A) and Anisandrus
sayi, female (B). ................................................................................... 214

Figure 2.5: Lateral habitus of Cnestus improcerus n. comb., female (A) and Cnestus
pseudosuturalis, female (B). .................................................................... 215

Figure 2.6: Lateral (A) and dorsal (B) views of Xylosandrus abruptulus. 1.9 — 2.1 mm.
female lectotype. .................................................................................. 2 1 6

Figure 2.7: Lateral (A) and dorsal (B) views of Xylosandrus adherescens, 2.0 mm.
female holotype. ................................................................................... 217

Figure 2.8: Lateral (A) and dorsal (B) views of X ylosandrus amputatus n. comb., 2.7 —
2.9 mm, female. .................................................................................. 218

Figure 2.9: Lateral (A) and dorsal (B) views of Xylosandrus arquatus, 2.3 — 2.5 mm,
female. .............................................................................................. 219

Figure 2.10: Lateral (A) and dorsal (B) views of Xylosandrus assequens, 1.6 — 2.3 mm,
female, holotype. ................................................................................. 220

Figure 2.11: Lateral (A) and dorsal (B) views of Xylosandrus borealis. 2.0 — 2.1 mm.
female. .............................................................................................. 221

Figure 2.12: Lateral (A) and dorsal (B) views of Xylosandrus borneensis n. sp., 1.5 - 1.6
mm, female holotype. ........................................................................... 222

Figure 2.13: Lateral (A) and dorsal (B) views of Xylosandrus brevis, 2.5 — 2.8 mm.
female. .............................................................................................. 223

Figure 2.14: Lateral (A) and dorsal (B) views of Xylosandrus compactus, 1.4 — 1.9 mm,
female. .............................................................................................. 224

Figure 2.15: Lateral (A) and dorsal (B) views of X ylosandrus corthyloides, 2.7 — 3.0
mm, female lectotype. ............................................................................ 225

Figure 2.16: Lateral (A) and dorsal (B) views of Xylosandrus crassiusculus, 1.7 — 2.9
mm, female. ....................................................................................... 226

Figure 2.17: Lateral (A) and dorsal (B) views of X ylosandrus curtulus, 1.3 — 1.5 mm,
female holotype of synonym X. biseriatus. ................................................... 227

Figure 2.18: Lateral (A) and dorsal (B) views of X ylosandrus derupteterminatus, 2.0 -
2.3 mm, female holotype. ....................................................................... 228

Figure 2.19: Lateral (A) and dorsal (B) views of Xylosandrus deruptulus. 1.8 mm.
female lectotype. .................................................................................. 229

Figure 2.20: Lateral (A) and dorsal (B) views onylosandrus discolor, 1.8 — 2.0 mm,
female. .............................................................................................. 230

Figure 2.21: Lateral (A) and dorsal (B) views of X ylosandrus diversepilosus, 1.8 - 2.3
mm, female holotype. ............................................................................. 231

Figure 2.22: Lateral (A) and dorsal (B) views of X ylosandrus eupatorii, 1.8 —— 2.1 mm,

female cotype. ..................................................................................... 232
Figure 2.23: Lateral (A) and dorsal (B) views of Xylosandrusferinus, 1.6 — 1.8 mm,
female lectotype. .................................................................................. 233
Figure 2.24: Lateral (A) and dorsal (B) views of Xylosandrus germanus. 1.9 — 2.5 mm.
female. .............................................................................................. 234
Figure 2.25: Lateral (A) and dorsal (B) views of X ylosandrus hirsutipennis. 1.9 — 2.2
mm, female paratype. ............................................................................. 235
Figure 2.26: Lateral (A) and dorsal (B) views of Xylosandrus hulcri n. sp., 2.4 — 2.7mm.
female holotype. .................................................................................. 236
Figure 2.27: Lateral (A) and dorsal (B) views of Xylosandrus jaintianus. 3.0 mm. female
holotype. ........................................................................................... 237
Figure 2.28: Lateral (A) and dorsal (B) views of Xylosandrus mancus, 2.9 - 3.3 mm,
female. .............................................................................................. 238
Figure 2.29: Lateral (A) and dorsal (B) views of X onsandrus mediocris, 1.4 mm, female.
....................................................................................................... 239
Figure 2.30: Lateral (A) and dorsal (B) views of Xylosandrus mesuae, 1.1 — 1.3 mm,
female. .............................................................................................. 240
Figure 2.31: Lateral (A) and dorsal (B) views of X ylosandrus metagermanus, 1.8 mm,
female holotype. ................................................................................... 241
Figure 2.32: Lateral (A) and dorsal (B) views of Xylosandrus mixtus n. comb.. 2.6 — 2.7
mm, female holotype. ............................................................................ 242
xi

Figure 2.33: Lateral (A) and dorsal (B) views of Xylosandrus monteithi, 3.0 — 3.4 mm.
female paratype. .................................................................................. 243

Figure 2.34: Lateral (C) and dorsal (D) views of Xylosandrus monteithi, 2.5 mm. male
allotype. . ........................................................................................... 244

Figure 2.35: Lateral (A) and dorsal (B) views of X ylosandrus morigerus. 1.5 — 2.0 mm,
female. .............................................................................................. 245

Figure 2.36: Lateral (A) and dorsal (B) views of Xylosandrus pusillus, 1.5 - 1.7 mm.
female holotype. ................................................................................... 246

Figure 2.37: Lateral (A) and dorsal (B) views of Xylosandrus pygmaeus, 1.3 — 1.4 mm,
female holotype. ................................................................................... 247

Figure 2.38: Lateral (A) and dorsal (B) views of Xylosandrus queenslandi, 1.6 — 1.9 mm,
female paratype. .................................................................................. 248

Figure 2.39: Lateral (A) and dorsal (B) views of X ylosandrus rotundicollis n. comb., 3.7
- 4.1 mm, female holotype. ..................................................................... 249

Figure 2.40: Lateral (A) and dorsal (B) views of Xylosandrus subsimiliformis, 2.8 mm,
female cotype. .................................................................................... 250

Figure 2.41: Lateral (A) and dorsal (B) views of Xylosandrus subsimilis, 2.6 mm, female
cotype. ............................................................................................. 251

Figure 2.42: Lateral (A) and dorsal (B) views of Xylosandrus terminatus, 1.5 — 1.9 mm,
female cotype. ..................................................................................... 252

Figure 2.43: Lateral (A) and dorsal (B) views of Xylosandrus woodi. 2.3 -— 2.4 mm,
female paratype. ................................................................................... 253

Figure 3.1: Species accumulation curves for Onkone Gare + Tiputini for total data
(Hylesinini + Xyleborina + Premnobina). .................................................... 280

Figure 3.2: Species accumulation curves for Onkone Gare + Tiputini for bark beetles
(Hylesinini). ....................................................................................... 281

Figure 3.3: Species accumulation curves for Onkone Gare + Tiputini for ambrosia
beetles (Xyleborina + Premnobina). ........................................................... 282

Figure 3.4: Richness estimators for Onkone Gare + Tiputini for total data (Hylesinini +
Xyleborina + Premnobina). ...................................................................... 283

xii

 

Figure 3.5: Richness estimators for Onkone Gare + Tiputini for bark beetles (Hylesinini).
........................................................................................................ 284

Figure 3.6: Richness estimators for Onkone Gare + Tiputini for ambrosia beetles
(Xyleborina + Premnobina). .................................................................... 285

xiii

CHAPTER 1

Polyphyly of Xylosandrus Reitter inferred from nuclear and mitochondrial genes
(Coleoptera: Curculionidae: Scolytinae)

ABSTRACT

The monophyly of the Xyleborina ambrosia beetle genus X ylosandrus was tested
using a data set comprised of multiple gene loci: 28S rDNA; the mitochondrial gene
cytochrome oxidase I (C01); and the nuclear genes arginine kinase (ArgK), rudimentary
(CAD), and Elongation Factor I-a (EF-l or). The nuclear protein-coding genes CAD and
ArgK were used for the first time in phylogenetics of Scolytinae. Analyses were
performed using Parsimony and Bayesian optimality criteria. X ylosandrus is a
widespread genus containing 54 species, several of which are of economic importance.
Our analyses included 43 specimens representing 15 X ylosandrus species and 19 species
from Amasa, A nisandrus, C nestus, and Xyleborus, and two species from the outgroup
genus Coccotrypes. All analyses recovered a polyphyletic X ylosandrus. Several species
of Xylosandrus were consistently placed in clades with the genera A nisandrus and
Cnestus with high support values (100% bootstrap support). Among these, was the
economically important invasive species X. mutilatus, which was consistently recovered
as part of the “Cnestus” clade. In our analyses, both CAD and ArgK demonstrated
phylogenetic utility across varying nodal depths. Despite the selection of genes with
signals at complementary phylogenetic depths. the data set used herein did not resolve the
phylogeny of Xylosandrus and related genera. Since the taxon sample available for
molecular work represents only a fraction of X ylosandrus species, it is recommended that
the genus be the subject of a complete revision that combines molecular and

morphological data in total evidence approach.

INTRODUCTION

The beetle subfamily Scolytinae (Coleoptera: Curculionidae) is comprised of 26
tribes containing approximately 225 genera and 6,000 species worldwide (Wood and
Bright 1992; Mandelshtam and Beaver 2003). Due to the present classification of
Scolytinae as a subfamily of the Curculionidae (Marvaldi 1997; Kuschel et al. 2000)
previously recognized tribes (e. g. Xyleborini) are currently treated as subtribes (e.g.
Xyleborina). Although the Curculionoidea are a relatively young group (152 MYO),
discovery of a fossilized scolytine from Cretaceous amber dates the origin of the
subfamily to at least 100 million years ago (Grimaldi and Engel 2005; Cognato and
Grimaldi in press). This long history likely contributes to the diversity of Scolytines
which occupy two ecological groups: bark beetles and ambrosia beetles. Bark beetles
bore into the phloem of trees and feed on tree tissues (Wood and Bright 1992). Ambrosia
beetles bore into the xylem and feed on symbiotic fungi, which grows on the walls of
their galleries (Beaver 1989; Wood and Bright 1992; Kirkendall 1993). This ambrosia]
habit has evolved multiple times within the Scolytinae and also in the related weevil
subfamily Platypodinae (Farrell et a1. 2001). The scolytine subtribe Xyleborina contains
approximately 1,300 described species and constitutes one of the largest radiations of
ambrosia beetles (Wood and Bright 1992; Jordal 2002). Xyleborina are absent in the
Dominican amber fossil record, suggesting that their radiation began in the Miocene
(Bright and Poinar 1994; Jordal et al. 2000). The xyleborine mating system, which
includes haplodiploidy and extreme inbreeding, is believed to be the cause of this

dramatic radiation (Normark et al. 1999). Ambrosia beetles tend to be less host-specific

 

than their bark beetle counterparts, because adaptation to host secondary chemistry has
not influenced their radiation. Hence they are particularly suited for the invasion of new
habitats and establishment as introduced exotic species. Ten species of exotic xyleborine
beetles became established in the United States between 1985 and 2005 (Haack 2006:
Rabaglia et al. 2006). These invasions, along with the interesting biology and ecology of
xyleborine beetles, have prompted much research interest. Historically, the classification
of the Xyleborina has been chaotic, with Wood (1986) describing his own classification
of the tribe as “tentative and flawed.” However, recent taxonomic work has begun to
correct this, as the subtribe is finally being studied within a phylogenetic context (Jordal
et al. 2000; Jordal 2002; Hulcr et al. 2007).

Xylosandrus Reitter (1913) is a large genus of xyleborine ambrosia beetles with a
widespread distribution primarily in tropical, but also in temperate regions of the world.
In their worldwide catalog of the Scolytinae, Wood and Bright (1992) list 52 species of
Xylosandrus. Subsequent descriptions, new synonymies, and new combinations have
brought the present number to 54 species (Bright and Skidmore 1997; Saha et al. 2002;
Wood 2007; Dole and Beaver in press).

Several Xylosandrus species cause economic losses in nursery and agricultural
settings in their native and introduced ranges. In Brazil, X compactus causes losses in
several economically important host species, including avocado, cacao, coffee, and
mango (Oliveira 2008). In North America, three Xylosandrus species (X. compactus, X.
crassiusculus, and X germanus) have caused “considerable economic damages” since
their introductions (Oliver and Mannion 2001). As an example, nursery managers in

Maryland reported individual losses of $3,650 - $8,400 in stock to Xylosandrus species in

the spring of 2008 alone (R. J. Rabaglia per. mm.) A revised understanding of
Xylosandrus generic boundaries and species diagnosis would aid in the identification and
control of these economically important beetles.

Xylosandrus currently contains species with highly variable morphologies, several
of which are similar to those of other genera, leading to confusion concerning the generic
boundaries of X ylosandrus and its relationship to and distinction from the genera Amasa
Lea, Anisandrus Ferrari, and Cnestus Sampson. The underlying cause of this confusion
has been the incorrect placement of species within X ylosandrus, which has blurred the
boundaries between it and other genera (Dole and Beaver in press). Commonly cited
diagnostic characters for Xylosandrus include widely separated procoxae; very stout,
cylindrical bodies; and antennal club that is obliquely truncate with the first segment of
the antennal club forming a circular costa and with segments two and three not visible on
posterior face of the club (Bright 1968; Wood 1986; Hulcr et al. 2007; Dole and Beaver
in press). However, even the frequently cited diagnostic character of widely separated
procoxae does not function as a synapomorphy for X ylosandrus and the genus is
presently defined by a collection of homoplastic xyleborine characters (Hulcr et al. 2007).

Both morphological (Dole and Beaver in press) and molecular data suggest that
Xylosandrus is not monophyletic (Jordal 2002). A cladistic review of the generic
taxonomic characters of Xyleborina recovered a monophyletic X ylosandrus, but it is
important to note that this study did not include species with morphologies that deviate
from the sensu stricto concept of the genus (Hulcr et al. 2007). In their review of
Australian Xylosandrus, Dole and Beaver (in press) contributed to the revision of

X ylosandrus by defining characters that distinguish it from C neslus and moving two

species from Xylosandrus to Cnestus. This was only a beginning, given the narrow
geographic perspective of their review. Xylosandrus is in need of a comprehensive
phylogenetic revision that includes other suspect genera, such as A masa, A nisandrus, and
Cnestus. The prevalence of homoplastic morphological characters within X ylosandrus
and related genera necessitates the inclusion of alternative characters, such as DNA. to
help reconstruct the phylogeny of this group.

Molecular studies of scolytine phylogenetics have been based mainly on
mitochondrial genes (168, C01), the ribosomal encoding gene 28S, and the nuclear
protein encoding genes elongation factor 1 a (EF-Ioc) and Enolase (Normark et a1. 1999;
Jordal et a1. 2000; Cognato and Vogler 2001; Farrell et a1. 2001; Jordal 2002; Jordal and
Hewitt 2004; Jordal et al. 2008). Of these genes, EF-la has shown the greatest potential
for resolving the phylogenetic relationship of Scolytinae (Jordal 2002, 2007). However,
use of these genes alone has not resolved a Scolytinae phylogeny. Recent studies have
indicated that the use of multiple independent gene loci may be necessary to recover
resolved phylogenies (Rokas and Carroll 2005; Edwards et a1. 2007; Wild and Maddison
2008). The thousands of protein-coding genes present in the nuclear genomes of
eukaryotes offer a rich source of new loci to include in phylogenetic studies of arthropods
(Wiegmann et a1. 2000; Jordal 2007; Wild and Maddison 2008). Protein-coding nuclear
genes have been used in the inference of Lepidoptera phylogenetics for over a decade
(Friedlander et al. 1998; Regier et a1. 1998; Fang et al. 2000; Friedlander et al. 2000;
Mitchell et a1. 2000; Wiegmann et al. 2000). They have also proven useful in the
reconstruction of higher-level relationships among the Arthropods (Schultz and Regier

2000a; Schultz and Regier 2000b; Regier and Schultz 2001). Nuclear protein-coding

genes are being used in Coleoptera phylogenetics, but a recent literature survey found
only 24 studies of beetle molecular phylogenetics (out of a total of 106) that utilized them
(Wild and Maddison 2008). Advantages of nuclear protein-coding genes include slower
rates of evolution, a lower tendency toward base-composition bias. and ease of alignment
(Wiegmann et al. 2000; Wild and Maddison 2008).

Recognizing the underutilized potential of nuclear protein-coding genes, Jordal
(2007) screened multiple gene loci for the phylogenetic reconstruction of scolytines. The
nuclear genes CAD, also known as rudimentary, and arginine kinase (ArgK) were found
to be the most promising and were recommended for use in future phylogenetic studies of
Scolytinae. The CAD gene codes for several enzymes involved in the de novo synthesis
of pyrimidines. In Insecta, the carbamolyphosphate synthetase locus of CAD has been
used in phylogenetic studies of Diptera (Moulton and Wiegmann 2004) and Hymenoptera
(Danforth et al. 2006). The ArgK gene encodes for a metabolic phosphostransferase
enzyme that aids in the maintenance of ATP levels. In Insecta, ArgK has been used in
phylogenetic studies of Hymenoptera (Kwakita et al. 2003; Banks and Whitfield 2006).

Wild and Maddison (2008) empirically tested the utility of CAD and ArgK, along
with other nuclear protein-coding genes, for the inference of beetle phylogenies against a
presumably known phylogeny of Coleoptera which spanned several taxonomic levels.
Within this study, CAD was the highest performing gene fragment of the eight tested,
because it recovered the greatest number of clades of the known phylogeny and had high
support values. ArgK reconstructed deeper clades of the known phylogeny with more
accuracy than shallower clades. Also, there is no evidence for the existence of multiple

COpies of CAD and ArgK within beetle genomes (Jordal 2007; Wild and Maddison

2008). Thus, the inclusion DNA from both genes in a phylogenetic analysis of
Xylosandrus could help resolve multiple levels of the resulting phylogeny.

Here, we use the CAD and ArgK genes for the first time for phylogenetic
inference in the Scolytinae. Using a data set comprised of multiple gene loci (28S, ArgK,
CAD, C01, and EF-la) we construct a phylogenetic hypothesis of the species
relationships within Xylosandrus, test the monophyly of the genus, and test the

relationships among X ylosandrus and the genera Amasa, Anisandrus, and C nestus.

MATERIALS AND METHODS

T axa, DNA sequences, and alignment

The majority of Xylosandrus species are relatively difficult to collect, thus it was
a challenge to include a broad taxon sampling in this study. Specimens for DNA
extraction were either collected by the authors in Ecuador, Thailand, Papua New Guinea,
and Borneo or obtained from colleagues. In nearly all cases, specimens were stored in
95% ethanol at — 80° C. However, DNA was successfully extracted from several
specimens that had been killed in 95% ethanol, but then pinned and stored dried (X.
monteithi andX. queenslandi). We included 43 specimens (Table 1.1) representing 15
X ylosandrus species 19 species from the genera Amasa, Anisandrus, C nestus, X yleborus,
and two species from the outgroup genus Coccotrypes. Species from a diverse group of
ge nera were chosen in order to test the generic limits of X ylosandrus.

DNA was extracted from dissected throacies following protocols of Cognato and

Vogler (2001) and the remaining body parts were pinned and vouchered at the A]. Cook

Arthropod Research Collection, Michigan State University. Using the purified DNA.
partial gene regions of mitochondrial cytochrome oxidase 1, nuclear ribosomal 28S (D2
and D3 regions), elongation factor-1a, CAD (rudimentary), and Arginine Kinase (ArgK)
were amplified with the listed PCR primers (Table 1.2). All PCR reactions first included
published COI, EF-I or and 28S primers (Normark et al. 1999; Hebert et al. 2003; Jordal
et al. 2008) and additional xyleborine specific primers were designed when PCR failed.
CAD and ArgK sequences were first generated with degenerate primers designed based
on bee sequences (Danforth et. a1 2006; Jordal 2007). We used these sequences to
subsequently create new xyleborine primers (Table 1.2). Each COI, EF-Ia and 28S PC R
reaction contained: 29.5 pl ddeO, 10 p1 5x TanNA polymerase buffer (Promega,
Madison, WI), 5 pl 25mM Promega MgClz, 1 pl 40 mM deoxynucleotide triphosphates
(dNTPs), 0.5 pl of 5 units/ pl Promega Taq DNA polymerase, 10 picomoles/ pl of forward
and reverse primers and 2 pl of DNA template (5—50 nanograms). Each CAD and ArgK
PCR reactions contained: 17.75 pl ddHZO, 2.5 pl 10x Taq DNA polymerase buffer
(Qiagen Inc., Santa Clara, CA), 1 pl 25mM MgClz, 0.5 pl 40 mM deoxynucleotide
triphosphates (dNTPs), 0.25 pl of 5 units/ pl Qiagen Hotstar Taq DNA polymerase, 5
picomoles/ pl of forward and reverse primers and 2 pl of DNA template (5-50
nanograms). PCR was performed on a thermal cycler (MJ Research, Waltham, MA)
under the following conditions: one cycle for 2 minutes at 95°C (15 min. for Hotstar
Taq), 34 cycles for 1 min. at 95°C, 0.75 min. at 45-55°C (see Table 1.2 for specific
annealing temperatures), 1 min. at 72°C, and a final elongation cycle of 5 min. at 72°C.
PCR products were visualized via electrophoresis. For each reaction, 5.0 pl of

PCR product was mixed with 2.00 pl 5X loading buffer (Promega. Madison. WI). The 7

pl mixture was applied to an ethidium bromide stained 1.5% agarose gel at 100 volts for
30 minutes. The gel was visualized under an ultraviolet light source and photographed.
Unincorporated oligonucleotides and T aq were destroyed in each reaction with ExoSAP-
IT following the manufacturer’s protocols (USB Corp., Cleveland, OH). The clean PC R
reactions were directly sequenced using BigDye® Terminator v.1.1 (Applied Biosystems.
Foster City, CA) cycle sequencing kit and visualized on an ABI 3730 or 3700 (Applied
Biosystems, Foster City, CA) at the Research Technology Support Facility at Michigan
State University. Primers used in PCR were used to sequence sense and antisense strands
for all reactions.

Sense and antisense DNA sequences were compiled with the computer software
Sequencer (Ann Arbor, MI), in order to inspect for ambiguities and create consensus
sequences. No sequence length variation was observed among the protein coding genes
for the included taxa, however considerable variation occurred in the 28S sequences.
These sequences were aligned by two methods; manually with reference to a scolytine-
specific secondary structure model (Jordal et al. 2008) and by a multiple progressive pair-
wise alignment with secondary refinement using the computer software, MUSCLE v.3.52
(Edgar 2004). We did not specify stems and loop regions for the secondary structure
alignment. Instead, we used the scolytid secondary structure model to guide the
alignment by identifying conserved and length variable regions. For the other alignment.
We used the default settings of the web version of MUSCLE
(http://www.ebi.ac.uk/Tools/muscle/index.html). Nexus files of both alignments are

available at http://www.hisl.ent.msu.edu/research/publications.php.

10

 

 

Phylogenetic Analyses
We conducted several analyses, which used different optimality criteria, alignments, and
accounted for potential bias in nucleotide substitution patterns. Parsimony analyses of
data aligned by secondary structure and the MUSCLE software were conducted with the
software PAUP* (Swofford 2002). For all parsimony analyses, most-parsimonious
reconstructions were obtained by a heuristic search with 300 random stepwise addition
replicates using PAUP* default settings. Bootstrap values were determined by
performing 1000 pseudo-replicates with simple sequence addition. Bremer support values
were calculated by constructing constraint trees with the software TreeRot (Sorensen
1 996) followed by subsequent analysis with PAUP. For analyses of both alignments. we
c()nsidered gap positions as missing data and as a 5th character state because previous
Studies have showed that gap positions are phylogenetically informative (Cognato and
V Ogler 2001 ; Lee 2001).
Several researchers have proposed direct optimization of nucleotide homology
ClLlring phylogenetic reconstruction given that static nucleotide alignments may not
accurately reflect positional homology of nucleotides for all taxa (Kruskal 1983; Sankoff
and Cedergren 1983; Wheeler 1996). We performed direct optimization on the data using
the program POY ver. 4.0.2881 (Wheeler 1996; Varén et al. 2008) in order to evaluate
the relative signal within length variable regions. We used the alignment guided by
Secondary structure to divide the 288 data into seven input files, which corresponded to
conserved and length variable regions. The other gene sequences were kept as separate
input files. These files were inputted into POY and analyzed under a parsimony

OI’timality criterion using the following tree search commands:

11

transform(tcm.'(1,2),gap_0pening:1), build (300), swap (threshold:5.0). select 0.
perturb (transform (static_approx), iterations: 1 5, ratchet: (0. 2, 3)), select().
fuse(iterati0ns:200,swapO), selectO, report (trees: (total)). An additional analysis
considering gap cost of 3 was conducted with the same tree search strategy as above.
Bremer support values were calculated for the tree recovered in the first analysis using

the following command: calculate_support(bremer, build(trees:0), swap(tbr, trees.'2)).
Data and command files are available at (http://www.hisl.ent.msu.edu

/research/publications.php).

We analyzed the data under maximum likelihood using Bayesian estimation of

phylogeny as implemented in Mr. Bayes 3.1.2 (Huelsenbeck and

Ronquist 2001) in order to evaluate the effect of nucleotide substitution models on
phylogenetic reconstruction. We conducted a separate analysis for each of the static
alignments. For both analyses, we partitioned the protein coding genes by codon
resulting in 13 partitions thus allowing each to independently evolve under a general time

reversal (GTR) model with a proportion of invariant sites and a gamma distribution. Four

I\’Ietropolis-Coup1ed Markov chain Monte Carlo searches (1 cold, 3 heated) were
performed twice for 20 x 106 generations each with sampling every 100th iteration.
Approach to stationarity (“bum-in”) of each search was determined with the graphical
interface of Tracer v. 1.4 (Rambaut and Drummond 2007) (graph not shown). All
parameters reached stability within 2,000,000 generations and parameters between runs
did not vary. Bayesian posterior probabilities were calculated by a majority-rule

Consensus of those trees after the burn-in (for both runs, 180,000 trees).

12

 

In order to assess the contribution of support of each gene to the total support of the
tree, the dependency of partitioned Bremer support (PBS) and nodal distance (the branch
length from a given node to the tip of the tree) were tested for each gene (Baker et a1.

1998; Cognato and Vogler 2001 ). In PAUP*, maximum likelihood was used to fit branch
lengths to a molecular clock in order to scale branch lengths throughout the most
parsimonious tree found in analysis 1 (Table 1.3). Maximum likelihood estimations were
performed using the following settings: general time-reversible model, base frequencies
determined empirically, proportion of invariable sites estimated, among-site rate variation
approximated to a gamma distribution with 4 rate categories and shape parameter = 0.5,
and molecular clock enforced. Linear regression analyses were then performed on the
nOdal distance versus partitioned Bremer support for each gene using PopTools (Hood

2008).

Intraspecific distances using .1 ukes-Cantor nucleotide substitution model were
Qalculated for each data partition in PAUP* for all taxa with more than one individual
Sequenced (X. compactus, X. crassiusculus, X. germanus, X. morigerus, and Xyleborus
”Otundicollis). Distances for specimens of each of these species were also calculated with
1‘e spect to Xyleborus pelliculosus as a means of comparing intra- and interspecific
distances. The presence of pseudogenes can affect analyses of genetic distance if they
I‘esult in the comparison of non-homologous gene copies (Song et al. 2008). For this
I‘eason, the presence of C01 pseudogenes was ruled out using several methods. First,
Sequence chromatograms were checked for the presence of double peaks. The presence
0f double peaks would indicate the possible amplification of pseudogenes. Next, the COI

Sequences were translated into amino acids using the invertebrate mitochondrial genetic

13

‘————-—1

 

code in the software Mesquite (Maddison and Maddison 2006) and checking for stop
codons in the sequences. The presence of stop codons throughout the amino acid
sequence would indicate that it has been translated from a non-functioning gene copy.
Lastly, nucleotide substitution rates were calculated for each codon position. In
functioning gene copies, there should be a strong substitution rate bias in the third codon

position, whereas this phenomenon may not be observed in non-functioning pseudogene

copies.

RESULTS

All analyses recovered a polyphyletic Xylosandrus (Table 1.3, Figs. 1.1 — 1.7). The
Varied placement of the “Anisandrus” and “C nestus” clades, along with the placement of
Several X onsandrus species within these clades, was responsible for rendering the genus
pelyphyletic. The phylogenetic placement of “Anisandrus” and “C nestus” changed,
depending on the parameters of the analysis performed, and the various placements were
usually weakly supported (e.g. S 50% bootstrap support). The uncertain position of these
C lades made it impossible to determine the phylogenetic relationships among
Xylosandrus, Anisandrus, and C nestus within the context of this study. While the
phylogenetic position of these clades was weakly supported, the placement of several

Species of Xylosandrus within the “A nisandrus” and “C nestus” clades had very strong

SUpport (e. g. 100% bootstrap support).

Homoplasy as measured by Consistency Indices (CI) and Retention Indices (RI)

 

indicated that homoplasy was the lowest for 28S of all the data partitions (Table 1.4).

However, CAD, ArgK, and EFl-a, had similar CI’s (0.450 — 0.486) and RI’s (0.631 —

l4

 

0.677). Homoplasy as measured by node and gene partition provided details of character
congruence. Bremer supports were affected by the alignment method used and whether
gaps were treated as missing data or as a 5th character in the analysis (Tables 1.5 — 1.8).
Overall, 28S gave the highest Bremer support values. Static alignment using MUSCLE
combined with gaps treated as missing data had a dramatic effect on 288 Bremer support
values (Table 1.7). The branch support provided for the various phylogenies by ArgK
was also very high (Tables 1.5 — 1.8). The support provided by CAD varied depending
on the alignment method used for the 28S data. Alignments informed by secondary
Structure produced trees with lower CAD Bremer support values (Tables 1.5 and 1.6).
However, when the alignment was produced by the software MUSCLE, the CAD Bremer
Support values were considerably higher (Tables 1.7 and 1.8). The comparison of PBS
811d nodal distances showed no significant relationship for all of the genes, except for
C01, which was significant at a = 0.05 (r = 0.1170, P = 0.0307) (Figs. 1.8). The COI
data exhibited stronger support for shallower nodes. The phylogenetic information

Obtained from the majority of the genes (EFl-a, 28S, ArgK, CAD) occurs throughout the

phylogeny, regardless of branch length.

Intra- and interspecific distances were expressed as the proportion of sites differing
between sequences, with interspecific distances given as the average observed between
Species (Table 1.9). Intraspecific distances for EF-la, ArgK, CAD, and 28S were
relatively low, ranging from 0 — 1.18% of sites differing between sequences. In
Comparison, the average interspecific distances observed for the five genes in the dataset

rarrged from 3.39 — 11.48%. Overall, sequences of C01 showed high intraspecific

di Stances, ranging from 0 — 13.16%. In comparison, the average interspecific distances

observed for COI ranged from 14.61 — 17.29%. COI sequence chromatograms were
clean and without double peaks, translated COI sequences did not contain stop codons
and there was a strong nucleotide substitution rate bias observed for the third codon

position. Therefore, the possibility of high CO] sequence divergences being artifacts

caused by the amplification of non-homologous pseudogenes was ruled out.

DISCUSSION

Phylogenetics of X ylosandrus

According to these data, the present classification of X ylosandrus contains species
from three different genera: Xylosandrus, Anisandrus, and C nestus. Although tree
topologies differed some, depending on analysis type, alignment parameters, and
tl‘eatment of gaps, the following clades were consistently recovered with high support

Values: “Anisandrus”, “Cnestus”, and (((X. germanus + X. n. sp. Borneo) X. morigerus.

X compactus) (Table 1.3). Membership within these clades is also consistent with
tTlorphological characters that separate other xyleborine genera from Xylosandrus sensu
Stricto, such as widely separated procoxae (see Chapter 2). Three additional clades were
recovered by multiple analyses with varying levels of support, but were not consistently

present on all of the tree topologies: “Cnestus” + X. crassiusculus, X. n. sp. PNG + X.

monteithi, X mancus + X. discolor (Table 1.3).

“Anisandrus” Clade

16

The “Anisandrus” clade consists of four species of Anisandrus (A. dispar, A. hirtus,
A. obesus, and A. sayi) along with two species currently classified as X ylosandrus (X.
ursa and X. ursinus). This clade was recovered by all phylogenetic analyses with high
support values (Table 1.3, Figs. 1.1 — 1.7). Bootstrap support for the “Anisandrus" clade
was 100% in the four analyses for which bootstrap support was calculated. Likewise, the
“A nisandrus” clade was recovered by both Bayesian analyses with a posterior probability

of 1 00. Bremer supports for this clade ranged from 22 to 67.

The genus Anisandrus was recently resurrected in a phylogenetic study of generic
taxonomic characters in the Xyleborina (Hulcr et al. 2007). Morphological characters
distinguishing Anisandrus from Xylosandrus include contiguous procoxae and lateral
protibial margins with 7 — 8 socketed teeth. In Xylosandrus sensu stricto the procoxae

are widely separated and the lateral protibial margins bear 4 — 5 socketed teeth (see
Chapter 2). The two Xylosandrus species included in the “A nisandrus” clade are
morphologically consistent with Anisandrus. This clade is also supported by a molecular
Synapomorphy in 28S in the form of a 9 — 50 base pair insertion comprised almost

exclusively of T’s and Cs.
“Cnestus” Clade

The “Cnestus” clade consists of two species of Cnestus (C. bimaculatus and C.
Pseudosuturalis) along with three species currently classified as Xylosandrus (X. ater, X.
irreprocerus, and X. mutilatus). This clade was also recovered by all phylogenetic
analyses with high support values (Table 1.3, Figs. 1.] — 1.7). Bootstrap support for the
c‘C'nestus” clade was 100% in the four analyses for which bootstrap support was

Cal culated. Likewise, the clade was recovered by both Bayesian analyses with a posterior

17

 

probability of 100. Bremer supports for the “Cnestus” clade were relatively high. ranging

from 53 — 120.

Dole and Beaver (in press) recognized the incorrect placement of Cnestus species
within X ylosandrus and made tentative steps toward correcting these taxonomic errors by
transferring two species from X ylosandrus to C nestus in their review of the Australian
species of Xylosandrus. Morphological characters support the placement of the three
Xylosandrus species within the “C nestus” clade. Characters distinguishing C nestus from
X yl osandrus sensu stricto include subcontiguous procoxae and a four segmented antennal
funicle. In Xylosandrus sensu stricto the antennal funicle is always five segmented.
Additionally, in C nestus the anterior margin of the pronotum bears four or fewer
asperities, with a coarse pair medially, and the pronotum is often produced anteriorly. In
Xylosandrus sensu stricto the anterior margin of the pronotum bears six or more
asperities of approximately equal size and is never produced anteriorly (see Chapter 2).
The two Xylosandrus species placed in the “Cnestus” clade are morphologically

Consistent with Cnestus.

“Xylosandrus sensu stricto” C lade

The clade (((X. germanus + X. n. sp. Borneo) X. morigerus) X. compactus) was
also consistently recovered by all analyses with high support values (Table 1.3, Figs. 1.1
\ 1.7). Given that it contains X. morigerus, the type species of Xylosandrus, this clade is
the highest supported grouping of species belonging to Xylosandrus sensu stricto
resolved by these data. Bootstrap support for this clade was 100% in all analyses for

which bootstrap support was calculated. Likewise, this X ylosandrus sensu stricto clade

18

 

.—

was recovered by both Bayesian analyses with a posterior probability of 100. Bremer
supports for this clade ranged from 21 — 28.
The four species included in the Xylosandrus sensu stricto clade are all
morphologically consistent with the strict definition of the genus. This clade also

contains three economically important species of X ylosandrus: X. compactus, X.

germanus, and X. morigerus.

X ylosandrus sensu lato
Several Xylosandrus species included in this analysis varied in their phylogenetic

placement: X. crassiusculus, X. discolor, X. mancus, X. monteithi, X. queenslandi, andX.
n- sp. Papua New Guinea. These species did not form a larger monophyletic Xylosandrus
Clade along with the Xylosandrus sensu stricto clade. However, none of the various
placements of these species were supported by the phylogenetic analyses. Despite their

Unresolved positions in the molecular phylogeny, the placement of these species in
JCylosandrus may be correct, since their morphologies are consistent with the defining

Characters of Xylosandrus sensu stricto. A phylogenetic analysis of morphological data
Will be necessary to further test the placement of these species within the genus. In
addition, Xyleborus rotundicollis was included in this analysis because its morphology is
Consistent with that of Xylosandrus and not Xyleborus. This species was never recovered

i n a clade with other Xyleborus species and its possible transfer into X ylosandrus should

be examined within the context of a morphological study of the genus.

Contributions of Data Partitions

19

 

Commonly used genes, such as 288, C01, and EF-la, demonstrated similar
phylogenetic utility as compared with other scolytine studies (Normark et al. 1999;
Cognato and Sperling 2000; Jordal et al. 2000; Cognato and Vogler 2001: Farrell et al.
2001; Jordal 2002; Jordal and Hewitt 2004; Cognato et al. 2005; Cognato and Sun 2007:
Jordal 2007; Jordal et al. 2008). However, all genes included in this analysis, except
COI, showed no significant correlation between nodal distance and PBS, indicating that
they had utility for resolving multiple levels of the phylogeny (Figs. 1.8). Typically. the
ribosomal-encoding gene 28S has been shown to resolve relationships within tribes of
Scolytinae, but offers little phylogenetic signal for deeper nodes (Jordal et al. 2008).
Furthermore, alignment guided by secondary structure increases the phylogenetic utility
of 28S (Jordal 2007; Jordal et al. 2008). Here, we found no significant correlation
between nodal distance and PBS for 288, but these results are not unexpected, given that
the relationships being tested were all bellow the tribal level. Mitochondrial genes, such
as COI, typically do not offer much signal for deeper divergences and offer better
resolution for shallower, species-level nodes (Normark et al. 1999; Cognato and Sperling
2000; Cognato and Vogler 2001; Farrell et al. 2001; Jordal 2002; Jordal and Hewitt 2004;
Cognato et al. 2005; Cognato and Sun 2007; Jordal 2007). For COI, there was a
significant correlation between nodal distance and PBS, with higher support for shallower
nodes, as is predicted for mitochondrial genes (Fig. 1.8). Interestingly, this significance
was largely due to the inclusion of sequences from multiple specimens of several species.
With the removal of the data points representing X. crassiusculus, X. germanus, and X.
morigerus the regression analysis found no significant relationship between nodal

distance and PBS. Nuclear genes, such as EF-l a. CAD and ArgK, are more slowly

20

 

evolving and have demonstrated phylogenetic utility across several levels of divergence
mortuark et al. 1999; Jordal et al. 2000; Cognato and Vogler 2001; Farrell et al. 2001;
Jordal and Hewitt 2004; Jordal 2007; Wild and Maddison 2008). In our analyses. both
CAD and ArgK showed phylogenetic utility across varying nodal distances (Figs. 1.8).
Although the genes used herein were chosen for their complementary phylogenetic
signals, they did not resolve the phylogeny of Xylosandrus and related genera. The
unavailability of the majority of X ylosandrus species for DNA sequencing may have
contributed to the lack of phylogenetic resolution. It is difficult to untangle whether the
genes or lack of taxa contributed to the unresolved phylogeny. However, the addition of
both could help to remedy this situtation (Rokas and Carroll 2005; Edwards et al. 2007;

Wild and Maddison 2008).

Genetic Divergence

The results of the comparisons of intra- and interspecific distances for the data
partitions provided a few interesting findings (Table 1.9). Xylosandrus crassiusculus is
the most widespread species of Xylosandrus, with an almost circumtropical distribution.
Specimens of X. crassiusculus from Madagascar and Thailand showed very high COI
sequence divergence from specimens collected in North America (10.09 — 10.94%).
Similarly, X. crassiusculus specimens from Madagascar and Thailand showed a relatively
high level of CO] sequence divergence (4.44%). The intraspecific divergences of these
specimens observed for the other four genes were similar to those observed in other
species. The high intraspecific divergences found for COI were consistent with the

findings of other studies of the gene in the Scolytinae (Cognato and Sun 2007: Menard

21

 

and Cognato 2007). For instance, COI sequence divergences observed for Ips DeGeer
species range from 0-10.0%, however, cryptic species were suspected in several cases
where COI sequence divergences were greater than 10%.

There was no difference observed for the 28S data between X. crassiusculus
specimens from Madagascar and Thailand. Similarly, specimens from introduced
populations in Maryland and North Carolina showed zero sequence divergence for all
five genes. The biogeographic history of X. crassiusculus is not entirely known and the
population genetics of the species is presently being investigated (Cognato et al.
unpublished data). However, these findings are consistent with the genetic bottleneck
that can occur with exotic species introductions, especially in the case of species with
haplodiploidy and extreme inbreeding, such as X ylosandrus.

High levels of intraspecific COI sequence divergence were observed in all other
species, except for X ylosandrus germanus. However, the findings for X germanus were
consistent with those for X. crassiusculus. The two specimens of X. germanus sampled
were from introduced populations in North America (Maryland and Michigan) and these
showed zero sequence divergence for all genes, expect for COI, which had a divergence
of 0.34%. The COI divergences for X. compactus and X. morigerus ranged from 7.35 —
9.74 %. Interestingly, the highest level of C01 sequence divergence was observed in the
two populations of Xyleborus rotundicollis sampled from Papua New Guinea (13.16%).
The possibility that this divergence is an artifact of the comparison of non-homologous
gene copies due to the presence of pseudogenes was tested and no evidence of
pseudogenes was found. These specimens were both collected from the highlands of

Papua New Guinea, where a continuous size gradient has been observed in X.

22

rotundicollis. The two specimens sequenced here represent the two extremes of the size
range. Further sequencing and comparisons of specimens across the entire size range
would be necessary to establish whether X. rotundicollis represents a single species or
two morphologically similar species with overlapping ranges.

As mentioned in the case of X. rotundicollis above, DNA sequences can be used
to help resolve species boundaries in cases where morphology provides little taxonomic
utility (Simon et al. 1994; Cognato and Sperling 2000; Scheffer and Lewis 2001; Pons et
al. 2006; Vogler and Monagham 2006). However, the findings of this study add to the
growing body of evidence suggesting that the implementation of DNA taxonomy must be
carefirlly tailored to suit the genetic characteristics of each taxonomic group (DeSalle et
al. 2005; Rubinoff 2006; Cognato 2007; Cognato and Sun 2007). Some authors have
proposed using a standard percentage sequence divergence as a means of differentiating
species (Hebert et al. 2003, 2004). However, inter- and intraspecific divergences can
vary dramatically, depending on the taxa. Furthermore, inter- and intraspecific distances
have been found to overlap in many groups of insects (Cognato 2006). Intraspecific
divergences in the Scolytinae have been observed to range between 0.04 — 13.16%
(Cognato 2006; Cognato and Sun 2007; Menard and Cognato 2007, this study). A
haplodiploidy system of inheritance and inbreeding, as is seen in the Xyleborina, could
have a dramatic effect on the divergence observed in these beetles. The effective
p0pu1ation size of haplodiploid organisms is at least twice that of diploid organisms
(Kirkendall 1993), thus new or rare haplotypes can accumulate faster. In addition, the
potential for introduced populations founded by a single female could lead to striking

patterns of divergence, as may be the case with species such as X. crassiusculus.

23

Therefore, any use of DNA taxonomy in xyleborines needs to take into consideration the

effects of haplodiploidy and inbreeding on genetic divergence and haplotype frequency.

Taxonomic Implications and Future Directions

The molecular characters utilized by this study have agreed with previously cited
morphological evidence that X ylosandrus is polyphyletic and that the current
classification of the genus contains species belonging to several genera (Dole and Beaver
in press). However, the taxon sample available for molecular work represents only a
fraction of the 54 species currently included in Xylosandrus. Due to the unavailability of
many species for molecular sequencing, morphological data will be the only way to
assess the proper generic placement of the majority of X ylosandrus species. For instance.
several species of X ylosandrus have morphologies which suggest that they should be
transferred to Amasa (see Chapter 2), but these species are so rarely collected that none
were available for DNA sequencing. Furthermore, the economic importance of
X ylosandrus requires the development of a morphology-based classification with clear
diagnostic characters and a key to the worldwide species. Such a revision is currently in
progress and will include the five gene data set used here in combination with
morphology to provide a more complete phylogenetic classification of X ylosandrus (see

Chapter 2).

24

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30

Gene Partion

 

 

 

EF-la COI Arglj CAD 285 Total
node
I 7.75 -0.50 9.25 13.25 9.25 39.00
2 5.75 -1.25 13.00 8.13 10.38 36.00
3 4.00 -3.00 0.67 -2.00 1.33 1.00
4 7.40 12.10 26.40 13.30 0.80 60.00
5 0.00 14.00 1.00 0.00 1.00 16.00
6 3.80 24.00 3.60 1.40 1.20 34.00
7 14.00 7.67 10.67 12.00 18.67 63.00
8 0.00 3.67 0.67 0.00 0.67 5.00
9 -4.00 7.50 2.00 2.00 1.50 9.00
10 3.60 -2.00 3.40 -4.40 2.40 3.00
11 2.00 -0.50 0.17 0.00 0.33 2.00
12 2.00 -0.50 3 .50 1.00 8.00 14.00
13 3.00 0.00 6.33 5.33 13.33 28.00
14 2.00 -1.57 0.43 -0.57 2.71 3.00
15 3.25 -2.25 7.75 7.25 16.00 32.00
16 3.60 -2.00 3.40 -4.40 2.40 3.00
17 7.00 -10.33 0.00 -1.00 25.33 21.00
18 -1.00 -1.00 -1.00 -1.00 5.00 1.00
19 2.33 2.67 3.67 -1.67 18.00 25.00
20 2.82 -3.27 1.55 -1.73 3.64 3.00
21 3.80 7.40 6.80 -1.00 7.00 24.00
22 1.71 3.79 -0.07 1.00 1.57 8.00
23 -0.50 17.00 5.50 -0.50 5.50 27.00
24 6.77 33.46 3.92 3.85 8.00 56.00
25 1.67 -1.33 2.33 2.00 4.33 9.00
26 -0.50 -3.00 1.50 3.50 2.50 4.00
27 6.00 -3.00 3.00 1 1.00 3.00 20.00
28 3.09 -3.73 1.45 -1.45 3.64 3.00
29 9.05 -O.20 1.25 -0.30 -5.80 4.00
30 1.20 3.80 -0.60 -1.20 ~0.20 3.00
31 0.00 1.00 -3.00 -1.00 5.00 2.00
32 0.00 -3.00 0.00 -1.00 1 1.00 7.00
33 0.00 6.00 0.00 0.00 9.00 15.00
34 2.00 0.00 0.00 -1.00 0.00 1.00
35 0.00 -7.00 7.00 0.00 2.00 2.00
36 0.00 -l .00 19.00 0.00 6.00 24.00
37 0.00 7.00 3.00 0.00 -1.00 9.00
38 2.00 -4.00 4.00 6.00 1.00 9.00
39 0.00 3.00 0.00 0.00 0.00 3.00
40 3.00 4.00 0.00 6.00 0.00 13.00
Total 108.59 103.61 151.53 72.78 204.48 641.00

 

Table 1.5: Partition branch support based on tree found by phylogenetic analysis 1 (see
Materials and Methods). Nodes refer to Figure 1.1.

31

Gene Partition

 

 

 

EF—la COI ArgK CAD 285 Total
node
I 1.33 0.33 6.33 16.33 8.67 35.00
2 2.33 2.33 14.33 10.33 8.67 38.00
3 1.33 -1.67 0.33 -0.67 0.67 0.00
4 11.33 11.33 30.83 13.83 0.67 68.00
5 -2.67 17.33 0.33 0.33 -0.33 15.00
6 -2.67 28.33 1.83 4.83 -0.33 32.00
7 6.33 13.33 11.33 16.33 28.67 76.00
8 0.33 5.33 4.83 7.83 1.67 20.00
9 1.33 1.33 0.33 10.33 25.67 39.00
10 -0.79 0.46 -1.04 -3.67 14.04 9.00
I] -1.27 -0.47 -1.67 -5.27 17.67 9.00
12 -0.67 -0.33 2.67 4.00 11.33 17.00
13 1.33 -0.67 6.33 3.33 36.67 47.00
14 -2.00 2.00 1.00 1.00 0.00 2.00
15 -0.67 -5.67 0.33 1.33 30.67 26.00
16 0.33 2.00 2.67 5.33 9.67 20.00
17 14.33 9.83 0.83 1.33 32.67 59.00
18 1.33 -1.67 2.33 1.33 3.67 7.00
19 4.33 4.33 9.33 -0.67 6.67 24.00
20 1.33 6.33 -0.67 -0.67 0.67 7.00
21 1.33 11.33 8.33 3.33 2.67 27.00
22 6.33 33.33 4.33 4.33 8.67 57.00
23 1.53 2.13 2.53 1.73 1.07 9.00
24 0.00 -1.00 2.00 4.00 -1.00 4.00
25 9.33 -3.33 4.50 8.17 5.33 24.00
26 2.00 -4.50 1.50 1.00 3.00 3.00
27 0.00 3.00 3.50 0.50 -2.00 5.00
28 0.33 -2. 17 3.83 -0.67 8.67 10.00
29 -0.67 —0. 17 0.83 -0.67 1.67 1.00
30 4.33 4.33 7.33 1.33 0.67 18.00
31 4.33 4.33 7.33 1.33 0.67 18.00
32 —0.67 -0.17 3.83 0.33 14.67 18.00
33 1.33 4.33 0.33 -0.67 29.67 35.00
34 1.33 -1.67 0.33 -0.67 0.67 0.00
35 1.33 -8.67 7.33 -0.67 2.67 2.00
36 1.33 -3.67 19.33 -0.67 6.67 23.00
37 -0.67 7.83 2.83 0.33 -1.33 9.00
38 2.33 -4.67 4.33 5.33 1.67 9.00
39 -0.67 3.83 -0. 17 0.33 -0.33 3.00
40 2.33 4.83 -O.17 6.33 -0.33 13.00
Total 73.47 145.46 176.66 121.30 321.11 838.00

 

Table 1.6: Partition branch support based on tree found by phylogenetic analysis 2 (see
Materials and Methods). Nodes refer to Figure 1.2.

32

Gene Partition

 

 

 

EF—la COI ArgK CAD 285 Total
node
I 3.00 6.00 1 1.00 12.00 1.00 33.00
2 6.00 7.00 13.00 12.00 3.00 41.00
3 5.33 -2.67 -1.67 -O.67 2.67 3.00
4 6.00 8.00 31.00 18.00 -4.00 59.00
5 2.00 4.50 -2.00 -0.50 0.00 4.00
6 4.00 3.00 -2.00 0.00 5.00 10.00
7 2.75 6.75 4.75 0.50 8.25 23.00
8 4.00 1.00 -5.00 -3.00 4.00 1.00
9 0.00 4.00 6.00 3.00 9.00 22.00
10 4.00 2.00 -5.50 -2.50 3.00 1.00
11 -1.00 14.00 9.00 -3.00 9.00 28.00
12 4.00 2.00 -5.50 -2.50 3.00 1.00
13 4.00 2.00 -5.50 -2.50 3.00 1.00
14 -0.67 0.33 5.00 3.33 -6.00 2.00
15 3.00 7.50 9.00 0.00 4.50 24.00
16 0.00 14.00 1.00 -1.00 -5.00 9.00
17 0.00 21.00 7.33 -0.67 -O.67 27.00
18 5.00 39.00 6.00 4.00 2.00 56.00
19 2.60 6.20 1.40 -0.60 -0.60 9.00
20 0.00 3.00 4.00 3.00 -4.00 6.00
21 7.00 2.00 2.00 10.00 0.00 21.00
22 1.33 13.67 5.67 1.67 -7.33 15.00
23 0.00 25.00 6.00 9.00 -12.00 28.00
24 11.80 11.10 13.70 15.90 12.50 65.00
25 -6.00 1.00 13 .00 0.00 -2.00 6.00
26 -O.67 0.33 5.00 3.33 -6.00 2.00
27 2.00 3.50 -1.75 -1.75 -1.00 1.00
28 -0.67 0.33 5.00 3.33 -6.00 2.00
29 -0.50 -1.00 6.00 2.50 -5.00 2.00
30 0.00 10.00 -0.67 -3.33 -2.00 4.00
31 0.00 3.33 2.33 -0.67 -4.00 1.00
32 -1.00 -5.00 1 1.00 1.00 -2.00 4.00
33 -0.50 10.00 4.50 -1.50 2.50 15.00
34 0.00 5.00 2.00 -1.00 -5.00 1.00
35 0.00 -6.00 6.00 0.00 1.00 1.00
36 1.40 2.80 18.00 -1.60 5.40 26.00
37 0.00 7.00 3.00 0.00 0.00 10.00
38 3.40 -1.80 2.60 4.60 -0.80 8.00
39 0.00 3.00 0.00 0.00 0.00 3.00
40 3.00 4.00 0.00 6.00 0.00 13 .00
Total 74.60 236.87 184.69 86.37 5.42 588.00

 

Table 1.7: Partition branch support based on tree found by phylogenetic analysis 3 (see
Materials and Methods). Nodes refer to Figure 1.3.

33

Gene Partition

 

 

 

EF-la COI ArgK CAD 288 Total
node
I -3.00 0.00 10.00 1 1.00 6.00 24.00
2 6.00 0.00 14.00 10.00 8.00 38.00
3 5.00 -5.00 1.00 -l .00 3.00 3.00
4 6.00 9.00 26.00 13.00 -1.00 53.00
5 -4.00 5.50 2.00 0.00 0.50 4.00
6 -4.00 5.50 2.00 0.00 0.50 4.00
7 -4.00 8.00 4.00 -4.00 10.00 14.00
8 -2. 17 20.50 3.83 4.67 4.17 31.00
9 -1.00 25.00 14.00 22.00 -1.00 59.00
10 -3.00 -11.50 3.00 0.00 12.50 1.00
11 14.00 17.00 0.00 5.00 4.00 40.00
12 -2.00 -14.00 -1.00 3.00 21.00 7.00
13 -0.50 -5.00 -0.50 2.50 6.50 3.00
14 -1.50 -2.25 1.50 1.00 9.25 8.00
15 0.00 0.00 3.00 4.00 4.00 1 1.00
16 -3.00 -8.50 3.00 12.00 30.50 34.00
17 0.00 -2.00 1.00 0.00 8.00 7.00
18 -3.00 1.00 7.00 0.00 48.00 53.00
19 -3.00 -1 1.50 3.00 0.00 12.50 1.00
20 -3.00 -1 1.50 3.00 0.00 12.50 1.00
21 3.00 8.00 7.00 0.00 6.00 24.00
22 0.00 1.00 1.00 4.00 0.00 6.00
23 0.00 15.00 6.00 0.00 4.00 25.00
24 6.33 28.33 6.00 9.00 12.33 62.00
25 1.00 -4.00 3.00 5.00 1.00 6.00
26 0.00 -5.00 2.00 4.00 1.00 2.00
27 4.00 -7.50 5.00 8.00 12.50 22.00
28 -3.00 -1 1.50 3.00 0.00 12.50 1.00
29 -3.00 ~11.50 3.00 0.00 12.50 1.00
30 -1.00 8.00 -1.00 0.00 0.00 6.00
31 -2.00 -5.25 1.25 -O.25 7.25 1.00
32 -3.00 -17.00 6.00 0.00 17.00 3.00
33 0.00 5.00 0.00 0.00 18.00 23.00
34 0.00 1.00 0.00 0.00 -1.00 0.00
35 0.00 -8.00 7.00 0.00 4.00 3.00
36 0.00 -2.00 19.00 0.00 9.00 26.00
37 0.00 7.00 3.00 0.00 3.00 13.00
38 2.00 -6.00 4.00 6.00 1.00 7.00
39 0.00 2.00 0.00 0.00 1.00 3.00
40 3.00 3.00 0.00 6.00 1.00 13.00
Total 1.17 20.83 175.08 124.92 321.00 643.00

 

Table 1.8: Partition branch support based on tree found by phylogenetic analysis 4 (see
Materials and Methods). Nodes refer to Figure 1.4.

34

 

 

 

 

 

 

 

 

 

 

 

EF-la COI ArgK CAD 288
X glosamlrus congpactus
Brazil vs. Ghana 0.0018 0.0769 0.0064 N/A 0.0061
Ave. Dist. Xyleborus pelliculosus 0.0961 0.1461 0.1033 0.063 0.0355
X ylosandrus crassiusculus
Madagascar vs. Thailand 0.0092 0.0444 0.0026 0.0014 0
Madagascar vs. Maryland 0.0031 0.1094 0.0051 0.0084 0.0013
Madagascar vs. N. Carolina 0.0031 0.1094 0.0051 0.0084 0.0014
Maryland vs. N. Carolina 0 0 0 0 0
Maryland vs. Thailand 0.0121 0.1009 0.0098 0.007 0.0013
N. Carolina vs. Thailand 0.01 18 O. 1009 0.0097 0.007 0.0013
Ave Dist. Xyleborus pelliculosus 0.0706 0.1726 0.1 104 0.0676 0.0665
X ylosandrus germanus
Maryland vs. Michigan 0 0.0034 0 0 0
Ave. Dist. Xyleborus pelliculosus 0.0976 0.1729 0.0918 0.0728 0.0339
Xylosandrus morigflus
Papua New Guinea vs.Ecuador 0.0018 0.0735 0.0042 0.007 0.004
Singapore vs. Papua New Guinea N/A 0.0855 0.0106 N/A 0.0016
Singapore vs. Ecuador N/A 0.0974 0.0106 N/A 0.0031
Ave. Dist. Xyleborus pelliculosus 0.0932 0.1573 0.0969 0.0742 0.0351
X Lleborus rotundicollis
Papua New Guinea 1 vs. Papua
New Guinea 2 0.0234 0.1316 N/A N/A 0.0069
Ave. Distance X yleborus
pelliculosus 0.1006 0.1658 0.1 148 0.0714 0.0528

 

Table 1.9: COI intra- and interspecific distances expressed as the proportion of sites

differing between sequences, with interspecific distances given as the average observed

between species.

35

38 —AAmasa n. sp. 301

96 40 39 masa resectus
36 WAmasa striatotruncatus
‘ 71 Amasa versicolor

.35 100' 37CAmasa bicostatus
34 79 98 Amasa schlichi

 

 

 

 

 

 

 

 

3, Amasa n. sp. 349
" Amasa n. sp. 1
86 33 Xyleborus califomicus Amasa
100 Xyleborus pelliculosus
Cnestus bimaculatus
4 '—_|_0:Cnestus pseudosuturalis

 

 

 

2 Xylosandrus ater
8 100 3 100 Xylosandrus mutilatus
__1 56 Xylosandrus improcerus

5 leosandrus crassiusculus/ Thailand “C ,, l
10 ‘__+3—§Xylosandrus crassiusculus/ Madagascar HCStUS

 

 

 

 

 

 

 

 

 

 

 

 

     
 

 

 

 

 

 

 

 

 

 

Xylosandrus crassiusculus/ USA. MD
100 Xylosandrus crassiusculus/ USA: NC
3_1 Xylosandrus mancus
16 '—::———X Xylosandrus discolor
'— '2 Anisandrus dispar
14% An1sandrus obesus
18 _ 61 Amsandrus say1
_ 15 13 Xylosandrus ursa
100 flylosandms ursinus ., ' ‘
20 Anisandrus hirtus “ , ‘ ”
1—1 Xylosandrus monteithi Anlsandl'us
—I::Xylosandrus n. sp. / Papua New Guinea
Xyleborus rotundicollis/ Papua New Guinea
—f(l)_0':Xy1eborus rotundicollis/ Papua New Guinea
‘ 29 21 Xylosandrus compactus/ Brazil
_‘ ”EECXylosandrus compactus/ Ghana
27 7‘) Xylosandrus morigerus/ Papua New Guinea
T36” 23 ., Xylosandrus morigerus/ Papua New Guinea
28 100 “Xylosandrus morigerus/ Singapore
x—_ 25 24|X losandrus ermanus/ USA: MD
86 100 Xillosandrus gemanus/ USA: M1
92 Xylosandrus n. s ./Bomeo
Xylosandrus queenslani
30r—Xyleborus affinjs Xylosandrus sensu stricto
5 fl—— Coccotrypes longior

 

F—— C occotrypes dactyliperda
-— 50 changes

Figure 1.1: Most parsimonious tree found by parsimony analysis of data aligned using
28S secondary structure (J ordal et a1. 2008), with gaps treated as missing data. Clade
numbers are given above nodes. Numbers below nodes are bootstrap support values.

36.

 

Amasa n. sp. 301
Amasa resectus

Amasa striatotruncatus
Amasa versmolor

37l— Amasa bicostatus
98 L— Amasa schlichi
Amasa n. sp. 349

Amasa n. sp. 1
33 r—' Xyleborus califomicus
1001— Xyleborus pelliculosus
Xyleborus afiinis
Xylosandrus mancus
Xylosandrus discolor
1 C nestus bimaculatus
4 '00 Cnestus pseudosuturalis

2 Xylosandrus ater
@ybsandms mutilatus
3 . Xylosandrus improcerus

98 5 Xylosandrus crassiusculus/ Thailand
7 I Xylosandrus crassiusculus/ Madagascar

I00 6 Xylosandrus crassiusculus/ USA: MD

100 Xylosandrus crassiusculus/ USA: NC
Anisandrus dispar
Anisandrus obesus

Xylosandrus ursa
Xylosandrus ursinus

   
  
   
 
 
 
     

 

 

 

30

 

 

 

 

 

 

133
I:

97

  
 
  

16
98

 

 

Anisandrus say1
13 Anisandrus hittus
70 15 Xylosandrus monteithi .
100' Xylosandrus n. sp./ Papua New Gumea
17 Xyleborus rotundicollis/ Papua New Guinea
W: Xyleborus rotundicollis/ Papua New Guinea
27 Xylosandrus compactus/ Brazil
65 TégCXylosandrus compactus/ Ghana
Xylosandrus morigerus/ Papua New Guinea
Xylosandrus morigerus/ Papua New Guinea
3 Xylosandrus morigerus/ Singapore
2 Xylosandrus gennanus/ USA: MD
100 Xylosandrus germanus/ USA: M1
Xylosandrus n. sp./ Borneo
Xylosandrus queenslandi
Coccotrypes dactilyperda

Coccotrypes longior
50 changes

   

 

 

 

 

 

 
  
   

._E
100

 

2—6 74
57

 

 

 

 

Figure 1.2: One of three most parsimonious trees found by parsimony analysis of data
aligned using 288 secondary structure (Jordal et al. 2008), with gaps treated as a 5th
character state. Clade numbers are given above nodes. Numbers below nodes are
bootstrap support values.

37

   
 
 
 
   
 

Amasa striatotruncatus
Amasa versicolor
Amasa bicostatus
Amasa schlichi
Amasa n. sp. 349
Amasa n. sp. 1
Xyleborus califomicus
100 Xyleborus pelliculosus
' Cnestus bimaculatus
Cnestus pseudosuturalis
Xylosandrus ater
ylosandrus mutilalus
63 Xylosandrus improcerus
6 Anisandrus dispar
Anisandrus obesus
69 Anisandrus sayi
7 l——Xylosandrus ursa
100 l—-—Xylosandrus ursinus
Anisandrus hirtus
Xyleborus rotundicollis/ Papua New Guinea
Xyleborus rotundicollis/ Papua New Guinea
Xylosandrus monteithi
Xylosandrus n. sp./ Papua New Guinea
15 Xylosandrus compactus/ Brazil
Xylosandrus compactus/ Ghana
16 Xylosandrus morigerus/ Papua New Guinea
JECXylosandrus morigerus/ Papua New Guinea
100 Xylosandrus morigerus/ Singapore
18 Xylosandrus germanus/ USA: MD
' _i19 100 iXylosandrus germanus/ USA: M1
92 Xylosandrus n. sp./ Borneo
22 Xylosandrus crassiusculus/ Thailand
24 99 Xylosandrus crassiusculus/ Madagascar
100 23 Xylosandrus crassiusculus/ USA: MD
100 Xylosandrus crassiusculus/ USA: NC
25 Xylosandrus mancus
—65|:Xylosandrus discolor
Xylosandrus queenslandi
_30C::Xyleborus afiinis
61 Coccotrypes longior
Coccotrypes dactyliperda
50 changes

    
   
   
 
 
   
    
  

 
  

 

 

 

 

 

 

 

 

 

 

Figure 1.3: Single most parsimonious tree found by parsimony analysis of data aligned
in MUSCLE, with gaps treated as missing data. Clade numbers are given above nodes.
Numbers below nodes are bootstrap support values.

38

 

38 Amasa n. sp. 301
Amasa resectus
Amasa striatotruncatus
Amasa versicolor
IOOI—Amasa bicostatus

L—Amasa schlichi
Amasa n. sp. 349
Amasa n. sp. 1
33 .—-—Xyleborus califomicus
looL—Xyleborus pelliculosus

010 Cnestus bimaculatus
|——_[—-—Cnestus pseudosuturalis

Xylosandrus ater
mrT-L—{OO—I—TL—Xylosandrus mutilatus
‘ Xylosandrus improcerus
Xylosandrus mancus

Xylosandrus discolor
31 Xylosandrus crassiusculus/ Thailand
" 9 978 Xylosandrus crassiusculus/ Madagascar
100 3 Xylosandrus crassiusculus/ USA: Maryland
20 100 Xylosandrus crassiusculus/ USA: North Carolina
— ll l——Xyleborus rotundicollis/ Papua New Guinea
13 100 l—-—Xyleborus rotundicollis/ Papua New Guinea
12 Xylosandrus monteithi
60 Xylosandrus n. sp./ Papua New Guinea
Anisandrus dispar
Anisandrus obesus
Anisandrus sayi
__22 18 96 Xylosandrus ursa

100' 100L Xylosandrus ursinus
Anisandrus hirtus

_l_1CXylosandrus compactus/ Brazil
‘ Xylosandrus compactus/ Ghana

 
 
 
  
  
  

 

 

32
76

 

 

 

 

 

 

 

I;

 

 

 

 

 

 

 

.—
I.

   
 

 

 

 

 

 

       
   
  

27 Xylosandrus morigerus/ Papua New Guinea
100‘ 232 94 Xylosandrus morigerus/ Papua New Guinea

'00 Xylosandrus morigerus/ Singapore
24 Xylosandrus gemianus/ USA: MD
100 Xylosandrus germanus/ USA: Ml
Xylosandrus n. sp./ Borneo

Xylosandrus queenslandi
Xyleborus affinis

30
—_5?L—— Coccotrypes longior

Coccotrypes dactyliperda
50 changes

 

 

I83

 

 

 

 

 

 

Figure 1.4: One of two most parsimonious trees found by parsimony analysis of data
aligned 1n MUSCLE, with gaps treated as a 5th character state. Clade numbers are given
above nodes. Numbers below nodes are bootstrap support values.

39

 

Amasa n. sp. 301
Amasa resectus
Amasa striatotruncatus
Amasa versicolor
Amasa bicostatus
Amasa schlichi

 

 

l9

 

Amasa n. sp. 349
80r—Xyleborus califomicus

 

 

L——)(yleborus pelliculosus

 

Amasa n. sp. 1

53 Cnestus bimaculatus
ECnestus pseudosuturalis
130 Xylosandrus improcerus

 

 

__58_:Xylosandrus ater
Xylosandrus mutilatus

76 r—Xyleborus rotundicollis/ Papua New Guinea

 

 

 

I—Xyleborus rotundicollis/ Papua New Guinea
Anisandrus dispar

Anisandrus obesus

Xylosandrus ursa

Xylosandrus ursinus

Anisandrus sayi

Anisandrus hirtus

 

 

 

 

 

 

Xylosandrus n. sp./ Papua New Guinea

 

46

 

Xylosandrus monteithi
9 r——Xylosandrus mancus

 

 

1——-——Xylosandrus discolor

69 Xylosandrus crassiusculus/ Thailand
96 [—i:Xylosandrus crassiusculus/ Madagascar

 

 

57 Xylosandrus crassiusculus/ USA: M1)
Xylosandrus crassiusculus/ USA: NC

‘#Q:Xylosandms compactus/ Brazil

 

46

 

 

Xylosandrus compactus/ Ghana
46 Xylosandrus morigerus/ Papua New Guinea
43 Xylosandrus morigerus/ Papua New Guinea
33 Xylosandrus morigerus/ Singapore
17 87 Xylosandrus germanus/ USA: MD

Xylosandrus gennanus/ USA: MI
Xylosandrus n. sp./' Borneo

 

 

 

Xylosandrus queenslandi
6 r—Xyleborus affinis
Coccotrypes longior

 

Coccotrypes dactyl iperda

Figure 1.5: One of two post parsimonious trees found by dynamic alignment and
analysis using POY, with a gap cost = 2. Numbers above nodes are Bremer support

values.

40

 

lOO

 

 

100

 

it:

I‘—
L—

 

100

 

 

100 IOO

 

 

 

100

 

 

 

 

55 100 I'—

l—

A:

 

 

 

 

100

 

 

97

 

l00

 

54

 

 

100

 

 

 

99

 

100
—l:

Amasa n. sp. 301

Amasa resectus

Amasa striatotruncatus

Amasa versicolor

Amasa bicostatus

Amasa schlichi

Amasa n. sp. 349

Amasa n. sp. 1

Xyleborus califomicus

Xyleborus pelliculosus

Xyleborus affinis

Cnestus bimaculatus

C nestus pseudosuturalis

Xylosandrus ater

Xylosandrus mutilatus

Xylosandrus improcerus

Xylosandrus crassiusculus/ Thailand
Xylosandrus crassiusculus/ Madagascar
Xylosandrus crassiusculus/ USA: MD

 

lOO

 

 

Xylosandrus crassiusculus/ USA: NC

 

100

 

 

51

 

96

 

 

 

100

 

 

 

i1:

100 r—Xylosandrus ursa
l———Xylosandrus ursinus

Anisandrus dispar
Anisandrus obesus
Anisandrus sayi

Anisandrus hirtus

 

 

 

 

 

 

100

 

100 100

 

 

 

 

 

100

 

67

 

 

IOO

 

 

 

100 J—Xylosandrus n. sp./ Papua New Guinea
L—Xylosandrus monteithi

100 l——Xyleborus rotundicollis/ Papua New Guinea
l———Xyleborus rotundicollis/ Papua New Guinea
100 r—Xylosandrus compactus/ Brazil
L—Xylosandrus compactus/ Ghana

Xylosandrus morigerus/ Singapore

100 l——Xylosandrus morigerus/ Papua New Guinea
nglosandrus morigerus/ Papua New Guinea

100
—l::

Xylosandrus gennanus/ USA: MD
Xylosandrus gennanus/ USA: Ml
Xylosandrus n. sp./ Borneo
Xylosandrus queenslandi

 

 

 

100 I—Xylosandrus mancus
7—Xylosandrus discolor

 

 

Coccotrypes dactyliperda
Coccotrypes longior

 

Figure 1.6: Bayesian tree found by analysis of data aligned using 288 secondary
structure (Jordal et al. 2008). Numbers above nodes are Bayesian posterior probabilities.

41

 

Amasa n. sp. 301
LCAmasa resectus
Amasa striatotruncatus
Amasa versicolor
r-—-Amasa bicostatus
I——Amasa schlichi
Amasa n. sp. 349
Amasa n. sp. 1
1———Xyleborus califomicus
L—Xyleborus pelliculosus
Xyleborus afiinis
|00 Cnestus bimaculatus
100 100 Cnestus pseudosuturalis
X osandrus ater
99 {Xi/(losandrus mutilatus
i Xylosandrus improcerus
77 Xylosandrus crassiusculus/ Thailand
94 100 Xylosandrus crassiusculus/ Madagascar
100 —— 100 Xylosandrus crassiusculus/ USA: MD
—— Xylosandrus crassiusculus/ USA: NC
100 r—Xylosandrus mancus
I—Xylosandrus discolor
78 100 r—Xylosandrus compactus/ Brazil
l——Xylosandrus compactus/ Ghana
———Xylosandrus morigerus/ PNG
100 Xylosandrus morigerus/ PNG
97 100 {Xylosandrus morigerus/ Singapore
_1 100 Xylosandrus germanus/ USA: MD
'00 {Xylosandrus germanus/ USA: Ml

Xylosandrus n. sp./ Borneo
Xylosandrus queenslandi

100 Xyleborus rotundicollis/ PNG
93 {Xyleborus rotundicollis/ PNG
79 ——Xylosandrus n. sp./ PNG
Xylosandrus monteithi
89 100 .__Anisandrus dispar

flCAnisandrus obesus

Anisandrus sayi
Xylosandrus ursa

Xylosandrus ursinus
Anisandrus hirtus
Coccotrypes dactyliperda

C occotrypes longior

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

   
 

 

 

 

 

 

Figure 1.7: Bayesian tree found by analysis of data aligned in MUSCLE. Numbers
above nodes are Bayesian posterior probabilities.

42

(A)

40.00
35.00
30.00
25.00
20.00
15.00
10.00

5.00

-5.00
-10.00
-15.00

Partioned Bremer Support (PBS)

0.00 -

0.02

0 EF-la, r = 0.0158, P = 0.4398, ns.

I COI, r = 0.1170, P = 0.0307, sig.. (1 = 0.05

' A 285, r = 0.07l3. P = 0.0957. ns.
A
I A A A
A o
A 3 A . 3
0 -£ IA 0 A) o . U I o ‘3
I!) 29 A 0 § §
‘ Of . fi ' I A l a
. I I I o- . I
I A
I
0.04 0.06 0.08 0.1 0.12 0.l4
Nodal Distance

Figure 1.8 (Cont. on next page): Relationships between PBS values and nodal distance
(maximum likelihood branch length from a given node to the tip of the tree) for the gene
partitions: 288, C01, and EF-la (A), CAD and ArgK (B).

43

(B)
o ArgK, r = 0.0006, P = 0.8844. ns.

30.00 . CAD, r = 0.0054, P = 0.6508. ns.

0
25.00
20.00 0
15.00
0 O .0
‘8 10.00 0 ' °
9- . a
500 ’ '° ° 3 o
' o o a 00 o .0 o
o 0 o
0.00- 9g» or 1. ,8 0‘: 3 9 gt
-5.00 . .
40.00
0 0.02 0.04 0.06 0.08 0.1 0.12 0.l4

Nodal Distance

Figure 1.8 (Cont): Relationships between PBS values and nodal distance (maximum
likelihood branch length from a given node to the tip of the tree) for the gene partitions:
288, C01, and EF-la (A), CAD and ArgK (B).

44

CHAPTER 2

Phylogenetic revision of Xylosandrus Reitter (Coleoptera: Curculionidae:
Scolytinae: Xyleborina)

45

ABSTRACT

A phylogenetic revision of the Xyleborina genus Xylosandrus Reitter based on
morphological and molecular data sets is presented. The monophyly of the genus was
tested using a 43 character morphological data set analyzed separately and in
combination with a molecular data set comprised of five independent gene loci: 28S
rDNA; the mitochondrial gene cytochrome oxidase I (C01); and the nuclear protein-
coding genes arginine kinase (ArgK), CAD (rudimentary), and elongation factor 1-01 (EF-
la). Xylosandrus was recovered as polyphyletic with the present classification
containing species from four different genera: Amasa Lea, Anisandrus Ferrari, Cnestus
Sampson, and Xylosandrus. A taxonomic revision of Xylosandrus is presented based on
these results. The following new combinations are given: Amasa cylindrotomicus
(Schedl), A. omissus (Schedl), A. oralis (Schedl), Anisandrus butamali (Beeson), A. ursa
(Eggers), A. ursinus (Hagedom), A. ursulus (Eggers), Cnestus ater (Eggers), C. fijianus
(Schedl), C. gravidus (Blandford), C. improcerus (Sampson), C. laticeps (Wood), C.
mutilatus (Blandford), C. orbiculatus (Schedl), C. peruanus (Wood), C. retifer (Wood).
C. retusus (Eichhoff), C. testudo (Eggers), Cyclorhipidion squamulatus (Beaver and
Loyttyniemi), Xylosandrus amputatus (Blandford), X. mixtus (Schedl), and X.
rotundicollis (Browne). Two new species of Xylosandrus are described: X borneensis
and X. hulcri. An illustrated key to worldwide species of Xylosandrus is provided.

Biogeography, host plants, diagnosis, and images are presented for each species.

46

INTRODUCTION

The scolytine subtribe Xyleborina contains approximately 1,300 described species
and constitutes one of the largest radiations of ambrosia beetles (Wood and Bright 1992;
Jordal 2002). The xyleborine mating system, which includes haplodiploidy and extreme
inbreeding, is believed to be the cause of this dramatic radiation (Normark et al. 1999).
Xyleborina are absent from the Dominican amber fossil record, suggesting that their
radiation began in the Miocene (Bright and Poinar 1994; Jordal et al. 2000). Ambrosia
beetles bore into the xylem of host trees and feed on symbiotic fungus, which grows on
the walls of their galleries. This ambrosia] feeding habit has evolved multiple times
within the Scolytinae and also in the related weevil subfamily Platypodinae (Jordal et al.
2000; Farrell et al. 2001). Ambrosia beetles tend to be less host-specific than their bark
beetle counterparts because adaptation to host secondary chemistry has not likely
influenced their radiation. This lack of host specificity, along with their haplodiploidy
mating system and inbreeding, make Xyleborina beetles particularly suited for the
invasion of new habitats and establishment as introduced exotic species. In theory, a
single female beetle could establish an invasive population of beetles. Furthermore, low
host specificity means that invasive beetles do not need to locate a specific host and can
infest multiple species. The economic importance of invasive xyleborines, along with
their interesting biology and ecology, has prompted much research interest in the
subtribe. Recently, the classification of the Xyleborina has been studied within a
phylogenetic context to update their classification within an evolutionary framework

(Jordal et a1. 2000; Jordal 2002; Hulcr et al. 2007).

47

Xylosandrus Reitter (1913) is a large genus of xyleborine ambrosia beetles with a
widespread distribution primarily in tropical and also in temperate regions throughout the
world. In their worldwide catalog of the Scolytinae, Wood and Bright (1992) list 52
species of Xylosandrus. Subsequent descriptions, new synonymies, and new
combinations have brought the present number to 54 species (Bright and Skidmore 1997,
2002; Saha et al. 1992; Wood 2007; Dole and Beaver in press). Several Xylosandrus
species cause economic losses in nursery and agricultural settings in their native and
introduced ranges. In Brazil, X. c0mpactus(1€ichhoft) causes losses in several
economically important host species, including avocado, cacao, coffee, and mango
(Oliveira 2008). X ylosandrus crassiusculus may potentially impact native tropical fauna,
as the species is present as an invasive in Costa Rican primary forests (Kirkendall and
@degaard 2007). Five .\’_i'lus(uzu’rus species currently occur in North America and of
these. only one is native, X. curmlus (Eichhol‘t). The other four. X. compacms. X.
germanus (Blandford). X. crass'iusm1111s (l\«'lotschulsky). and X. mull/Yams (Blandlbrd)
have been introduced from the Old World tropics (Rabaglia 2002; Rabaglia et al. 2006).
In North America, three Xylosandrus species (X. compactus, X. crassiusculus, and X
germanus) have caused “considerable economic damages” since their introductions
(Oliver and Mannion 2001). It is difficult to get exact figures on the economic losses
attributed to Xylosandrus species, since these numbers are often grouped with damage
caused by other xyleborines. As an example, nursery managers in Maryland reported
individual loses of $3,650 - $8,400 in nursery stock to Xylosandrus species in the spring

of 2008 alone (R. J. Rabaglia per. com.).

48

The current classification of Xylosandrus contains species with highly variable
morphologies, several of which are similar to those of other xyleborine genera (Hulcr et
al. 2007; Dole and Beaver in press). Recent phylogenetic analyses of molecular data
have also suggested that Xylosandrus is polyphyletic (Jordal 2002; see Chapter 1).
Xylosandrus requires the development of a revised classification based on morphological
and molecular evidence given the apparent taxonomic ambiguities and the economic
importance of this group (Dole and Beaver in press; see Chapter 1). Furthermore, clear
diagnostic characters and a key to worldwide species are necessary for the classification
to be a predictive and useful tool in identification and control of invasive X ylosandrus
species.

In this study, we define 43 morphological characters in an effort to resolve
taxonomic ambiguities among Xylosandrus species and related genera. We conduct a
combined phylogenetic analysis of these taxa using these morphological characters and

molecular data in order to guide a revision of Xylosandrus.

Systematics
X ylosandrus was established as a monotypic genus by Reitter in 1913, with
Xylosandrus morigerus designated as the type species. The genus remained monotypic
for many years until Hoffmann (1941) transferred X yleborus germanus into X ylosandrus.
Following this, Nunberg (1959) transferred X erborus compactus into X ylosandrus and
scolytine taxonomists began to take notice of the genus. In 1962, Schedl disagreed with
the designation of the genus and listed Xylosandrus as a synonym of Xyleborus Eichhoff.

Browne (1963) noted character differences that he felt justified the genus, chiefly the

49

“broad, obtuse prosternal process separating the front coxae” and transferred 14 species
from X yleborus to X ylosandrus. Schedl (1964) agreed with Browne’s assessment and
transferred 8 more species from X yleborus to X ylosandrus. Later, Schedl (1971)
described the first species originally described as Xylosandrus, rather than being
transferred from Xyleborus: X. adherescens and X. assequens.

In 1982, Wood began transferring species into Xylosandrus, chiefly from
Xyleborus, and within a decade the genus grew to include 52 species (Wood 1982, 1984;
Wood and Bright 1992). At this point in its taxonomic history, the generic limits and
defining characters of Xylosandrus were blurred. Many species transferred to
X ylosandrus by Wood and Bright (1992) have subcontinguous or contiguous procoxae,
along with many other characters that set them apart from the species originally included
in the genus. Since Wood and Bright gave no discussion of characters supporting their
new combinations, it is difficult to ascertain their reasoning. Since 1992, additional
species have been described in or transferred to Xylosandrus, only some of which fit the
characters that originally defined the genus (Beaver 1998; Saha et al. 1992; Wood 2007;
Dole and Beaver in press).

The incorrect placement of many species within X ylosandrus has created some
taxonomic confusion between Xylosandrus and the genera Amasa Lea (1893) and
Cnestus Sampson (1911). In addition to this, the recent resurrection of the genus
Anisandrus Ferrari (1867) has highlighted similarities between it and several species of
Xylosandrus (Hulcr et al. 2007). In their review of the Australian species of X ylosandrus,
Dole and Beaver (in press) made tentative steps toward correcting the taxonomy of

X ylosandrus by defining characters that separate X ylosandrus sensu stricto from C nestus

50

and transferring two species, Cnestus pseudosolidus (Schedl) and C. solidus (Eichhoff)
from Xylosandrus to C nestus.

Xylosandrus sensu stricto can be distinguished from other xyleborine genera by
the following combination of characters: stout body, usually about twice as long as wide,
widely separated procoxae, flat scutellum that is flush with the surface of elytra, and
obliquely truncate antennal club with the first segment forming a circular costa and dense
pubescence on the oblique portion of the club (Reitter 1913; Browne 1963; Bright 1968;
Wood 1986; Hulcr et al. 2007; Dole and Beaver in press).

Amasa can best be distinguished from X ylosandrus by its antennal club, which is
oval, not truncate, with prominent 1St and 2nd segments, separated by sutures that are
visible on both the anterior and posterior face (Lea 1893; Wood 1986; Hulcr et al. 2007).
In all Xylosandrus, the antennal club is obliquely truncate, with the first segment forming
a circular costa, lacking sutures posteriorly and with segment one covering the entire
posterior face. Amasa also have contiguous procoxae (Lea 1893; Wood 1986; Hulcr et
al. 2007). In all Xylosandrus sensu stricto, the procoxae are separated by an intercoxal
piece that is at least half the width of the coxae.

Anisandrus can be distinguished from Xylosandrus by the contiguous procoxae
(Hulcr et al. 2007). The lateral margin of the protibae of Anisandrus are armed with 6 — 7
socketed teeth. In Xylosandrus sensu stricto there are always only 4 — 5 socketed teeth
on the lateral margin of the protibiae. Anisandrus species also have pronotal lateral
margins that are rounded, a character which separates them from C nestus, which has
carinate lateral margins (Hulcr et al. 2007). X ylosandrus sensu stricto contains species

with both rounded and carinate lateral margins of the pronotum.

51

Cnestus can be distinguished from Xylosandrus by the subcontiguous procoxae
(Sampson 1911; Hulcr et al. 2007). Cnestus also have antennae with four funicular
segments, whereas Xylosandrus species have five. Additionally, in C nestus the anterior
margin of the pronotum bears 4 or fewer asperities, with a pair of coarse asperities
medially, and the pronotum is often produced anteriorly. In Xylosandrus the anterior
margin of the pronotum bears 6 or more smaller asperities of approximately equal size
and is never produced anteriorly. Many species of Cnestus have elytra that are wider
than they are long, a character which is never observed in X ylosandrus sensu stricto.

Recent taxonomic work has examined the classification of xyleborine genera
within a phylogenetic context (Jordal et al. 2000; Jordal 2002; Hulcr et al. 2007; see
Chapter 1). A cladistic review of the generic taxonomic characters of Xyleborina
recovered a monophyletic Xylosandrus, but it is important to note that this study did not
include species with morphologies that deviate from the sensu stricto concept of the
genus (Hulcr et al. 2007). Dole and Beaver (in press) were the first to treat the
classification of problematic Xylosandrus species by defining characters that distinguish
Xylosandrus from Cnestus. Phylogenetic analyses of DNA sequence data have also
recovered a non-monophyletic X ylosandrus. Jordal (2002) used the nuclear gene
Elongation Factor -lu(EF-1 a) to test the monophyly of the Xyleborina and, while the
subtribe was recovered as monophyletic, Xylosandrus was found to be paraphyletic with
respect to Cnestus. Phylogenetic reconstruction of a subset of Xylosandrus taxa using
data from mitochondrial and nuclear genes recovered a polyphyletic X ylosandrus with
the placement of Anisandrus and Cnestus among Xylosandrus species (see Chapter 1). In

addition, several Xylosandrus species were placed within the “A nisandrus” and

52

“Cnestus” clades with strong support (e.g., 100% bootstrap support), further indicating

the need for a taxonomic revision of Xylosandrus.

Biogeography

Xylosandrus is widely distributed in primarily tropical, but also in temperate
regions worldwide. Whereas the inclusion of the problematic species discussed above
does not change the overall distribution in terms of biogeographical regions, the
biogeography discussed herein will pertain to Xylosandrus as it is defined in this
taxonomic revision (Table 2.1). Three species of Xylosandrus have almost
circumtropical distributions: X. compactus, X. crassiusculus, andX. morigerus. A fourth
species, Xylosandrus germanus is less widely distributed, occurring in the Nearctic
Region, Oceania, the Oriental Region, and the Palearctic Region. These four species
were all introduced as exotic invasive species into temperate regions of the world.
Xylosandrus has a high level of regional endemism, with 71% of species occurring in
only one biogeographical region. The Oriental Region is the most species rich, with 76%
of Xylosandrus species present and a high occurrence of endemic species. The next
richest is the Australian Region, with 29% of Xylosandrus species present. The
Afrotropical Region and Oceania each contain 13%, and the Palearctic Region contains
16% of Xylosandrus species. Contrary to its tendency towards remarkable levels of
species diversity, the Neotropical Region contains the same number of Xylosandrus
species as the Nearctic (where the vast majority of species are introduced). This is

congruent with the biogeography of the rest of Xyleborina, which tend to have their

53

highest levels of diversity in the Old World Tropics (Beaver 1979; Wood and Bright

1992).

Natural History

As is typical of ambrosial feeding scolytines, Xylosandrus beetles are usually not
host specific (Beaver 1979). Known host plants are listed for each species herein. All
xyleborine beetles are haplodiploid, inbreeding and sexually dimorphic (Figs. 2.33 and
2.34). The diploid males are dwarfed, flightless and bear little resemblance to their
female counterparts. All Xylosandrus species are xylomycetophagous. The female
carries ambrosial fungi within a mycangium between the pronotum and mesonotum.
Colonization of a host tree begins when a female beetle locates a suitable host and
initiates gallery construction. As she bores into the tree, the female inoculates the woody
tissue lining the gallery tunnels with a suite of microorganisms, including the ambrosial
fungi. Once the fungi have begun to grow along the walls of the gallery, the female
beetle lays eggs in small clusters at the end of the main tunnel. X ylosandrus larvae and
adults will feed exclusively on the ambrosial fungi. If the establishing female was
unmated, she will produce only male offspring. In this case, as soon as they have
pupated into adults, the flightless and dwarfed males (Fig. 2.34) will mate with their
mother to produce female offspring. If the establishing female had mated prior to gallery
construction, she will produce both male and female offspring, but with a highly skewed
sex ratio (Kirkendall 1993). In Xyleborina beetles, the female to male sex ratio has been

observed to be as high as 30:1 (Bright 1968; Krikendall 1993). When both male and

54

female offspring have been produced, the males will mate with their sisters to produce

the next generation of beetles (Kirkendall 1993).

MATERIALS AND METHODS

Approximately 1,300 specimens were studied from entomological collections.

The following acronyms are used for collections referenced in the text:

AMNH
BMNH
BPBM
CAS
CSCA
FICB
FMNH
FRCS
FRI
IRSNB
IZM
MNB
MRCB
MSUC
MTD

MZLU

American Museum of Natural History, New York, New York, USA.
The Natural History Museum, London, United Kingdom.

Bernice P. Bishop Museum, Honolulu, Hawaii, USA.

California Academy of Sciences, San Francisco, California, USA.
California State Collection of Arthropods, Sacramento, California, USA.
Forest Research Centre, Lae, Papua New Guinea.

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

Forest Research Centre, Sabah, Sandakan, Malaysia.

Indian Forest Research Institute, Dehra Dun, Uttar Pradesh, India.
Institut Royal des Sciences Naturelles de Belgique, Belgium, Brussels.
Institute of Zoology, Moscow, Russia.

Museum fur Naturkunde der Humbolt University, Berlin, Germany.
Musee Royal du Congo Belgiquem, Tervuren, Belgium.

Michigan State University Collection, East Lansing. Michigan, USA.
Musetun fur Tierkunde, Dresden, Germany.

Lund University, Lund, Sweeden.

55

MZUSP

NHMW

NHR

NZSI

UCDC

USNM

ZFMK

Museum de Zoologia, Universidade de Sao Paulo, Sao Paulo, Brazil.
Naturhistorisches Museum Wien, Wien, Austria.

Naturhistoriska riksmuseet, Stockholm, Sweden.

Zoological Survey of India, National Zoological Collection, Calcutta.
India.

Roger A. Beaver, personal collection, Chiang Mai, Thailand.

R. M. Bohart Museum of Entomology, University of California. Davis.
California, USA.

National Museum of Natural History, Washington, DC, USA (Including
Stephen L. Wood Collection).

Zoologische Forschungsinstitut und Museum Alexander Koenig, Bonn.

Germany.

Plant host species were compiled from the following publications: Bright and

Skidmore (1997, 2002), Cibrian et al. (1995), Ohno (1990), Schedl (1962), Wood and

Bright (1992). Hosts recorded since the publication of Bright and Skidmore (2002) were

collected from the literature by the authors and provided by Dr. Don Bright (per. com.).

Plant author names, when missing, were added using the International Plant Names Index

(www.ipni.org).

Morphological Characters

A total of 43 external morphological characters (24 binary, 19 multistate) were

coded and used in the phylogenetic analysis. Morphological characters were scored for

56

females only, since the morphological classification of Xyleborina beetles is based
entirely on females and males are rare or unknown for many species. Characters were
scored for 52 species of Xylosandrus (Table 2.2). In order to test the generic limits of
Xylosandrus, a diversity of taxa were included in the morphological analysis: three
Anisandrus, nine A masa, three Cnestus, one C occotrypes and three X yleborus species
(Table 2.2). Characters coded were distributed as follows: two habitus characters (five
states), five head characters (14 states), 14 pronotal characters (35 states), two elytral
characters (five states), and 20 elytral declivity characters (51 states). Characters that
were ambiguous or difficult to see on the specimen(s) examined were coded as “?” and
treated as missing data. Inapplicable characters were coded as “7” and treated as missing
data. All characters were treated as non-additive and unweighted in the phylogenetic
analysis.

The following morphological characters and character states were used in the
phylogenetic analysis. The character data matrix (Table 2.2) was prepared and coded

using the program MX (Yoder et al. 2006-Present).

Habitus
1. Body ratio

0 = less than 2.0 times longer than wide

1 = more than 2.0 times longer than wide
2. Pronotal to elytra color

0 = roughly the same color

1 = elytra distinctly darker than pronotum

57

2 = overall, same color, but elytra with testaceous patches
Head
3. F rons sculpture

0 = punctuate

1 = finely granulate

2 = reticulate or rugose

? = ambiguous or difficult to see
4. F rons median keel

0 = absent

1 = present

? = ambiguous or difficult to see
5. Antenna] funicular segment count

0 = 4-segmented

1 = 5-segmented

2 = 6-segmented

? = ambiguous or difficult to see
6. Antennal club type

0 = circle absent

1 = circle closed anteriad

2 = circle closed posteriad

? = ambiguous or difficult to see
7. Segments visible on posterior face of antennal club

0 = segments 2 and 3 not visible. segment 1 covering whole face

58

l = segment 2 visible, segment 1 covering most of face
2 = segments 1, 2, and 3 visible on face
? = ambiguous or difficult to see
Pronotum
8. Pronotum ratio
0 = Pronotum wider than long
1 = pronotum of equal length and width
2 = pronotum longer than wide
9. Pronotal type dorsal
0 = rounded (type 1, Hulcr et al. 2007)
1 = basic (type 2, Hulcr et al. 2007)
10. Pronotal type laterally
0 = basic
1 = rounded
2 = prolonged anteriorly
3 = prolonged posteriorly
11. Pronotal vestiture
0 = erect, hair-like setae
1 = semi-appressed, hair-like setae
12. Pronotal basal setae
0 = glabrous, without setae (except for mycangium, when present)
1 = moderately setose, with basal setae being less dense than anterior

2 = densely setose, with basal setae being at least as dense as anterior

59

13. Pronotal mycangial setae
0 = absent
1 = present
? = ambiguous
14. Pronotal basal sculpture
0 = punctate
1 = asperate-granulate
15. Pronotal basal sculpture density
0 = moderate sculpture, punctures or granules separated by a distance greater than
their size
1 = dense sculpture, punctures or granules separted by a distance equal to or less
than their size
16. Pronotal anterior serrations count
0 = 4 or fewer serrations
1 = 6 or more serrations
17. Pronotal anterior serrations course median pair
0 = absent
1 = present
? = only two anterior serrations present
18. Pronotal anterior margin produced
0 = not produced
1 = produced

19. Pronotal lateral carina

60

0 = absent
1 = present
20. Procoxal separation
0 = widely separated, intercoxal piece at least 1/3 the width of coxae
1 = narrowly separated to subcontiguous
2 = contiguous
? = procoxae not visible
21. Protibial teeth
0 = 4 or 5 socketed teeth
1 = 6 socketed teeth
2 = 7 socketed teeth
3 = 8 socketed teeth
? = ambiguous or difficult to see
Elytra
22. Elytral ratio
0 = elytra longer than wide
1 = elytra of equal length and width
2 = elytra wider than long
23. Discal interstrial sculpture distribution
0 = uniseriate
l = multiseriate, including biseriate
Elytral Declivity

24. Declivital origin

61

0 = declivity originating less than 1/3 the length of elytra from base
1 = declivity originating 2 1/2 the length of elytra from base
25. Decilivital slope
0 = elytral disc gradually curving into declivity
1 = declivital face abrupt and steeply separated from disc
26. Declivital shape
0 = not circular
1 = circular
27. Declivital surface
0 = convex
1= flattened
2 = concave, at least in part
28. Declivital carina
0 = absent, rounded
1 = granulate
2 = carinate
3 = serriate
29. Declivital carina length
0 = carina not extending beyond 71h interstriae
1 = carina extending beyond 7‘h interstriae, forming a circumdeclivital ring
? = absent
30. Declivital striae count

0 = 3 striae visible on declivity

62

1 = 3 striae visible on declivity
2 = 4 striae visible on declivity
3 = 5 striae visible on declivity
4 = 6 striae visible on declivity
? = ambiguous
31. Declivital strial impression
O = not impressed
= impressed
32. Declivital strial sculpture
0 = punctuate
1 = granulate
33. Declivital strial sculpture distribution
0 = straight
I = confused
34. Declivital strial setae type
0 = absent
1 = hair-like
2 = scale-like
? = ambiguous
35. Declivital strial setae length
0 = length less than or equal to the width of second declivital interstriae
1 = length greater than the width of second declivital interstriae

2 = length at least 2 times the width of second declivital interstriae

63

? = absent or ambiguous
36. Declivital strial setae profile
0 = appressed or semi-appressed
= erect
? = absent or ambiguous
37. Declivital interstrial sculpture
0 = punctuate
l = coarsely granulate
2 = finely granulate
38. Declivital interstrial sculpture distribution
0 = uniseriate
1 = multiseriate, including biseriate

39. Declivital interstrial setae type

0 = absent
1 = hair-like
2 = scale-like

? = ambiguous
40. Declivital interstrial setae length
0 = length less than or equal to the width of second declivital interstriae
1 = length greater than the width of second declivital interstriae
2 = Length at least twice the width of second declivital interstriae
? = absent or ambiguous

41. Declivital interstrial setae profile

64

0 = appressed or semi-appressed
1 = erect
? = absent, ambiguous
42. First interstriae elevated at apex ofelytra
0 = not elevated
1 = elevated
43. Granules or tubercles near apex of first interstriae
0 = absent

1 = present

Molecular Characters

The molecular data set used herein (see Chapter 1) is comprised of multiple gene
loci, chosen for their complementary phylogenetic signals at varying nodal depths: 28S
rDNA; the mitochondrial gene cytochrome oxidase I (C01); and the nuclear protein-
coding genes argenine kinase (ArgK), CAD (rudimentary), and Elongation Factor -101
(EF-la). For extraction, sequencing and alignment protocols see Chapter I. For our
combined analysis, we used the molecular data set aligned manually with reference to a
scolytine-specific secondary structure model (Jordal et al. 2008). The resulting combined
data set included only taxa for which both morphological and molecular data were
available (Table 2.3). This combined data set includes 27 taxa, representing 15
X ylosandrus species, 12 species belonging to the genera A masa. Anisandrus, C nestus,

Xyleborus, and the oustgroup genus Coccotrypes (Table 2.3).

65

Phylogenetic Analysis

Phylogenetic analysis of the morphological data set was conducted with the
software TNT (Goloboff et al. 2003). A new technology driven search was employed for
the parsimony analysis with all search modules employed: sectorial search (RSS and
C SS), ratchet, drift, and tree fiising. Default TNT setting were used, with the following
exceptions: tree fusion was conducted globally after every hit and the search was set to
end after the minimal tree length was found 10 times. Traditional (heuristic) search
methods were then used to conduct tree bisection and reconnection branch swapping on
the 43 most parsimonious trees found in the new technology search. Maximum trees
being held by TNT was set to 10,000. Bootstrap support values were calculated by
performing 1,000 pseudo-replicates with simple sequence addition in the program
Winclada (Nixon 1999).

Phylogenetic analysis of the combined data set was conducted with the software
PAUP* (Swofford 2002). A heuristic search was employed with 300 random stepwise
addition replicates using PAUP* default settings. Gaps were treated as missing data in
the analysis. Bootstrap values were calculated by performing 1,000 pseudo-replicates
with simple addition sequence in PAUP*. Bremer support values for each data partition
were calculated by constructing a constraint tree with the software TreeRot (Sorensen

1996) followed by subsequent analysis with PAUP*.

RESULTS

Xylosandrus was recovered as polyphyletic by analyses of both the morphological

and the combined data set. Phylogenetic analysis of the morphological data set produced

66

10,000+ equally parsimonious trees of 318 steps (max trees in TNT set to 10,000) (Fig
2.1). The strict consensus tree was mostly unresolved, but recovered several clades with
high support values. The monophyly of the clade containing Amasa, Anisandrus,

C nestus, and Xylosandrus with respect to the genus Xyleborus was recovered with 100%
bootstrap support (see discussion of X yleborus rotundicollis below). The unresolved
placement of the “Amasa”, Anisandrus, and “Cnestus” clades, as well as the placement of
several species of Xylosandrus within these clades, was responsible for rendering

X ylosandrus paraphyletic. While morphology did not resolve their phylogenetic
placement, the data did support the monophyly of the “Amasa” and “C nestus” clades and
the inclusion of several X ylosandrus species within them. These clades were both
recovered with 100% bootstrap support. Anisandrus was recovered as monophyletic with
100% bootstrap support. The Consistency Index (Cl) indicates that the occurrence of
homoplasy is high in the morphological data set (CI = 0.190). Furthermore, the
Retention Index (RI) indicates that character state changes are occurring predominantly
on the internal nodes (R1 = 0.671 ).

Phylogenetic analysis of the combined data set recovered a single most
parsimonious tree of 4524 steps (Fig. 2.2). This tree was well resolved, with high support
values (e. g. 2 90% boostrap support) toward the terminal nodes and poorer support (e. g.
S 74 % bootstrap support) for the deeper relationships among the clades. The placement
of the “Anisandrus”, and “C nestus” clades, the inclusion of several X ylosandrus species
within these clades, as well as the placement of X ylosandrus mancus and X. discolor in a
clade with Amasa and Xyleborus, rendered Xylosandrus polyphyletic. The low support

for deeper nodes made it impossible to determine the phylogenetic relationships among

67

Xylosandrus, Amasa, Anisandrus, Cnestus, and Xyleborus. However, the placement of
several species of Xylosandrus within the “Anisandrus” and “Cnestus” clades had very
strong support (e. g. 100% bootstrap support). The genus A masa and a X ylosandrus sensu
stricto clade (containing the type species X. morigerus) were also recovered with 100%
bootstrap support.

Homoplasy as measured by C1 and RI indicated that homoplasy was lowest for
the 28S data partition (Table 2.4). The nuclear protein-coding genes ArgK, CAD, and
EF-lu had similar CI’s (0.474 — 0.522) and RI’s (0.460 — 0.600). The mitochondrial
gene COI had the highest level of homoplasy (C I = 0.265, R1 = 0.246). The low RI value
for COI indicates that the character state changes for this gene largely occurred towards
the terminal nodes of the tree. The morphological data demonstrated lower levels of
homoplasy in the combined analysis (CI = 0.346) than it did when analyzed alone (CI =
0.200). However, the RI observed for the morphological data was lower in the combined
analysis (R1 = 0.492), indicating that the occurrence of character state changes shifted
more toward the terminal nodes when the data were combined with molecular data.
Overall, 288 gave the highest bremer support values, with CAD having the next highest
values (Table 2.5). Interestingly, the overall bremer supports for C01 and morphology

were similar.

DISCUSSION

All analyses conducted herein recovered a polyphyletic Xylosandrus, with the
present classification of the genus containing species from at least four different genera:

Xylosandrus, Amasa, Anisandrus, and Cnestus. These findings are consistent with those

68

of other studies of Xylosandrus (Jordal 2002; Dole and Beaver in press; see Chapter 1).
Separate and combined analyses of morphological and molecular data sets have
recovered the following clades with high support values: “A nisandrus”, “A masa“,
“Cnestus” and “X ylosandrus sensu sticto” (see Chapter 1). However, even a data set
combining morphology and five gene partitions was not sufficient to resolve the
relationships among these genera (Fig. 2.2). Despite this, these trees provide a valuable

framework with which to revise the present classification of X ylosandrus.

Amasa Clade

The “Amasa” clade has been consistently recovered by all parsimony analyses
(Bayesian and POY analyses of molecular data recovered an unresolved or polyphyletic
Amasa, with respect to Xyleborus) (see Chapter 1) (Figs. 2.1 and 2.2). The genus is
rendered monophyletic by the inclusion of three X ylosandrus species: X cylindrotomicus,
X. omissus, and X, oralis (Fig. 2.1). The placement of these species within Amasa is
supported by the morphological characters used to define Amasa and distinguish it from
other Xyleborina genera (Hulcr et al. 2007) (Fig. 2.3). Similarly, the species A. mixtus
and A. amputatus were included among X ylosandrus species, rather than Amasa. by the
analysis of the morphological data set. The transfer of these species into X ylosandrus is
consistent with the characters defining X ylosandrus sensu stricto. The combined
phylogenetic analyses have indicated that Amasa is more closely related to X yleborus
than it is to Anisandrus, C nestus, and Xylosandrus. Thus, it appears that confusion
between Xylosandrus and Amasa was simply the result of taxonomic error and not an

indication of close phylogenetic relationship.

69

 

Anisandrus Clade

The “Anisandrus” clade has been recovered by all analyses that included
molecular data (see Chapter 1) (Fig. 2.2). The only analysis that did not recover this
clade was that of morphological data alone (Fig. 2.1). However, the membership of
species included in this clade is consistent with characters that support the separation of
Anisandrus from other Xyleborina genera (Hulcr et al. 2007) (Fig. 2.4). All other
analyses recovered the “Anisandrus” clade with high support (100% boostrap support,
Bayesian posterior probability of 100, and Bremer supports ranging from 22-67) (see
Chapter 1) (Fig. 2.2; Table 2.5). The genus Anisandrus is rendered monophyletic by the
inclusion of three Xylosandrus species: X. ursa, X ursinus, X. ursulus. Transfer of these
species to Anisandrus is also supported by the morphological characters that distinguish
the genus (Hulcr et a1. 2007). Xylosandrus butamali is a fourth species with the
morphological characteristics of A nisandrus rather than those of X ylosandrus. This
species was not available for DNA sequencing and was only included in the
morphological analysis. This analysis placed X borealis in a larger clade with the
“A nisandrus” and “Cnestus” clades with 93% bootstrap support, but the relationships
within this clade were unresolved in the strict consensus tree. Based on this phylogenetic
evidence, in combination with the morphological characters that distinguish A nisandrus.

X. borealis is transferred to Anisandrus herein.

Cnestus Clade

70

 

The “Cnestus” clade has been recovered by all phylogenetic analyses (see Chapter
1) (Figs. 2.1 and 2.2). The genus is rendered monophyletic by the inclusion of 1 1
Xylosandrus species: X. ater,X. fijianus, X. gravidus, X. improcerus, X. laticeps. X.
mutilatus,X. 0rbiculatus,X. peruanus,.X. retifer,X. retusus, andX. testudo. These are in
addition to two species already transferred from Xylosandrus to C nestus by Dole and
Beaver (in press): X. pseudosolidus and X. solidus. The inclusion of these species in
C nestus is also supported by morphological characters used to distinguish the genus
(Hulcr et al. 2007; Dole and Beaver in press) (Fig. 2.5). Support values for the “C nestus”
clade were very high (100% bootstrap support, Bayesian posterior probability of 100, and

Bremer supports ranging from 53-120).

The transfer of the above species from X ylosandrus to Cnestus is of some
importance to scolytine control, considering the establishment of X. mutilatus as an
invasive species in North America. The inclusion of X. mutilatus in C nestus constitutes a
new generic record for North America (Rabaglia et al. 2006). Likewise, the transfer of
several South American species (X. laticeps, X. peruanus, X. retifer, X. retusus) to
Cnestus establishes, for the first time, the presence of the genus in the Neotropics (Wood

2007).

X ylosandrus sensu stricto C lade
The “Xylosandrus sensu stricto” clade was consistently recovered by all analyses
that included molecular data (see Chapter 1) (Fig. 2.2). The analysis of morphological
data did not recover this clade, but the resolution for all Xylosandrus species not placed in

clades with other genera was poor (Fig. 2.1). The “X ylosandrus sensu stricto” clade is

71

comprised of (((X germanus + X. borneensis n. sp.) X. morigerus) X. compactus). Given
that it contains X. morigerus, the type species of X ylosandrus, this clade is the highest
supported grouping of species belonging to Xylosandrus sensu stricto found by
phylogenetic analyses of the genus. Support values for this clade were very high (100%
bootstrap support, Bayesian posterior probabilities of 100, Bremer supports ranging from
21-28). The species included in this clade are morphologically consistent with the strict
definition of the genus. This clade also contains three economically important species of

Xylosandrus: X. compactus, X germanus, and X morigerus.

Xylosandrus sensu lato

The phylogenetic placement of the remaining Xylosandrus species was largely
unresolved by these analyses. However, a couple of clades were recovered by the
combined data analysis with high support (2 92 % bootstrap support): X hulcri n. sp. +
X. moteithi and X discolor + X mancus (Fig. 2.2). The placement of a few species calls
into question the monophyly of X ylosandrus, even after this revision. The placement of
X crassiusculus as sister to Cnestus, which has been recovered by multiple analyses with
high support (2 97 % bootstrap support, Bayesian posterior probabilities of 100), would
render the genus paraphyletic (see Chapter 1) (Fig. 2.2). Because of the wide distribution
and economic importance of the species any taxonomic changes to X crassiusculus
should be made with strong phylogenetic support. A study of the species’ relationship to
other Xyleborina genera not considered in this analysis should be made before it is hastily
established as a monotypic genus. Furthermore, X ylosandrus crassiusculus forms a

morphologically distinct group with X hirsutipennis and any analysis of its phylogenetic

72

placement should consider this species as well. A phylogenetic study of Xyleborina
genera is presently being completed and may shed more light on this taxonomic issue
(Cognato et al. in prep.)

Several X ylosandrus species groups may require further consideration as the
phylogenetics of Xyleborina genera is resolved. The species X amputatus, X beesoni. X.
borealis, X brevis, X discolor, X diversepilosus, X jaintianus, X mancus, X
squamulatus, X subsimilis, and X subsimiliformis all form a distinct morphological
group with declivital faces that are steep and abruptly separated from the elytral disc.
This grouping was recovered in a clade with Amasa by the analysis of morphological data
with 97% bootstrap support. Phylogenetic analysis of the combined data set recovered a
subset of these species (X discolor + X mancus) as more closely related to X yleborus
aflinis (92% bootstrap support) and to a clade containing Amasa + X yleborus
califomicus (64 % bootstrap support) than to Xylosandrus. Within this species group, X
amputatus, X mancus, and X squamulatus form perhaps the most distinct group of
Xylosandrus species. These three species have lateral declivital margins with a carina or
a raised rim of granules that extends beyond the 71h declivita interstriae, forming a
circumdeclivital ring, a character often observed in Amasa. However, these species have
X ylosandrus—type antennae and pronotal-mesonotal mycangia, two characters that are
never observed in A masa. Future work on the phylogenetics of Xylosandrus and the
generic classification of Xyleborina should address these issues with more thorough

taxon sampling and the expansion of DNA data sets.

Key to the females of the species of Xylosandrus

73

3 (2)

4 (3)

5 (4)

6 (5)

Margin of elytral declivity carinate or with a raised rim of granules. ...2
Margin of elytral declivity rounded, tuberculate, or serrate but without a

continuous carina or rim. ...32

Margin of elytral declivity carinate to 7th interstriae. ...3
Margin of elytral declivity with carina extending beyond 7‘h interstriae. forming a

circumdeclivital ring. ...37

Declivital face of elytra steep and abruptly separated from disc. ...4

Elytral disc gradually curving into declivity. ...16

Declivital striae punctate. Five or six striae visible on declivity. ...5

Declivital striae granulate. Four or five striae visible on declivity. ...9

Declivital striae impressed. ...6

Declivital striae not impressed. ...7

Elytral declivity with deeply impressed striae, giving the appearance of six
distinct ridges on face. Six striae visible on declivity. Declivital striae with very
appressed, hair-like setae, shorter than the width of second declivital interstriae.
Interstriae very finely granulate, giving the declivity a matte appearance, with

erect, hair-like setae, shorter than the width of second declivital interstriae.

74

7(5)

8 (7)

Pronotum with a lateral costa, but not carinate. 1.5 — 1.6 mm long. Oriental
Region. ..X. boreensis n. sp. (Fig. 2.12)

Elytral declivity with striae less impressed. Five striae visible on declivity.
Declivital striae with erect or semi-erect, hair-like setae, longer than the width of
second declivital interstriae. Interstiae more coarsely granulate, declivity shining.
Pronotum with a lateral costa and carina. 1.3 — 1.5 mm long. Oriental Region.

...X. pygmaeus (Eggers) (Fig. 2.38)

Elytral declivity flattened. Five striae visible on declivity. ...8

Elytral declivity convex. Six striae visible on declivity. Declivital striae with
setae. Interstriae uniseriate punctate, with erect, hair-like setae, longer than twice
the width of second declivital interstriae. 1.2 — 1.8 mm long. Afrotropical
Region, Australian Region, Neotropical Region, Oceania, Oriental Region,

Palearctic Region. ...X. morigerus (Blandford) (Fig. 2.35)

Declivital striae without setae. Interstriae uniseriate granulate, with erect, hair-
like setae, longer than the width of second declivital interstriae. Larger species,
2.0 — 2.3 mm long. Oriental Region. ...X. derupteterminatus (Schedl) (Fig. 2.18)
Declivital striae without setae. Interstriae unieriate punctate, with erect, hair-like
setae, longer than the width of second declivital interstriae. Smaller species. 1.5 —

1.9 mm long. Oriental Region. ...X. terminatus (Eggers) (Fig. 2.41)

75

9 (4) Declivity covered with a dense vestiture of appressed, flattened, scale-like setae.
Striae and interstriae granulate. Pronotum granulate and pubescent basally.
Lateral pronotum costate and carinate. Frons rugose. 2.6 mm long. Oriental
Region. ...X. subsimilis (Eggers) (Fig. 2.41)

-— Declivital setae hair-like, not flattened. 10

10(9) Lateral pronotum costate and carinate. ...ll

— Lateral pronotum costate, but without a carina. 14

11 (10)Pronotum uniformly covex dorsally. Declivital face convex. Smaller species. 1.5
— 2.0 mm long. Australian Region, Oceania, Oriental Region. ..X. discolor
(Blandford) (Fig. 2.20)

— Pronotum with conspicuous summit on basal third. Declivital face flattened,

convex, or sometimes depressed in areas. Larger species, 2.8 — 3.0 mm long.

..12

12 (11)Four striae visible on elytral declivity. Declivital interstriae without a row of
longer, erect setae, bearing only a vestiture of short, appressed setae. 1 3

- Five striae visible on elytral declivity, with striae 4 and 5 forming a loop.
Declivital interstriae with a single row of long, erect, hair-like setae, along with a
dense vestiture of shorter appressed setae. 2.8 ~— 2.9 mm long. Oriental Region.

..X. beesoni Saha, Maiti, and Chakraborti (Illustrated in Saha et al. 1992)

76

13 (12) First and second declivital interstriae elevated toward apex, with depressed areas
on each side of raised interstriae. Frons rugose, with a distinct median keel. 3.0
mm long. Oriental Region, Palearctic Region. ..X. jaintianus (Schedl) (Fig.
2.27)

— Declivital face flattened, without interstriae elevated toward apex. F rons
punctate, without a distinct median keel. 2.8 mm long. Oriental Region. ...X.

subsimiliformis (Eggers) (Fig. 2.40)

14 (10) Declivital striae and interstriae granulate with appressed, hair-like setae. 15
— Declivital striae coarsely granulate, without setae. Interstriae granulate, with
erect, hair-like setae, longer than twice the width of second declivital interstriae.

1.8 — 2.3 mm long. Oriental Region. ...X. diversepilosus (Eggers) (Fig. 2.21)

15(14) Granules on interstriae dense and closely placed, giving the declivity a matte
appearance. Smaller species, 2.0 — 2.1 mm long. Oriental Region. Palearctic
Region. ..X. borealis Nobuchi (Fig. 2.11)

— Granules on interstriae less densely and closely placed, giving the declivity a
shining apprearance. Larger species, 2.5 — 2.8 mm long. Oriental Region,

Palearctic Region. ...X. brevis (Eichhoff) (Fig. 2.13)

16(3) Declivital striae and interstriae densely, finely. and confusedly granulate.

Pronotum of equal length and width. 17

77

— Declivital striae and interstriae not densely, finely, and confusedl y granulate.

Striae punctate. Pronotum wider than long. 1 8

17 (16) Pronotum with a lateral costa, but without a carina. Elytral disc multiseriate
punctate. Six striae visible on elytral declivity. Striae with erect, hair-like setae.
shorter than the width of second declivital interstriae. Interstriae with semi-
appressed, hair-like setae, longer than the width of second declivital interstriae.
Frons rugose. 1.7 — 2.9 mm long. Afrotropical Region, Australian Region,
Nearctic Region, Neotropical Region, Palearctic Region. Oceania. Oriental
Region. ...X. crassiusculus (Motschulsky) (Fig. 2.16)

— ‘ Pronotum with a lateral costa and carina. Elytral disc uniseriate punctate. Five
striae visible on elytral declivity. Striae and interstriae with semi-appressed, hair-
like setae, longer than the width of second declivital interstriae. Frons punctate.

1.9 — 2.2 mm long. Afrotropical Region. ...X. hirsutipennis (Schedl) (Fig. 2.25)

18 (16) Pronotum with a lateral costa and carina. ...19

— Pronotum with a lateral costa. but without a carina. ...27

19 (18)At least first interstriae on elytral disc multiseriate punatate. ...20

— All interstriae on elytral disc uniseriate punctate. ...21

20 (19) Elytral disc with interstriae densely punctured. Declivital interstriae multiseriate

granulate. Declivital surface matte in appearance. Stouter species, 2.0 times

78

 

longer than wide; elytra of equal length and width. 1.6 — 2.3 mm long. Oriental
Region. ...X. assequens Schedl (Fig. 2.10)

— Elytral disc with interstriae more sparely punctured, multiseriate only on first
interstriae. Declivital interstriae uniseriate granulate. Declivital surface shining.
More elongate species, 2.2 times longer than wide; elytra 1.4 times longer than

wide. 1.8 mm long. Oriental Region. ...X. deruptulus (Schedl) (Fig. 2.19)

21 (19)Declivital striae with semi-appressed, hair-like setae. shorter than the width of
second declivital interstriae. ...22

— Declivital striae without setae. ...24

22 (21) Pronotal disc glabrous, except for a dense patch of short, erect setae basally.
indicating the presence of a pronotal-mesonotal mycangium. Elytra strongly
arched from base to middle of declivity. 1.3 — 1.5 mm long. Nearctic Region.
Neotropical Region. ...X. curtulus (Eichhofl) (Fig. 2.17)

— Pronotal disc more evenly pubescent, with a dense patch of short, erect setae
basally, indicating the presence of a pronotal-mesonotal mycangium. Elytra more

evenly arched from middle of disc to apex. ...23

23 (22) Body very stout, 1.9 times as long as wide. 1.5 — 1.7 mm long. Oriental Region.

...X. pusillus (Schedl) (Fig. 2.36)

79

— Body less stout, 2.3 times longer than wide. 1.4 — 1.9 mm long. Afrotropical
Region, Nearctic Region, Neotropical Region, Oceania, Oriental Region. ...X.

compactus (Eichhoff) (Fig. 2.14)

24 (2'1)Smaller species, 1.4 mm long. Pronotum of equal length and width. Declivital
interstriae uniseriate granulate, with erect, hair-like setae, longer than the width of
second declivital interstriae. Oriental Region. ...X. mediocris (Schedl) (Fig.

2.29)

— Larger species, 1.8 — 2.5 mm long. ...25

25 (24) Pronotum wider than long or of equal length and width. Pronotal disc glabrous.
except for a dense patch of short, erect setae basally, indicating the presence of a
pronotal-mesonotal mycangium. ...26

—— Pronotum 1.1 times longer than wide. Pronotal disc more evenly pubescent, with
a dense patch of short, erect setae, indicating the presence of a pronotal-mesonotal
mycangium. 1.9 — 2.5 mm long. Nearctic Region, Oceania, Oriental Region,

Palearctic Region. ...X. germanus (Blandford) (Fig. 2.24)

26 (25)Pronotum wider than long. Declivital interstriae uniseriate granulate, with semi-
appressed, hair-like setae, longer than the width of second declivital interstriae.
2.0 mm long. Oriental Region. ...X. adherescens Schedl (Fig. 2.7)

— Pronotum of equal length and width. Declivital interstriae uniseriate granulate,

with erect, hair-like setae. longer than twice the width of second declivital

80

interstriae. 1.8 — 2.1 mm long. Oriental Region. ...X. eupatorii (Eggers) (Fig.

2.22)

27 (18)Body bicolored, with pronotum distinctly lighter than elytra or with a testaceous
patch on elytra basally and laterally. ...28

— Body uniformly colored, light to dark brown. ...31

28 (27) Pronotum and elytral apices dark brown, elytra with a testaceous patch basally
and laterally. Elytral disc with interstriae mutliseriate. Declivital interstriae
multiseriate punctate, with erect, hair-like setae, longer than twice the width of
second declivital interstriae. 2.4 — 2.7 mm long. Australian Region. ...X. hulcri
n. sp. (Fig. 2.26)

— Pronotum distinctly lighter than elytra; elytra without a testaceous patch. ...29

29 (28)Smaller species, 1.1 — 1.3 mm long. Declivital striae with setae. Interstriae with
erect, hair-like setae, longer than the width of second declivital interstriae.
Australian Region, Oriental Region. ...X. mesuae (Eggers) (Fig. 2.30)

— Larger species, 1.6 — 2.5 mm long. ...30

30 (29)Six striae visible on elytral declivity. Declivital striae with erect, hair-like setae.

longer than the width of second declivital interstriae. Interstriae punctate.

Pronotum distinctly lighter than elytra; pronotum light brown and elytra black.

81

 

Body 2.3 times longer than wide. Larger species, 2.3 — 2.5 mm long. Oriental
Region. ...X. arquatus (Sampson) (Fig. 2.9)

— Five striae visible on elytral declivity. Declivital striae without setae. Interstriae
granulate. Pronotum distinctly lighter than elytra; pronotum light brown and
elytra dark brown. Smaller species, 1.6 - 1.8 mm long. Oriental Region. ...X.

ferinus (Schedl) (Fig. 2.23)

31 (27) Five striae visible on elytral declivity. Declivital striae with semi-appressed,
hair-like setae. Interstriae multiseriate. Larger species. 2.6 — 2.7 mm long.
Australian Region. ...X. mixtus (Schedl) (Fig. 2.32)

— Six striae visible on elytral declivity. Declivital striae without setae. Interstriae
uniseriate. Smaller species, 1.8 mm long. Oriental Region. ...X. metagermanus

(Schedl) (Fig. 2.31)

32 (1) Margin of elytral declivity rounded or with a discontinuous row of small
tubercules. Elytral disc with multiseriate interstrial punctures. ...33
— Margin of elytral declivity serrate. Elytral disc with uniseriate interstrial

punctures. . . .35

33 (32) Margin of elytral declivity with a discontinuous row of small tubercles. Five

striae visible on declivity. Smaller species, 2.3 — 2.4 mm long. Australian

Region. ...X. woodi Dole and Beaver (Fig. 2.42)

82

— Margin of elytral declivity rounded. Six striae visible on declivity. Larger

species 3.0 — 4.1 mm long. ...34

34 (33)Basal pronotum lacking a dense patch of setae. Declivital striae with semi-
appressed, hair-like setae, longer than the width of second declivital interstriae.
Interstriae multiseriate granulate, with semi-appressed, hair-like setae, longer than
the width of second declivital interstriae. Smaller species, 3.0 — 3.4 mm long.
Australian Region. ...X. monteithi Dole and Beaver (Fig. 2.33)

— Basal pronotum with a dense patch of short, erect setae, indicating the presence of
a pronotal-mesonotal mycangium. Declivital striae with semi-appressed, hair-like
setae, shorter than the width of second declivital interstriae. Interstriae uniseriate
granulate, with erect, hair-like setae, longer than twice the width of second
declivital interstriae. Larger species, 3.7 — 4.1 mm long. Australian Region. ...X.

rotundicollis (Browne) (Fig. 2.39)

35 (32) Basal pronotum with a dense patch of short, erect setae, indicating the presence of
a pronotal-mesonotal mycangium. Body unicolorous. Six striae visible on elytral
declivity. Striae granulate. Smaller species, 1.6 — 2.1 mm long. ...36

—— Basal pronotum lacking a dense patch of setae. Body bicolored, pronotum
distinctly darker than elytra. Five striae visible on elytral declivity. Striae
punctate, with erect, hair-like setae, longer than the width of second declivital

interstriae. Interstriae multiseriate granulate, with erect, hair-like setae, longer

83

than the width of second declivital interstriae. Larger species, 2.7 — 3.0 mm long.

Oriental Region. ...X. corthyloides (Schedl) (Fig. 2.15)

36 (35)Elytral disc gradually curving into declivity. Declivity shining. Declivital striae

37(2)

with appressed, hair-like setae, shorter than the width of second declivital
interstriae. 1.9 — 2.1 mm long. Australian Region. ...X. abruptulus (Schedl)
(Fig. 2.6)

Declivital face steep and abruptly separated from disc. Declivity matte. Striae
with erect, acutely tapering, hair-like setae, shorter than the width of second
declivital interstriae. 1.6 — 1.9 mm long. Australian Region. ...X. queenslandi

Dole and Beaver (Fig. 2.38)

Declivital striae with a row of large, shallow punctures, arranged in a somewhat
wavy line. Declivital interstriae shining, not densely granulate. Stouter species,
1.2 — 1.4 times as long as wide. Smaller species, 2.9 — 3.3 mm long. Afrotropical
Region and Oriental Region. ...X. mancus (Blandford) (Fig. 2.28)

Declivital striae with smaller punctures arranged in perfectly straight rows.
Declivital interstriae densely and finely granulate-punctate, giving the declivity a
matte appearance. More elongate species, 2.5 times as long as wide. 2.7 — 2.9

mm long. ...X. amputatus (Blandford) n. comb. (Fig. 2.8)

Taxonomy

84

Genus X ylosandrus Reitter
Xylosandrus Reitter 1913: 80, 83. Type-species: Xylosandrus morigerus Blandford.
Apoxyleborus Wood 1980: 90. Type-species: X yleborus mancus Blandford. original
designation. Synonymy: Wood 1984: 229.
Diagnosis. Xylosandrus sensu stricto can be distinguished from other xyleborine genera
by the following combination of characters: the stout body, usually about twice as long
as wide, widely separated procoxae, flat scutellum that is flush with the surface of elytra,
and obliquely truncate antennal club with the first segment forming a circular costa and

dense pubescence on the oblique portion of the club.

X ylosandrus abruptulus (Schedl)
(Fig. 2.6)

X yleborus abruptulus Schedl, 1953: 81. Lectotype 9: Australia, Wongabel, 2 May 1941 .
A. R. Brimblecombe, from Loranthus sp.; NHMW; designated by Schedl, 1979a: 9.
Xylosandrus abruptulus (Schedl): Schedl, 1964: 213.
Notes. Schedl (1953) failed to designate a holotype in his original description of X
abruptulus and subsequently designated a lectotype (Schedl 1979a).
Diagnosis. Female 1.9 — 2.1 mm long; 2.1 times longer than wide. Body brown:
antennae and legs same color as body. F rons punctate. Antennae with 5 funicular
segments. Antenna] club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.7 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of

semi-appressed hair-like setae; setae less dense on disc. Basal pronotum with a dense

85

patch of short, erect setae, indicating the presense of a pomotal-mesonotal mycangium.
Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type 0, Hulcr et a1.
2007). Pronotum with lateral costa, not carinate. Procoxae widely separated. Protibiae
with 4 socketed teeth on lateral margin; meso- and metatibiae with 7 - 8 socketed teeth.
Elytra 1.3 times longer than wide; 1.7 times longer than pronotum. Discal striae
punctate; interstraie uniseriate punctate. Elytral disc gradually curving into declivity.
Declivity convex, lateral margin with coarse serrations. Six straie visible on declivity.
Striae granulate, with appressed, hair-like setae, shorter than the width of second
declivital interstriae. Interstriae granulate, uniseriate, with erect. hair-like setae. longer
than the width of second declivital interstriae.

This species is one of three Xylosandrus with lateral declivital marigins that are
marked by coarse serrations: X abruptulus, X corthyloides (Fig. 2.15), and X
queenslandi (Fig. 2.3 8). X ylosandrus abruptulus can be distinguished from these species
by the following characters: body unicolorous; basal pronotum with a dense patch of
short, erect setae, indicating the presence of a pronotal-mesonotal mycangium; elytral
disc gradually curving into declivity; six striae visible on elytral declivity; and declivital
striae punctate with appressed. hair-like setae. shorter than the width of second declivital
interstriae.

Distribution. AUSTRALIAN REGION: Queensland.

Hosts. Loranthus L. sp.

Specimens Examined. (12 Q; 0 (3‘)

Type material: Lectotype Xyleborus abruptulus (92; NHMW). Syntype: Australia,

Wongabel, 2 May 1941, A. R. Brimblecombe. from Loranthus sp. (S2; BMNH).

86

 

Other material: AUSTRALIAN REGION: Queensland: N. Qld., Mt Finnigan Summit
via Helenvale, 28-30 Nov 1985, G. Monteith and D. Cook, Pitfall traps, rainforest (1 Q:
RAB); N. QLD, Wallaman Falls Rd, 600 m, 14 Dec 1986 - 2 Jan 1987, Monteith.
Thompson, and Hamlet, RF, Flight intercept trap (1 Q; RAB); N. Qld., Mossman Bluff
Track, 5-10 km W Mossman, Site 9, 1260 m, 1-17 Jan 1989, Monteith, Thompson, and
Anzses, flt. intercept (2 E2; RAB); N. Qld., Mossman Bluff Track, 5-10 km W Mossman.
Site 7, 7100 m, 20 Dec 1989 - 15 Jan 1990, Monteith, Thompson. and Anzses, flt.
intercept (1 Q; RAB); N. E. QLD, Cardwell Range, Upper Broadwater Ck Valley, 750 m,
17-20 Dec 1986, Monteith, Thompson, and Hamlet, Flight intercept trap (1 Q; RAB);
NEQ: 17°26'S, 145°42'E, Hughes Road, Topaz, 650 m, 6 Dec 1993 - 25 Feb 1994,
Monteith, Cook, Janetzki, RF Pitfalls (1 9; RAB); NEQ: 16°24'Sx14501 713, Upper High
Falls Ck., 1000 m, 25 Jan - 12 Feb 1996, R. Wertz, Flight intercept trap (1 Q; RAB); N.
E. Qld, Kirrama Range, (Douglass Ck Rd, 800 m), 10 Dec 1986 - 1 1 Jan 1987, Monteith.

Thompson, and Hamlet, RF, Flight intercept trap (2 Q: RAB).

Xylosandrus adherescens Schedl
(Fig. 2.7)
Xylosandrus adherescens Schedl, 1971: 375. Holotype 9: Hui (‘7). Chuo Chan [Tonkin],
Nov. 03; NHMW.
Diagnosis. Female 2.0 mm long; 2.0 times longer than wide. Body light brown to
brown; antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;

segment one covering entire posterior face. Pronotum 0.8 times longer than wide.

87

Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presence of a pronotal-
mesonotal mycangium. Pronotal disc moderately puncatate. Lateral aspect of pronotum
basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa and carina. Procoxae
widely separated. Protibiae with 4 socketed teeth on lateral margin; meso- and metatibial
teeth not visible on specimen examined with. Elytra 1.3 times longer than wide; 1.5
times longer than pronotum. Discal striae punctate; interstraie uniseriate punctate.
Elytral disc gradually curving into declivity. Declivity convex, lateral margin carinate to
7th interstriae. Six striae visible on declivity. Striae punctate, without setae. Interstriae
granulate, uniseriate, with semi-appressed, hair-like setae, longer than the width of
second declivital interstriae.

This species is morphologically similar to X eupatorii (Fig. 2.22) and X
germanus (Fig. 2.24). These three species share the following characters: elytral disc
gradually curving into declivity; pronotum with a lateral carina; interstriae on elytral disc
uniseriate punctate; and declivital striae with semi-appressed, hair-like setae, shorter than
the width of the second declivital interstriae. X ylosandrus adherescens can be
distinguished from these species by the following characters: pronotum wider than long:
pronotal disc glabrous, except for a dense patch of short, erect setae basally, indicating
the presence of a pronotal-mseonotal mycangium; and declivital interstriae with semi-
appressed, hair-like setae, longer than the width of second declivital interstriae.
Distribution. ORIENTAL REGION: Vietnam (Tonkin Island).

Hosts. Unknown.

88

Specimens Examined. (1 Q; 0 3)

Type material: Holotype X ylosandrus adherescens (Q; NHMW).

Xylosandrus amputatus (Blandford), new combination
(Fig. 2.8)

Xyleborus amputatus Blandford. 1894b: 575. Holotype 92: Japan: Higo; BMNH.
Amasa amputatus (Blandford): Wood and Bright, 1992: 682.
Notes. This species was first included in Amasa by Wood and Bright (1992), but the
authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Diagnosis. Female 2.7 — 2.9 mm long; 2.5 times longer than wide. Body light brown to
brown; declivity distinctly darker than rest of elytra; legs and antennae the same color as
body. F rons punctate. Antennae with 5 funicular segments. Antennal club obliquely
truncate; first segment forming a circular costa; segment one covering entire posterior
face. Pronotum 1.0 — 1.1 times longer than wide. Dorsal aspect of pronotum rounded
(type 1, Hulcr et al. 2007). Pronotal vestiture of semi-appressed, hair-like setae; setae
less dense on disc. Basal pronotum with a dense patch of short, erect setae, indicating the
presence of a pronotal-mesonotal mycangium. Pronotal disc densely punctate, with
punctures separated by a distance less than or equal to their size. Lateral aspect of
pronotum basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa, not carinate.
Procoxae widely separted. Protibiae with 5 socketed teeth on lateral margin; meso- and
metatibiae with 11 socketed teeth. Elytra 1.3 times longer than wide; 2.3 — 2.4 times

longer than pronotum. Discal striae punctate; interstriae multiseriate punctate. Declivital

89

face of elytra steep and abruptly separated from disc. Declivity concave, lateral margins
carinate, with carina extending beyond 7th interstriae, forming a circumdeclivital ring.
Four striae visible on declivity. Striae punctate, without setae. Interstriae finely
granulate-punctate, without setae.

This species is one of two Xylosandrus with the margin of the elytral declivity
with a carina that extends beyond the 7th interstriae, forming a circumdeclivital ring: X
amputatus and X mancus (Fig. 2.28). X ylosandrus amputatus can be distinguished from
X mancus by the following characters: 2.5 times as long as wide; declivity without
setae; declivital striae with smaller punctures arranged in perfectly straight rows; and
declivital interstriae densely and finely granulate-punctate, giving the declivity a matte
appearance.
Distribution. ORIENTAL REGION: China, Japan, Taiwan.
Hosts. Acer L. sp., Cinnamomum mairei H. Lév., C innamomum L. sp., Machilus Nees
sp., Pelargonium hortorum L. H. Bailey, Ziziphusjujuba Lam.
Specimens Examined. (3 92; 0 6‘)
Type material: Unable to examine type material.
Other material: ORIENTAL REGION: China Gang-keu, SW. F ukien, S, China, VII-
26-36, L. Gressit Collection (1 Homotype Q; USNM); Japan: Okinawa Id, June 23.

1945, F. N. Young, No 54 (l SQ; USNM); Japan: Kagoshima Pref., Tarumizu Oonohara.

Broadleaf forest, 425 m, 14 Aug 2000, Yoshikazu Sato Coll., Ex; ETOH-baited trap (1 Q:

USNM).

Xylosandrus arquatus (Sampson)

90

an-..” i.

 

(Fig. 2.9)

Xyleborus arquatus Sampson, 1912: 246. Holotype Q: Ceylon [Sri Lanka]; BMNII.
Xylosandrus arquatus (Sampson): Schedl, 1964: 213.
Diagnosis. Female 2.3 — 2.5 mm long; 2.3 times longer than wide. Pronotum distinctly
lighter than elytra; pronotum light brown and elytra black; antennae and legs light brown.
Antennae with 5 funicular segments. Antennal club obliquely truncate; first segment
forming a circular costa; segment one covering entire posterior face. Pronotum 0.9 times O
longer than wide. Dorsal aspect of pronotum basic (type 2, Hulcr et al. 2007). Pronotal
vestiture of semi-appressed, hair-like setae; setae less dense on disc. Base of pronotum
with a dense patch of short, erect setae, indicating the presence of a pronotal-mesonotal
mycangium. Pronotal disc moderately granulate. Pronotum moderately punctuate
basally. Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotum with
lateral costa, not carinate. Procoxae widely separated. Protibiae with 4 socketed teeth on
lateral margin; mesotibiae with 8 socketed teeth; metatibiae with 8 socketed teeth. Elytra
1.4 times longer than wide; 1.5 times longer than pronotum. Discal straie punctate;
interstriae uniseriate punctate. Elytral disc gradually curving into declivity. Declivity
convex, lateral marigins carinate to 7‘h interstriae. Six striae visible on declivity. Striae
punctate, with erect, hair-like setae, longer than the width of second declivital interstriae.
Interstriae punctate, uniseriate, with erect. hair-like setae. shorter than the width of
second declivital interstraie.

This species is one of four bicolored Xylosandrus with the pronotum distinctly
lighter than the elytra: X arquatus, X discolor (Fig. 2.20), X ferinus (Fig. 2.23). and X.

mesuae (Fig. 2.30). Xylosandrus arquatus can be distinguished from X discolor by the

91

following characters: elytral disc gradually curving into declivity. X ylosandrus arquatus
can be distinguished from X ferinus and X mesuae by the following characters: 2.3 — 2.5
mm long; pronotum light brown and elytra black; six striae visible on elytral declivity:
declivital striae with erect, hair-like setae; and declivital interstriae punctate.
Distribution. ORIENTAL REGION: Sri Lanka.

Hosts. Cinnamomum L. spp., “Kududavula sp.” (Sri Lanka). Symplocos loha Buch.-
Ham. ex D. Don.

Specimens Examined. (29 Q; 0 g?)

Type material: Unable to examine type material.

Other material: ORIENTAL REGION: Sri Lanka: Ceylon [Sri Lanka] (1 Q;
NHMW); Sri Lanka: Bad. Dist., Pattopola, 200 mtrs., 3 June 1975, S. L. Wood,
Kududavula sp. (2 S9; USNM); Sri Lanka: Mat. Dist., Enselwatte, 800 mtrs., 25 May
1975. S. L. Wood, misc. hosts (1 Q; USNM); Sri Lanka: N. E. Dist., 11 km SE Nuara
Eliya, 1 June 1975, 2000 m. S. L. Wood, collected from branches (1 Q; USNM); Sri
Lanka: N. E. Dist., 11 km SE Nuara Eliya, 1 June 1975, 2000 m. S. L. Wood. Host:
Symplocos loha (18 $2; USNM); Sri Lanka: N. E. Dist., 11 km SE Nuara Eliya, 1 June

1975, 2000 m. S. L. Wood, collected from twigs (6 S2; USNM).

Xylosandrus assequens Schedl
(Fig. 2.10)
Xylosandrus assequens Schedl, 1971: 376. Holotype $2: Malaya, Kelantan, Bukit

Kabong, 14.ii.1947, in Xanthophyllum sp., F. G. Browne; BMNH.

92

 

Diagnosis. Female 1.6 —— 2.3 mm long; 2.0 times longer than wide. Body brown to dark
brown; antennae and legs light brown. Frons rugose. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.7 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Pronotum punctate basally;
lacking a dense patch of setae. Pronotal disc densely punctate, with punctures separated
by a distance less than or equal to their size. Lateral aspect of pronotum basic (type 0,
Hulcr et al. 2007). Pronotum with lateral costa and carina. Procoxae widely separated.
Protibiae with 4 socketed teeth on lateral margin; mesotibiae with 9 socketed teeth;
metatibiae with 10 socketed teeth. Elytra 1.3 times longer than wide; 1.7 times longer
than pronotum. Discal striae punctate; interstriae multiseriate puncate; striae and
interstriae confused on disc. Elytral disc gradually curving into declivity. Declivity
convex, lateral margins carinate to 7th interstriae. Six striae visible on declivity. Striae
punctate, with erect, hair-like setae, longer than the width of second declivital interstriae.
Interstriae granulate, multiseriate, with erect, hair-like setae, longer than the width of
second declivital interstriae.

This species is morphologically similar to X deruptulus (Fig. 2.19). Xylosandrus
assequens can be distinguished from X deruptulus by the following characters: 2.0 times
longer than wide; elytra of equal length and width; elytral disc with interstriae densely
punctured; declivital interstriae multiseriate; and declivital surface matte in appearance.
Distribution. ORIENTAL REGION: Brunei, Malaysia.

Hosts. Xanthophyllum Roxb. sp.

93

Specimens Examined. (7 S9; 0 6‘)

Type material: Holotype Xylosandrus assequens (Q; BMN H).

Other material: ORIENTAL REGION: Brunei: BRUNEI: Temburong: Nr. K.
Belalong Field Study Centre 150 m, 4°33'N 155°09'E, 5-8.iii.1992, R. A. Beaver.
RGS/UBD Exped. Pitfall trap (1 Q; RAB); BRUNEI: Temburong: Nr. K. Belalong Field
Study Centre 100 m, 4°33'N 155°09'E, 12-14.iii.1992, R. A. Beaver, RGS/UBD Exped. (2
Q; RAB); BRUNEI: Temburong: Nr. K. Belalong Field Study Centre 150 m, 4°33'N
155°09'E, 22-24.ii.l992, R. A. Beaver, RGS/UBD Exped. Pitfall trap (1 SB: RAB);
BRUNEI: Temburong: Nr. K. Belalong Field Study Centre 150 m, 4°33'N 155°09'E, 26-
29.ii.1992, R. A. Beaver, RGS/UBD Exped. Pitfall trap (1 £2; RAB); Malaysia:
MALAYSIA: Sabah Sipitang, Mendolong, T6/R, 31.iii.1989, leg. S. Abdebratt (1 Q;

RAB).

X ylosandrus beesoni Saha, Maiti, and Chakraborti
(Illustrated in Saha et al. 1992)

Xylosandrus beesoni Saha, Maiti, and Chakraborti, 1992: 11. Holotype 9: India:
Rangirum (1846 m), Darjiling Dist., coll. J. C. M. Gardner, 8.ix.1929, ex. “Symplocos
theaefolia ” [Symplocos theifolia].
Diagnosis. Female 2.8 — 2.9 mm long; 2.0 times as long as wide. Body yellowish
brown. Frons reticulate. Antennae with 5 funicular segements. Antennal club obliquely
truncate; frist segment forming a circular costa; segment one covering entire posterior
face. Pronotum wider than long. Dorsal aspect of pronotum rounded (type 1, Hulcr et al.

2007). Pronotal vestiture of semi-appressed, hair-like setae; pronotal disc densely setose.

94

setae as dense as on anterior pronotum. Basal pronotum with a dense patch of short, erect
setae, indicating the presence of a pronotal-mesonotal mycangium. Pronotal disc densely
asperate-granulate, with sculpture separated by a distance less than or equal to their size.
Lateral aspect of pronotum basic (type 0, hulcr et al. 2007). Pronotum with lateral costa
and carina. Procoxae widely separated. Protibiae with 5 socketed teeth on lateral
margin; meso- and metatibiae with 8 socketed teeth. Elytra longer than wide; 1.4 times
longer than pronotum. Discal striae punctate; interstriae multiseriate punctate. Declivital
face of elytral steep and abruptly separated from disc. Declivity convex, lateral margins
carinate to 7th interstriae. Five striae visible on declivity with striae 4 and 5 forming a
loop. Striae granulate, with semi-appressed, hair-like setae, shorter than the width of
second declivital interstriae. Interstriae granulate, multiseriate, with a single row of long
hair-like setae along with a dense vestiture of shorter appressed setae.

This species is morphologically similar to X. discolor (Fig. 2.20), X jaintianus
(Fig. 2.27), and X subsimiliformis (Fig. 2.40). Xylosandrus beesoni can be distinguished
from these species by the following characters: declivital face covex, five striae visible
on declivity with striae 4 and 5 forming a loop, and declivital interstriae with a single row
of long, erect, hair-like setae, along with a dense vestiture of shorter appressed setae.
Distribution. ORIENTAL REGION: India.
Hosts. Symplocos theifolia D. Don.
Specimens Examined. (0 SB; 0 6)
Type material: Unable to examine type material.
Discussion. This species was described by Saha, Maiti, and Chakraborti (1992) from 4

female specimens from a single collecting event. All specimens are deposited in

95

collections in India (FRI and ZSI) and were not available for examination as part of this
revision. However, based on the original species description and corresponding images.
this X beesoni is clearly a member of the X ylosandrus senso stricto group. The species
description was also detailed enough that it could be utilized to score morophological

characters for the phylogenetic analysis.

X ylosandrus borealis Nobuchi
(Fig. 2.11)

Xylosandrus borealis Nobuchi, 1981: 34. Holotype Q: Honshu, Kyushu (Japan);
Nobuchi Collection, Ibaraki, Japan.
Diagnosis. Female 2.0 — 2.1 mm long; 1.8 — 1.9 times as long as wide. Body yellowish
brown to light brown; antennae and legs the same color as body. Frons rugose with a
distinct median keel. Antennae with 5 funicular segments. Antennal club obliquely
truncate; first segment forming a circular costa; segment one covering entire posterior
face. Pronotum 0.9 times longer than wide. Dorsal aspect of pronotum rounded (type 1.
Hulcr et al. 2007). Pronotal vestiture of semi-appressed, hair-like setae; pronotal disc
densely setose, setae as dense as on anterior pronotum. Basal pronotum with a dense
patch of short, erect setae, indicating the presence of a pronotal-mesonotal mycangium.
Pronotal disc densely asperate-granulate, with sculpture separated by a distance less than
or equal to their size. Lateral aspect of pronotum prolonged anteriorly (type 9, Hulcr et
al. 2007). Pronotum with lateral costa, not carinate. Procoxae widely separated.
Protibiae with 4 socketed teeth on lateral margin; meso- and metatibiae with 8 socketed

teeth. Elytra 1.0 — 1.3 times longer than wide; 1.1 times longer than pronotum. Discal

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striae punctate; interstriae multiseriate punctate. Declivital face of elytral steep and
abruptly separated from disc. Declivity flattened, lateral margins carinate to 7th
interstriae. Four striae visible on declivity. Striae granulate, with appressed, hair-like
setae, shorter than the width of second declivital interstriae. Interstriae granulate,
multiseriate, with appressed, hair-like setae shorter than the width of second declivital
interstriae.

This species is morpholocially similar to X brevis (Fig. 2.13). X ylosandrus
borealis can be distinguished from X brevis by the matte appearance of its elytral
declivity, due to dense and closely placed interstrial granules, and by its smaller size of i
2.0 — 2.1 mm.

Distribution. ORIENTAL REGION: Japan. PALEARCTIC REGION: Korea.
Hosts. Camellia sasanqua Thunb., Styrax obassia Siebold and Zucc.

Specimens Examined. (4 $2; 0 6)

Type material: Unable to examine type material.

Other material: ORIENTAL REGION: Japan: Okusa, Japan, 20.VII.1935, Coll. K.

Baba (4 $2; USNM).

Xylosandrus borneensis Dole and Cognato, new species
(Fig. 2.12)
Description. Female (Fig. 2.12): 1.5 — 1.6 mm long, 2.1 times longer than wide, with
pronotum yellowish brown, slightly darker toward apex, elytra brown, ventral side and
appendages yellowish brown. F rons convex, shining, coriaceous, punctate between eyes.

with deep, vertically elongate punctures, a single hair-like setae originating from each

97

puncture. Epistoma with row of short hair-like setae along lower margin. Eyes
emarginate. Antennal funicle 5-segmented, scape and funicle with sparse, short, hair-like
setae; club obliquely truncate, first segment sclerotized, forming a circular costa (type 1.
Hulcr et al. 2007), circular costa closed anteriad, oblique part of club densely pubescent,
second segment not comeous; posterior face of club covered entirely by first segment
(type 1, Hulcr et al. 2007). Pronotum of equal length and width, basic shape dorsally
(type 2, Hulcr et al. 2007), widest about two-thirds pronotal length from base, anterior
third broadly rounded toward apex, basal angles rounded, anterior margin with 8-10
asperities; anterior slope aspirate, asperities smallest at summit and increasing in size
toward anterior margin; disc moderately punctate, patch of denser punctures medially at
base, hair-like setae originating from punctures, background sculpture finely granulate;
lateral aspect of pronotum basic (type 0, Hulcr et al. 2007), lateral costa extending two-
thirds pronotal length, lateral carina present only in basal ‘A of prontum; pronotal
vestiture of short, semi-appressed, hair-like setae. Scutellum triangular, flush with
surface of elytra. Elytra 1.1 times longer than wide, 1.1 times as long as pronotum,
parallel-sided on basal two-thirds and then broadly rounded toward apex; disc shining;
declivity matte in appearance. Striae impressed beginning slightly before declivital
origin and becoming more deeply impressed on declivity, shallowly and regularly
punctate, punctures becoming less distinct on declivity, with very short, appressed setae
originating from punctures. Interstriae equal the width of striae, finely granulate. giving a
matte appearance, short (less than or equal to width of interstria), erect, hair-like setae in
uniseriate rows; interstriae 4 - 6 not reaching apex of the declivity. Declivity

commencing behind mid-point of elytra, abruptly and steeply separated from disc; lateral

98

margins carinate to 7th interstriae. Procoxae widely separated. Protibiae with 4 socketed
teeth on lateral margin; mesotibiae with 7 - 8 socketed teeth; metatibiae with 8 - 9
socketed teeth. Abdominal ventrites evenly punctured, punctures with short and long.
moderately appressed, hair-like setae.
Specimens Examined. (11 92; 0 6)
Type Material. Holotype Q: MALAYSIA, Sabah, Danum Valley, 120 m asl. July 2006
Hulcr coll. Burseraceae, twig. Vial 1694. In MSU.
Paratypes Q: MALAYSIA, Sabah, Danum Valley, 120 m asl. July 2006 Hulcr coll.
Burseraceae, twig. Vial 1694. One paratype in F RCS. Nine paratypes in MSU. One
paratype female was used for DNA extraction and remains are vouchered in MSU.
BRUNEI [DARUSSALAM]: Temburong: nr. K[uala] Belalong Field Stud[ies]. Centre.
4.33'N 115.09'E, 150m, 21.ii.1992 (R.A.Beaver). One paratype in RAB. MALAYSIA:
Sabah, Sipitang, Mendolong, 11.v.1988 (2); 10.iii.1989 (l); 14.iii.1989
(2) (S.Adebratt).Three paratypes in MZLU; two in RAB.

Male: Unknown
Etymology. This species is named borneensis after the type locality.
Diagnosis. This species can be distinguished from all other known Xylosandrus by the
deeply impressed declivital striae, which form six distinct ridges on the declivity. It is
morphologically similar to X pygmeaus (Fig. 2.3 8). Xylosandrus borneensis can be
distinguished from X pygmaeus by the following characters: pronotum without lateral
carina; declivital striae more deeply impressed; six striae visible on elytral declivity;

declivital striae without setae; and declivital interstriae finely granulate, giving the

99

declivital face a matte appearance, with erect, hair-like setae, shorter than the width of
second declivital interstriae.

Distribution. ORIENTAL REGION: Malaysia (Sabah).

Hosts. Burseraceae.

Discussion. This species was found in association with the mycocleptes species

Xyleborus mucronatulus Eggers (Hulcr 2008).

Xylosandrus brevis (Eichhoff)
(Fig. 2.13)

Xyleborus brevis Eichhoff, 1877: 121. Syntypes SB: Nipon (Hagi Hiller) and Nipon lnsula
Asiatica; IRSNB.
Xylosandrus brevis (Eichhofi): Browne, 1965: 204.
Xyleborus cucullatus Blandford, 1894c: 121. Syntypes Q: Kurigahara. andKonose in
Higo, Japan; BMNH. Synonymy: Murayama 1954: 176.
Diagnosis. Female 2.5 — 2.8 mm long; 2.1 times longer than wide. Body dark brown;
antennae and legs light brown. Frons rugose, with distinct median keel between eyes.
Antennae with 5 funicular segments. Antennal club obliquely truncate; first segment
forming a circular costa; segment one covering entire posterior face. Pronotum 0.9 times
longer than wide. Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007).
Pronotal vestiture of dense, semi-appressed, hair-like setae; pronotal disc densely setose,
setae as dense as on anterior pronotum. Basal pronotum with a dense patch of short, erect
setae, indicating the presense of a pronotal-mesonotal mycangium. Pronotal disc densely

asperate-granulate, with sculpture separated by a distance less than or equal to their size.

100

Lateral aspect of pronotum prolonged anteriorly (type 9, Hulcr et al. 2007). Pronotum
with lateral costa, not carinate. Procoxae widely separated. Protibiae with 4 socketed
teeth on lateral margin; meso- and metatibiae with 10 — 11 socketed teeth. Elytra 1.1
times longer than wide; 1.2 times longer than pronotum. Discal striae punctate;
interstraie multiseriate punctate. Declivital face of elytra steep and abruptly separated
from disc. Declivity convex, lateral margins carinate to 7‘h interstriae. Four striae visible
on declivity. Striae coarsely granulate, with appressed, hair-like setae, shorter than the
width of second declivital interstriae. Interstriae granulate, multiseriate, with appressed,
hair-like setae, shorter than the width of second declivital interstriae.

This species is morpholocially similar to X borealis (Fig. 2.1 1). Xylosandrus
brevis can be distinguished from X borealis by the shining appearance of its elytral
declivity and by its larger size of 2.5 — 2.8 mm.

Distribution. ORIENTAL REGION: Japan , Taiwan, Thailand. PALEARCTIC
REGION: China (Xizang [Tibet]), Korea, Nepal.

Hosts. Berberis L. sp., Camellia japonica L., C. sasanqua Thunb., Cinnamomum
japonicum Siebold, Diospyros kaki Thunb., F agus crenata Blume, Grevillia Knight sp.,
Hamamelis L. sp., Lindera Thumb. spp., Machilusjaponica Siebold and Zucc., Machilus
thunbergii Siebold and Zucc., Maesa tenera Mez, Meliosma cuneifolia F ranch.,
Parabenzoin praecox Nakai, Quercus L. spp., Smilax china L., Styrax obassia Siebold
and Zucc., Viburnum L. sp., Weigela hortensis C. A. Mey.

Specimens Examined. (25 Q; 0 6‘)

Type material: Unable to examine type material.

101

Other material: ORIENTAL REGION: Japan: Japan: Kagoshima Pref, Tarumizu
Oonohara, Broadleaf forest, 425 m, 14 May 2001, Yoshikazu Sato Coll. Ex; ETOH-
baited trap (4 E2; MSU); Japan, Ookusa, 20.VI.1933. Coll. K. Baba (3 Q; USNM);
JAPAN: RYUKYUS, Mt. Yonaha-dake, Okinawa-honto Is., 5.VI.1997, H. Goto leg..
Host tree: Michilusjaponica sleb. Et Zucc. (4 92; RAB); Japan, Tamagowa. 29.Vll.1980.
Fagus crenatus (3 E2; USNM); Japan, Tokyo, Takao, Vll.31.1957, Coll. A. Nobuchi (4
Q; USNM). Taiwan: TAIWAN: Taichung, Hsien: Shei-Pa N.P., 10.5.2004, J—T. Yang.
Pitfall (5 SQ; RAB). PALEARCTIC REGION: Nepal: Nepal: Sikha 83°40'E, 28°26'N.

8000 ft. 24—26.v.1954. K. H. Hyatt. Litter in oak forest (2 SB; BMNH).

X ylosandrus compactus (Eichhoff)
(Fig. 2.14)

Xyleborus compactus Eichhoff, 1875: 201. Syntypes 2 Q, 1 6‘: Japan; NHMW (syntypes
in Hamburg Museum lost).
Xylosandrus compactus (Eichhoff): Nunberg, 1959: 434.
.Xyleborus morstatti Hagedom, 1912: 37. Syntypes SE: Amani, Deutsch-Ostafrika;
Hamburg Museum, lost.
Xylosandrus morstatti (Hagedom): Browne, 1963255. Synonymy: Murayama and
Kalshoven, 1962: 247.
Diagnosis. Female 1.4 — 1.9 mm long; 2.3 times longer than wide. Body brown to dark
brown; antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;

segment one covering entire posterior face. Pronotum of equal length and width. Dorsal

102

aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-
appressed hair-like setae; setae less dense on disc. Basal pronotun with a dense patch of
short, erect setae, indicating the presence of a pronotal-mesonotal mycangium. Pronotal
disc moderately punctate basally. Lateral aspect of pronotum basic (type 0, Hulcr et al.
2007). Pronotum with lateral costa and carina. Procoxae widely separated. Protibiae
with 4 socketed teeth on lateral margin; mesotibiae with 7 socketed teeth; metatibiae with
8 socketed teeth. Elytra 1.3 times longer than wide; 1.3 times longer than pronotum.
Discal striae punctate; interstriae uniseriate punctate. Elytral disc gradually curving into
declivity. Declivity convex, lateral margins carinate to 7th interstriae. Six striae visible
on declivity. Striae punctate, with semi-appressed, hair-like setae, shorter than the width
of second declivital interstriae. Interstriae punctate-granulate, uniseriate, with erect, hair-
like setae, longer than twice the width of second declivital interstriae.

This species is morphologically similar to X curtulus (Fig. 2.17) and X pusillus
(Fig. 2.36). It can be distinguished from X curtulus by a pronotal disc that is evenly
pubescent, rather than mostly glabrous. X ylosandrus compactus is nearly
morphologically identical to X pusillus. The only character distinguishing the two
species is the degree of body stoutness, with X compatcus being 2.3 times as long as
wide and X pusillus being 1.9 times as long as wide. However, this is too large a
disparity to warrant synonymizing the two species without further investigation.
Distribution. AF ROTROPICAL REGION: Cameroon, Cameroon Islands (Grande
Comoro), Equatorial Guinea, Fernando Poo, Gabon, Ghana, Ivory Coast, Liberia,
Madagascar, Mauritania, Nigeria, Reunion Islands, Senegal, Seychelles Islands, Sierra

Leone, South Afi'ica, Tanzania, Uganda. NEARCTIC REGION (Introduced) United

103

States (Alabama, Florida, Louisiana, Mississippi North Carolina, East Texas).
NEOTROPICAL REGION: Brazil, Cuba, Peru, Peutro Rico, Virgin Islands.
OCEANIA: Hawaiian Islands (Hawaii, Kauai, Lanai, Maui, Molokai, Oahu), Samoan
Islands. ORIENTAL REGION: Bonin Islands, China (Guangdong), India (Tamil
Nadu), Indonesia (Java), Japan, Malaya, Malaysia (Sabah), Philippine Islands, Ryukyu
Islands, Sri Lanka, Taiwan, Thailand, Vietnam (Tonkin Island).

Hosts. Acacia mangium Willd., Acalypha L. sp., Acer L. sp., Acrocarpusfi'axinifolius
Wight and Am., Adenanthera pavonina L., Albizzia chinensis Merr., A. lebbeck Benth..
A. zygia (DC.) Macbride, Ardisia paniculata Roxb., Aucoumea klaineana Pierre,
“Bamboo Orchid” (Hawaii), Bauhinia L. sp., 8. tomentosa L., Bombax malabaricum
DC ., Cajanus cajan (L.) Millsp., Camellia sinensis Kuntze, Cassia hirta Wild., C.
mutijuna Rich., C. siamea Lam., C. tora L., Cattleya Lindl. sp., C innamomum camphora
(L.) J. Presl, C. iners Reinw., C. zeylanicum Broyn., Clerodendron Burm. sp., Coflea L.
sp., C. arabica L., C. bukobensis Zimmerrnann, C. canephora Pierre, C. liberica Bull...
C. quillon Wester, C. robusta L. Lind., C. stenophylla G. Don., Cola nitida (Vent) Schott
and End., Crotalaria L. sp., Cryptocaria Gay sp., Dendrobium Sw. sp., D. phalaenopsis
Fitz., D. veratrifolium Lindl., Desmodium ovalifolium Guill., Drypetes phyllanthoides
(Rock) Sherff, Elaeis guineensis J acq., “Elderberry” (Singapore), Enthandrophragma
utile Sprague, Erythroxylon novagranatense Hieron., Eupatorium pallescens DC.,
Eusideroxylon zwageri Teij sm., Ficus aurea Nutt., F. soroceoides Bar., Gossypium L.
sp., Haasia Nees. sp., Hopea parviflora Bedd., Ichthyomethia communis S. F. Blake,
Indigofera sujj’ruticosa Mill., Jacobinia Moric. sp., Khaya grandifoliola C. DC ., K.

senegalensis A. Juss., Leucaena glauca Benth.. Litsaea cassiaefolia Blume. Mangifera

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indica L., M odorata Griff, Melia azedarach L., Muntingia calabura L., Myrciaria
dubia (H. B. and K.) McVaugh, Nectandra angustifolia Nees. and Mart., Olea europaea
L., Persea Mill. sp., Persa Americana Miller, P. gratissima Gaertn., P. indica Sprang..
“Prosopis nudiflora” (Java: Schedl 1962), Quercus myrsinaefolia Blume, Rhizophora L.
sp., Sambucus L. sp., S. canadensis L., S. javanica Reinw., Shorea Roxb. ex. C. F.
Gaertn. sp., S. sumatrana (Slooten) Desch., Spathodea campanulata P. Beauv., Swietenia
macrophylla King, S. mahagoni Jack., Tephrosia maxima Pres., Thea sinensis L.,
Theobroma cacao L., Toona sureni Merr., Turraeanthus africana Pellegr., Vanda coerula
Griff, V. teres Lindl., V. tricolor Lindl., Vitex L. sp., “Wild Grape” (Florida)..
Specimens Examined. (123 SE; 8 6)

Type material: Syntypes Xyleborus compactus (2 Q, 1 6; NHMW).

Other material: AFROTROPICAL REGION: Ghana: Ghana: Kumasi, 12.9.61 (6 $2.
1 6; USNM). Madagascar: Madagascar: Prov. F lanarantsoa, 7 km W Ranomafana,
1100 m, 1-7 November 1988, W. E. Steiner (1 Q; USNM). NEARCTIC REGION:
United States: Florida: Dade Co. Coral Gables, Matheson Hammock Pk., 27 June 1980.
O’Brien and Wibmer (1 9; CAS); Florida: Key Largo, 6.25.1951, Price Beamers —
Wood, Ichthyomethia communis (1 SB; USNM); Florida: Key Largo, 6.25.1951, Price
Beamers — Wood (4 Q, 1 6; USNM); Florida: Key Largo, 6.25.1951, Price Beamers —
Wood, taken on wild grape (7 Q; USNM); Florida: Key Largo, 6.25.1951, Price Beamers
—- Wood, Ardisia paniculata (4 S2; USNM); Florida: Key Largo, 6.25.1951, Price
Beamers — Wood, Cajanus cajan (6 S2; USNM); Florida: Key Largo, 6.25 .1951, Price
Beamers -— Wood, taken on Ficus aurea (1 Q; USNM); Florida: Tallahassee, Fall, 1979,

C. W. O’Brien, reared ex dogwood flags. emer. Sum 1980 (23 SB; CAS); Florida. W.

105

Palm Beach, 1.27.61 (6?; USNM). NEOTROPICAL REGION: Peru: Perou-Loreto.
Iquitos, Juin 1990, G. Couturier Col., Plante — Héte, Myrciaria dubia (1 9; USNM).
Peurto Rico: Peurto Rico: Carite St. F or., VII.28.1999, C. W. O’Brien, P. Kovarik (1 9:
CAS). OCEANIA: Hawaiian Islands: Hawaii: Mt. Puu Puae, Waianae Mts.. C ahu 425
m., in twigs of Drypetes phyllanthrides, 1.VII.1970. W. C. Gagne Collector (1 Q;
NHMW); Hawaii: Oahu, Kailua, I. 1962, Ex Vitex sp. (10 9; CAS); Hawaii: Oahu,
Kailua, 1262., ex. Vitex sp. Roy Hirata C011. (2 SB, 3 6; USNM); Hawaii: Oahu, Nuuanu,
May 8, 1931, H. L. Lyon, Elderberry stems imported from Singapore (1 S2; USNM);
USA: HI: O’ahu 1., N. Halawa Valley, NW of Honolulu, 390 m, 21025’N, 157051 ’W,
11-29.VI.1991, FMHD#91-4, Met. Polym.-Psidium-Hibiscus mixed nat./exotic forest, A.
Newton and M. Thayer, #869, window trap (4 92; F MNH); Quarantine from Hawaii at
Carpenteria, California, VI.20.2001, Ex. Bamboo Orchid (2 Q; CSCA). ORIENTAL
REGION: India: India: Coffee Research Station, Chilemagalur dist., Mysore, 2.1.1966
(7 S2; CAS); South India: Nilgiri Hills, Devala, 3200 ft., Xl.60, P. S. Nathan (23 Q; 2
6; USNM). Indonesia: Java: Boger, VIII.1964, N. L. H. Krauss, Coffee (1 Q; USNM);
E. Java, Ma Lang, 10.1951, Planta nutrix, Coffea (4 Q; USNM). Sri Lanka: Sri Lanka:

Col. Dist., Labugama, 23 June 1975. S. L. Wood, collected from branches (1 EB; USNM).

X ylosandrus corthyloides (Schedl)
(Fig. 2.15)
Xyleborus corthyloides Schedl, 1934: 86. Lectotype 9: Java, Mount Gede, 800 m. ex
Zingiberaceae; NHMW; designated by Schedl, 1979a: 66.

Xylosandrus corthyloides (Schedl): Wood and Bright 1992: 790.

106

Xyleborus percorthyloides Schedl, 1957: 85. Lectotype S2: Java, Mount Gede, 800 m;
NHMW; designated by Schedl, 1979a: 66. Synonymy: Wood and Bright 1992: 790.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors failed to indicate it as a “new combination.” X yleborus percorthyloides was
also listed as a synonym (an “uneeded replacement of corthyloides”) by Wood and Bright
(1992), but was not indicated as a “new synonymy.”

Diagnosis. Female 2.7 — 3.0 mm long; 2.1 times longer than wide. Body bicolored,
pronotum distinctly darker than elytra; antennae and legs yellowish brown. F rons
punctate. Antennae with 5 funicular segments. Antennal club obliquely truncate; first
segment forming a circular costa; segment one covering entire posterior face. Pronotum
0.9 times longer than wide. Dorsal aspect of pronotum rounded (type 1, Hulcr et al.
2007). Pronotal vestiture of semi-appressed, hair-like setae; setae less dense on disc.
Lacking a dense patch of setae at base of pronotum. Pronotal disc moderately punctate.
Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa.
not carinate. Procoxae widely separated. Protibiae with 4 socketed teeth on lateral
margin; mesotibial teeth not visible on specimens examined; metatibiae with 10 socketed
teeth. Elytra 1.1 times longer than wide; 1.2 times longer than pronotum. Discal striae
punctate; interstriae uniseriate punctate. Declivital face of elytra steep and abruptly
separated from disc. Declivity flattened, lateral margins with small serrations. Five
striae visible on declivity. Striae punctate, with erect, hair-like setae, longer than the
width of second declivital interstriae. Interstriae granulate, multiseriate. with erect, hair-

like setae, longer than the width of second declivital interstriae.

107

This species is one of three Xylosandrus with lateral declivital marigins that are
marked by coarse serrations: X abruptulus (Fig. 2.6), X corthyloides, and X queenslandi
(Fig. 2.3 8). Xylosandrus corthyloides can be distinguished from these species by the
following characters: 2.7 — 3.0 mm long; basal pronotum lacking a dense patch of setae:
body bicolored, pronotum distinctly darker than elytra; five striae visible on elytral
declivity; and declivital striae punctate.

Distribution. ORIENTAL REGION: Indonesia (Java).

Hosts. Lingiberaceae, Zingiberaceae.

Specimens Examined. (2 92; l 6)

Type material: Holotype Xyleborus corthyloides (SQ; NH MW). Paratype X yleborus
percorthyloides Java: Mt. Gede, l4-IX-1922, 800 m, L. G. E. Kalshoven, Lingiberaceae
(Q; NHMW). Allotype Xyleborus percorthyloides Java: Mt. Gede, 14-IX-1922. 800 m.

L. G. E. Kalshoven , Lingiberaceae (6; NHMW).

X ylosandrus crassiusculus (Motschulsky)
(Fig. 2.16)

Phlaeotrogus crassiusculus Motschulsky, 1866: 403. Syntypes 9: published as Des
Montagnes de Nura-Ellia, Ceylon types labeled India Occidentale; IZM.
Xylosandrus crasssiusculus (Motschulsky): Wood, 1982: 766.
Xyleborus semiopacus Eichhoff, 1878: 334. Syntypes Q: Nipom insula Japonica;
Hamburg Museum, lost. Synonymy: Wood, 1969: 119.
Xyleborus semigranosus Blandford, 18 96a: 211. Holotype Q: Sumatra; BMNH.

Synonymy: Schedl 1959: 496.

108

Xyleborus ebriosus Niisima, 1909: 154. Holotype SB: Sapporo, Japan; Nobuchi
Collection, Ibaraki. Synonymy: Choo, 1983: 98.

Dryocoetes bengalensis Stebbing, 1908: 12. Syntypes Q: Goalpara. Assam; FRI.
Synonymy: Beeson, 1915: 297.

Xyleborus mascarenus Hagedom, 1908: 379. Syntypes Q: Mauritius, and Bomole et
Amani in Deutsch-Ostafrika; NHMW (syntypes in Hamburg Museum lost). Synonymy:
Eggers, 1923: 130.

Xyleborus okoumeensis Schedl, 1935: 271. Syntypes Q: imported Okoume logs,
Carlshafen, Hessen-Nassau [Germany]; NHMW. Synonymy: Schedl, 1959: 496.
Xyleborus declivigranulatus Schedl, 1936: 30. Lectotype 92: Selangor, Malay Peninsula;
NHMW; designated by Schedl, 1979a: 76. Synonymy: Schedl, 1959: 496.

Diagnosis. Female 1.7 — 2.9 mm long; 2.2 times longer than wide. Body light to dark
brown; antennae and legs yellowish brown. F rons rugose. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum of equal length and width. Dorsal
aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-
appressed, hair-like setae; pronotal disc densely setose, setae as dense as on anterior
pronotum. Basal pronotum with a dense patch of short, erect setae, indicating the
presence of a pronotal-mesonotal mycangium. Pronotal disc densely punctate, with
punctures separated by a distance less than or equal to their size. Lateral aspect of
pronotum basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa, not carinate.
Procoxae widely separated, though less so than in most Xylosandrus species. Protibiae

with 4 socketed teeth on lateral margin; mesotibiae with 11 socketed teeth; metatibiae

109

with 14 — 16 socketed teeth. Elytra 1.2 times longer than wide; 1.2 times longer than
pronotum. Discal striae punctate; interstriae multiseriate punctate. Elytral disc gradually
curving into declivity. Declivity convex, cariante to 7th interstriae. Six striae visible on
declivity. Striae granulate, with erect, hair-like setae, shorter than twice the width of
second declivital interstriae. Interstriae granulate, multiseriate, with semi-appressed.
hair-like setae, longer than the width of second declivital interstriae.

This species is morphologically similar to X hirsutipennis (Fig. 2.25). Both
species have a declivital face that is matte in appearance due to densely, finely, and
confusedly granulate striae and interstriae. X ylosandrus crassiusculus can be
distinguished from X hirsutipennis by the following characters: frons rugose; pronotum
without a lateral carina; elytral disc multiseriate punctate; six striae visible on the elytral
declivity; declivital striae with erect, hair-like setae shorter than the width of second
declivtial interstriae; and declivital interstriae with semi-appressed, hair-like setae. longer
than the width of second declivital interstriae.

Distribution. AF ROTROPICAL REGION: Cameroon, Congo, Equatorial Guinea.
Fernando Poo, Gabon, Ghana, Ivory Coast, Kenya, Madagascar, Mauritania, Nigeria,
Sierra Leone, Seychelles Islands, Tanzania, Zaire. AUSTRALIAN REGION: New
Guinea. NEARCTIC REGION (Introduced): United States (Alabama, Delaware.
Florida, Georgia, Kansas, Louisiana, Maryland, Mississippi, North Carolina, Oregon,
South Carolina, Tennessee, Texas). NEOTROPICAL REGION: Costa Rica, Panama.
OCEANIA: Hawaiian Islands (Hawaii, Kauai, Maui, Oahu), Micronesia (Palau Islands),
New Caladonia, Samoan Islands. ORIENTAL REGION: Bonin Islands, Burma, Hong

Kong (F ujian), China (Hunan, Sichuan), India (Andaman Islands. Assam. Bengal.

11.0

Himachal Pradesh, Madhya Pradesh, Maharashtra, Tamil Nadu, Uttar Pradesh), Indonesia
(Borneo, Celebes, Java, Sumatra), Japan, Malaya, Malaysia (Sabah, Sarawak), Philippine
Islands, Sri Lanka, Taiwan, Thailand, Vietnam. PALEARCTIC REGION: Bhutan.
China (Xizang [Tibet]), Germany, Korea, Nepal.

Hosts. Acacia Mill. sp., A. decurrens Willd., A. mangium Willd., Acrocarpus J. R. F orst
and G. Forst spp., Adina rubescens Hemsl., Adinandra dumosa Jack., Adinobotrys
atropurpureus Dunn., Afzelia bipindensis Harms, A gathis Salisb. sp., Albizia Durazz. sp..
A. chinensis Merr., A. ferruginea Benth., A. gummifera (Gmel.) C. A. Sm., A. lebbek L.,
A. moluccana Miq., A. stipilata Boivin, A. zygia (DC.) Macbride, Alnus Mill. sp.,
Alstonia R. Br. sp., Altingia excelsa Nor., A moora Roxb. sp., Angylocalyx pynaertii De
Wild., Anisoptera Korth. sp., Anthonotafiagrans Baker, Antiaris africana Engl.,
Antrocaryon micraster A. Chev. and Guill., Artocarpus J. R. Forst. and G. Forst. sp., A.
chaplasha Roxb., Aucoumea klaineana Pierre, Barteria nigritiana Hook., Bauhinia
tomentosa L., Bischofiajavanica Blume, Buchanania arborescens Blume, B. sessilifolia
Blume, Cacao Mill. sp., Calamus L. sp., Calophyllum tetrapterum Miq., Camelia
sinensis (L.) O. Kuntze, C. thea Link., Canarium L. sp., Cannabis sativa L., Carapa
procera DC., Carica papaya L., Carya illinoinensis (Wangenh.) Koch, Caryota L. sp.,
Castanea argentea Bl., C. javanica Blume, Castanopsis (D. Don) Spach spp., Castilla
elastica Cerv., Casuarina equisetifolia L., Cecropia L. leaf petiole, Cedrela toona Roxb.
ex Rottler and Willd., Ceiba pentandra Gaertn., C. thonningii A. Chev., Celtis brownii
Rendl., C. luzonicus Warb., C. mildbraedii Engl., C. zenkeri Engl., Chlorophora excelsa
Benth, and Hook., Chloroxylon swietenia DC., Chrysophyllum L. sp., C inchona L. sp..

C innamaum camphora (L.) J. Presl, C innamomum L. sp., C. camphora L., C istanthera K.

111

Schum. sp., Cleistopholis patens Benth., Coelocaryon preussii Warb., C offea L. sp.. C.
robusta L. Linden, C ylicodiscus gabunensis Harms, C ynometra hankie Harms.
Dacryodes pubescens (Vermoesen) H. J. Lam., Dactylocladus stenostachya Oliv.,
Dalbergia latifolia Roxb., D. sissoo Roxb., Dialium corbisieri Staner, D. pachyphjlillum
Harms, Dillenia pentagyna Roxb., Dimocarpus longan Lour., Dipterocarpus C. F.
Gaertn. spp., D. baudii Korth., Distemonanthus benthamianus Baill., Doona zeylanica
Thwaites., Dryobalanops C. F. Gaertn. sp., Drypetes leonensis Pax. Var. glabra J.
Le'onard, Elaeis guineensis J acq. (leafstalks), Elaeocarpus sericeus Stapf., “Elaeocarpus
tetrapterum” (Wood and Bright 1992), E. tuberculatus Roxb., Entandrophragma
angolense C. DC., E. cylindricum Sprague, E. utile Sprague, Erythrina L. spp.. E.
lithosperma Miq. var. inermis de. and Val., Erythrophleum guineense G. Don,
Eucalyptus deglupta Blume, Eucalyptus L’Hér. sp., E. robusta Sm., Eugenia caryophylla
St. Lag., E. jambolana Lam., Eupatorium pallescens DC., F agara macrophylla (Oliv.)
Engl., Fagus crenata Blume, Ficus L. spp., Garcinia polyantha Oliv., G. punctata Oliv.,
Gliricidia maculate H. B. and K., Gluta tourtour Marchand, G. travancorica Bedd.,
Gmelina arborea Roxb., Gossweilerondendron balsamiferum Harms, Grevillea Knight
sp., G. robusta A. Cunn., Guarea cedrata A. Chev., G. laurentii De Wild., Hannoa
klaineana Pierre, Haronga paniculata (Pert.) Lodd., Hevea brasiliensis (Willd. ex A.
Juss) Miill. Arq., Holigarna arnottiana Hook., Hopea beccariana Burck, H. ferrea Heim,
H. odorata Roxb., H. parviflora Bedd., H. wightiana Wall., “Ilteocafus baudii” [?]
(China), Julbernardia sereti (De Wild.), Khaya ivorensis A. Chev., Kayeafloribunda
Wall., Koompassia malaccensis Maingay, Lagerstroemiaflos-reginae Retz., L. speciosa

Pres., Lannea grandis Enql., Lasiococca Hook. sp., Lauraceae sp.. Lecanodiscus

112

cupanoides Planch., Leea crispa L., L. sambucina (L.) Willd., “Liana” [woody vine] (Sri
Lanka), Lithocarpus wallichiana Rehder, Lophira procera A. Chev., Lovoa klaineana
Pierre, L. trichiliodes Harms, Lufla Mill. sp., Macaranga monandra Miill.-Arg., Machilus
odoratissima Nees., Macrolobium Schreb. sp., M. macrophyllum Harms, Malus pumila
Mill., Mangifera indica L., Mansom'a altissima A. Chev., Melanorrhoea Wall. sp.,
Melochia umbellata Stapf., Microcos coriacea (Mast.) Burret, M. pinnatifida (Mast.)
Burret, Mitragyna stipulosa O. Ktze. Rev., Murraya koenigii Spreng., Musanga
cecropopdes R. Br., Myrianthus arboreus P. Beauv., Myristica L. sp., Myristica
dactyloides Wall., Napoleana imperialis P. Beauv., Nauclea (Sarcocephalus) diderichii
De Wild., “Nayabu” (Sri Lanka), Ochthocosmus africanus Hook., “Octomeles
sumatrana” (Ohno 1990), Ongokea gore Engl., Pachylobus deliciosus Pellegr.,
Palaquium gutta Burck., Pancovia laurentii De Wild., Parinari kerstingii Engl., Parishia
Hook. f. sp., Parkia bicolor A. Chev., Pithecollobium lobatum Benth., Piptadenia
africana Hook, Piptadeniastrum afi'icanum Benth., Protium pittieri Engl., Pycnanthus
angolensis (Welw.) Exell., Quercus L. sp., Q. serrata Roxb., Randia congolana De
Wild. and Th. Dur., Ricinodendron heudelotii (Baill.) Pierre, Saccharum oflicinarum L.,
Sagraea laurina Dalz., Sandoricum Cav. sp., Sapium P. Browne sp., Sapotaceae sp.,
Scorodophloeus zenkeri Harms, Shorea Roxb. ex. C. F. Gaertn. sp., S. guiso (Blanco)
Blume, S. macroptera Dyer, S. robusta Gaertn., Sindora Miq. sp., Sorindeia lameirei De
Wild., Staidtia stipitata Warb., Sterculia macrophylla Vent., S. oblonga Mater, S. villosa
Roxb., Strombosia glaucesens Engl., Strombosiopsis tetrandra Engl., Styrax bensoin
Dryand., Swietenia macrophylla King, Synsepalum subcordatum De Wild., Tarieta utilis

Sprague, Tectona grandis L., Terminalia ivorensis A. Chev., T. superba Engl., T.

113

tomentosa W. and A., Tessmannia africana Harms, T. anomala Harms, T etrapleura
(Thonn.) Taub., Thalia geniculata L., Thea sinensis L., Theobroma cacao L., Topobea
maurofernandeziana Cogn., Trichilia heudelotii Planch., T. prieureana J uss.,
Triplochiton scleroxylon K. Schum., Turraneanthus africana Benth., Vernonia arborea
Ham., V. conferta Benth., V itis L. sp., Vochysiaferruginea Mart., Xanthophyllum affine
Korth.

Specimens Examined. (104 Q; 2 6)

Type material: Lectotype Xyleborus declivigranulatus (Q; NHMW). Holotype
Xyleborus mascarenus (Q; NHMW). Lectotype Xyleborus okoumeensis: In Okumé ( Q;
NHMW). Paratype Xyleborus okoumeensis: W. Africa: Gabun-vinde (Q; NHMW).
Allotype Xyleborus okoumeensis: Germany: Carlshafen, Hess. Nassau, 17.8.28 (6 ;
NHMW). Neotype Xyleborus semiopacus: Formosa, Akau, l-10.XII.1907, Hans Sauter
leg., vend. 23.IV.1908 (9; NHMW).

Other material: AFROTROPICAL REGION: Congo: Yangambi. 1952, C. Donis, z.
1345, C011. R. Mayne, Com. Et Bois Congo, R. 2598 (1 Q; USNM). NEOTROPICAL
REGION: United States: Alabama: Baldwin Co., Mobile — Apr 88, C. Kouskelekas (2
Q; USNM); Quarentine from Florida at Costa Mesa, California, IX.5.2000, Ex.
Dimocarpus longan (1 Q; CSCA); Mississippi: Harrison Co., Gulfport, 29 March 1985.
John Davis, ex peach (healthy) (1 Q; USNM); Mississippi: Stone Co. 29 March 1985, G.
Weaver, ex plum (healthy) (1 Q; USNM); Mississippi: Stoneville, Amer. Elm, 6-23-86. J.
D. Solomon (2 Q; USNM). OCEANIA: Hawaiian Islands: Hawaii, Hilo, Hav. Fern
Wood, V-l6-‘53, Working in cut wood, C. J. Davis (1 Q; USNM); Hawaii, Hilo, X-l7-

62, ex Eucalyptus robusta, R. Nelson (1 Q; USNM); Oahu. Haleauau Val.. 7-54. B. J.

114

Ford Collector (1 Q; USNM); Oahu: Waianae Mts., 2-55, Ford (3 Q; CAS); USA: HI:
Oahu 1., N. Halawa Valley, NW of Honolulu, 390 m, 21025’N, 157051’W, 1 l-
29.VI.1991, FMHD#9l-4, Met. Polym.-Psidium-Hibiscus mixed nat./exotic forest. A.
Newton and M. Thayer, #869, window trap (6 Q; FMNH). Micronesia: Koror I., Palau
Islds., Limestone ridge N. of inlet, 16 Jan. 1948 (1 Q; USNM). ORIENTAL REGION:
Burma: Burma: Mogaung, X.4.44, Cpt. L. C. Kuitert (1 Q; USNM). China: China:
Suisapa, 1000 m., Lichuan Distr., W. Hupah, VIII.21.48 (1 Q; CAS); Salango, Kafang,
3.X.1949., F. G. Browne, ex Ilteoeafus baudii [?] (1 Q; USNM). India: India: Dehra
Dun. U. P., C. F. C. Beeson, 25.X.1915, ex Cinnamaum cambhora (1 Q; USNM); S.
India: Animalia Hills, April 1956, Cichona 3500 ft., P. S. Nathan (2 Q; USNM); South
India: Animalia Hills, Cincohona, V-60, 3500 ft., P.S. Nathan (1 Q; USNM); S. India:
Cinchona, Anamalai Hills, 3500 ft., V.1959, P. S. Nathan (1 Q; NHMW); S. India:
Coffee Re. Sta, 12-29 ’59, Coflea robusta Lot 60-13752 (1 Q; USNM); S. India: Madras.
Coimbatore, IV-1956 1400 ft., P. S. Nathan (4Q; USNM); South India: Nilgiri Hills,
Devala, 3200 ft., X-60, P. S. Nathan (2 Q; USNM). Indonesia: Java, Leg. Kalshoven (2
Q; USNM); Java, Bantam, Leg. Kalshoven, Planta nutrix Hena (10 Q; USNM). Japan:
Japan: Matskawa, 27-VII-1980, S. L. Wood (1 Q; USNM); Okinawa: ID, Nov 16, 1945.
F. N. Young (2 Q; USNM); Japan: Tamagowa, 29-VII-1980, S. L. Wood Fagus crenatus
(7 Q; USNM); Japan: Tokyo, Takao, VlIl.31.l957, Coll. Akira Nobuchi (2 Q; USNM).
Philipine Islands: Philippines, Mindanao, 1965, Krauss, Cacao trunk (1 Q, l 6;
USNM). Taiwan: Taiwan: Raisya, 24.XI.1934, S. Issiki (1 Q; NHMW). Thailand:
Thailand: Chiengmai Prov., E. Fk. Mae Ping, 56 km N. Chiengmai, 1300’, 24-Xi-1964.

W. L. and J. G. Peters, at light (1 Q; CAS). Sri Lanka: Sri Lanka: Col Dist., Labugama,

115

23 June 1975, S. L. Wood, Collected from branches (10 Q; USNM); Sri Lanka: Col Dist..
Labugama, 23 June 1975, S. L. Wood, Collected from twigs (2 Q; USNM); Sri Lanka:
Col. Dist., Labugama, 23 June 1975, S. L. Wood, Collected from Liana (1 Q; USNM):
Sri Lanka: Col Dist., Labugama 23 June 1975, S. L. Wood, Misc. hosts (1 Q; USNM):
Sri Lanka: Gal. Dist., Ugugama, Kanneliya Jungle, 400 ft., 6-12-X-1973, at black light.
K. V. Krombein, P. E. Karunarante, P. Fernando, J. Fernando (1 Q; USNM); Sri Lanka:
Kal. Dist., Morapitiya, 250 mtrs., 27 May 1975, S. L. Wood, Misc. hosts (3 Q; USNM);
Sri Lanka: Ceylon: Kan. Dist., Kandy, 1-15 March 1971, Piyadasa and Sompala (1 Q;
USNM); Sri Lanka: Keg. Dis., Kitulhala, 200 m, 30 May 1975, S. L. Wood, Host:
Nayabu (1 Q; USNM); Sri Lanka: Keg. Dist., Kitulgala, 250 m., 27 May 1975, S. L.
Wood, Collected from pole (1 Q; USNM); Sri Lanka: Keg. Dist., Kitulgala, 200 m., 30
May 1975, S. L. Wood, Host: Doona sp. (2 Q; USNM); Sri Lanka: Mat. Dist.,
Enselwatte, 800 mtrs. 25 May 1975, S. L. Wood, Misc. hosts (7 Q: USNM); Sri Lanka:
Rat, Dist., Gilimale, 17 May 1975, S. L. Wood, Host: Myristica dactyloides (3 Q;
USNM); Sri Lanka: Rat. Dist. Gilimale, 17 May 1975, S. L. Wood, Collected from pole
(1 Q; USNM). PALEARCTIC REGION: Germany: Germany: Carlshafen, Hess.

Nassau, 17.8.28 (3 Q; NHMW).

X ylosandrus curtulus (Eichhoff)
(Fig. 2.17)
Xyleborus curtulus Eichhoff, 1869: 281. Holotype Q: Brazil: Patria; IRSNB.

Xylosandrus curtulus (Eichhoff): Wood, 1982: 770.

116

Xyleborus curtuloides Eggers, 1941a: 102. Holotype Q: Guadeloupe (Gourbeyre); Eggers
Collection (not listed by Anderson and Anderson, 1971 or Schedl, 1979a). Synonymy:
Wood, 1982: 770.

Xyleborus biseriatus Schedl, 1963: 226. Holotype Q: Nova Teutonia, Santa C atarina.
Brazil; NHMW. Synonymy: Wood, 1973: 187.

Xylosandrus strumosus Schedl, 1972: 73. Holotype Q: Brasilien, Corcovado. Guanabara;
NHMW. Synonymy: Wood and Bright, 1992: 793.

Anisandrus zimmermanni Hopkins 1915: 67. Holotype Q: Biscayne, Florida; USNM.
Synonymy: Wood, 2007: 467.

Xylosandrus zimmermanni (Hopkins): Wood, 1962: 79.

Notes. Wood and Bright (1992) first list Xylosandrus strumosus as a synonym of
Xylosandrus curtulus. The synonymy is referenced as “Wood 1992: (in press)”. but the
synonymy did not appear in another publication.

Diagnosis. Female 1.3 — 1.5 mm long; 2.1 times longer than wide. Body brown to dark
brown; antenne and legs yellowish brown. Foms punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.8 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et a1. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presence of a pronotal-
mesonotal mycangium. Pronotal disc morderately punctate. Lateral aspect of pronotum
rounded (type 1, Hulcr et al. 2007). Pronotum with lateral costa and carina. Procoxae

widely separated. Protibiae with 4 socketed teeth on lateral margin; meso- and metatibiae

117

with 7 socketed teeth. Elytra 1.3 times longer than wide; 1.5 times longer than pronotum.
Discal striae punctate; interstriae uniseriate punctate. Elytral disc gradually curving into
declivity. Declivity convex, lateral margins carinate to 7‘h interstriae. Six striae visible
on declivity. Striae punctate, with semi-appressed, hair-like setae, shorter than the width
of second declivital interstriae. Interstriae granulate, uniseriate, with erect, hair-like
setae, longer than the width of second declivital interstriae.

This species is morphologically similar to X compactus (Fig. 2.14) and X
pusillus (Fig. 2.36). Xylosandrus curtulus can be distinguished from these species by the
following characters: pronotal disc glabrous, except for a dense patch of short, erect, hair-
like setae basally, indicating the presence of a pronotal-mesonotal mycangium; and elytra
strongly arched from base to middle of declivity.

Distribution. NEARCTIC REGION (Introduced): United States (Florida).
NEOTROPICAL REGION: Antilles Islands (Guadeloupe), Brazil. Colombia, Costa
Rica, Guatemala, Honduras, Mexico (Chiapas, Colima, Hidalgo, Nayarit, Oaxaca. San
Luis Potosi, Varacruz), Venezuela.

Hosts. Acer rubrum L., “Anonillo” (Guatemala), Ardisia Gaertn. sp., .Byrsonima
cotinifolia H. B. and K., Calliandra confusa Sparque and Riley, C edrela odorata L.,
Chrysobalanus L. sp., Coffea canephora var. robusta (Linden) A. Chev., C upania
guatemalensis Radlk., Ficus L. spp., Inga Scop. sp., Nectandra Roll. ex Rottb. sp.,
Phoradendron robustissimum Eichler, Phoradendron Nutt. spp., Rheedia edulis Planch.
and Triana., Serjania Mill. sp., Spondias mombin L., T abebuia Gomes ex. DC. sp.,
Ocotea catesbyana Sarq., Dodonaea viscosa J acq..

Specimens Examined. (86 Q; 3 6)

118

Type material: Holotype X yleborus biseriatus; (Q; NHMW). Holotype X ylosandrus
strumosus (Q; NHMW). Holotype Anisandrus zimmermanni (Q; USNM).

Other material: NEARCTIC REGION: United States: Florida: Sebring. 6-20-1951.
Price Beamers — Wood (1 Q; USNM); Florida: Sebring, 6-20-1951, Price Beamers —
Wood, Ardisia (7 Q, l 6: USNM); Florida: Sebring, 6-20-1951, Price Beamers — Wood.
Ocotea catsibiana (l QUSNM); Florida: Sebring, 6-20-1951, Price Beamers — Wood.
Red Maple (2Q, 1 6; USNM). NEOTROPICAL REGION: Colombia: Colombia:
Caicedonia near Sevilla, Aug. 1959, Coffee branches, Leg. Duque (1 Q; USNM):
Colombia: El Bosque, Caicedonia (V.), Junio 1959, J. Restrepo, en Café (11 Q, 1 6 ;
USNM); Colombia: 1959, leg. Beteem (1 Q; USNM). Costa Rica: Costa Rica: Guapiles.
Lim. 300 ft., VIII-22-’66, S. L. Wood, unknown vine (2 Q; USNM); Costa Rica: Santa
Ana, S. J., 4000 ft., VII-l-l963, S. L. Wood, Coffee (1~ Q; USNM); Costa Rica: Santa
Ana, S. J ., 4000 ft., VIII-30-1963, S. L. Wood, Rheedia edulis (1 Q; USNM); Costa Rica:
Santa Ana, S. J ., 4000 ft., VIII-30-l963, S. L. Wood, unknown sapling (3 Q; USNM);
Costa Rica: Santa Ana, S. J., 400 ft., VIII-l-1963, S. L. Wood, Cupania guatemalensis (9
Q; USNM); Costa Rica: San Ignacio, S. J., 4700 ft., VIII-5-1963. S. L. Wood. unknown
sapling (1 Q; USNM); Costa Rica: Pandora, Lim., 150 ft., VIII-23-1963, S. L. Wood.
unknown shrub (4 Q; USNM); Costa Rica: Tapnti Cart., 400 ft., VIII-17-1963, S. L.
Wood, Calliandra confusa (1 Q; USNM). Guatemala: Guatemala: Palin, Esquintla, 100
ft., V-19-1964, S. L. Wood, Anonillo (1 Q; USNM); Guatemala: Rodeo Esquintla. 500
ft., Vl-4-1964, S. L. Wood, Unknown vine (1 Q: USNM); Guatemala: Volcan de Agua,
3000 ft., V-19-1964, S. L. Wood, unknown twigs (1 Q; USNM); Guatemala: Volcan de

Agua, 3000 ft., V-19-1964. S. L. Wood, unknown broken branch (1 Q: USNM).

119

Honduras: Honduras: Zamorano, Moraz, 2200 ft., lV-18-1964, S. L. Wood,
Phoradendron robustissimum (1 QUSNM); Honduras: Zamorano, Moraz, 2200 ft.,
Serjania (1 Q; USNM). Mexico: Mexico: 2 mi W Armeria, Col., VI-28-l965, 200 ft., S.
L. Wood, Phoradendron (2 Q; USNM); Mexico: El Salto, S. L. P., VI-19-53, taken on
Ficus (1 Q; USNM); Mexico: Morelos, Ruinas de Xochicalco, S-322, 22E 82, 1200 ms 11
m, Col Atkinson and Equihua, Hosp. Dodonaea viscosa (1 Q; USNM); Mexico: Laguna
Sta. Maria N., VII-6-1965, 3000 ft., S. L. Wood, unknown vine (1 Q; USNM); Mexico:
Romero 12 mi S, VI-24-1967, OAX, S. L. Wood, Phoradendron (1 Q; USNM); Mexico:
5 mi N Rosamorada N., VII-15-1965, 300 ft., S. L. Wood, unknown broken branch (2 Q;
USNM); Mexico: Romeo, 23 mi N VC, 300 ft., S. L. Wood, unknown branch (1 Q;
USNM); Mexico: Tapachula Chis, 21 VIII 82, C01. A. B. Celis, Coffea canephora var.
robusta (1 Q; USNM); Mexico: Vera Cruz, 16 mi S Tecolutla, VI-26.53 (8Q; USNM).
Venezuela: Venezuela: S. of Barrancas, Barinas, 150 m., XI-5-69, S. L. Wood, Spondias
mombin (1 Q; USNM); Venezuela: 9 km S. of Barrancas, Barinas, 150 m., XI-15-69, S.
L. Wood, Inga (1 Q; USNM); Venezuela 40 km, E. Canton-Barinas, 111-8-1970, 70 m., S.
L. Wood, unknown vine (1 Q; USNM); Venezuela: 20 km SW El Vigia, Merida, XII-10
69, e1. 50 m., S. L. Wood (1 Q; USNM); Venezuela: Rancho Grande, Aragua, 1100 m.,.
lV-9—l970, S. L. Wood, T abebuia (3 Q; USNM); Venezuela: Rancho Grande, Aragua.
1100 m., IV-9-1970, S. L. Wood, Nectandra sp. (5 Q; USNM); Venezuela: 40 km SE

Socopo, Barinas, 1-25-1970, 150 m., S. L. Wood, Bejuco Blanco (1 Q; USNM).

X ylosandrus derupteterminatus (Schedl)

(Fig. 2.18)

120

Xyleborus derupteterminatus Schedl, 1951: 64. Holotype Q: Java, Mount Gede
IX.1932L. G. E. Kalshoven Coll., NHMW.
Xylosandrus derupteterminatus (Schedl): Schedl, 1964: 213.
Diagnosis. Female 2.0 — 2.3 mm long; 2.0 times longer than wide. Body dark brown to
black; antennae and legs same color as body. F rons punctate, with distinct median keel
between eyes. Antennae with 5 funicular segments. Antennal club obliquely truncate;
first segment forming a circular costa; segment one covering entire posterior face.
Pronotum 0.8 times longer than wide. Dorsal aspect of pronotum rounded (type 1, Hulcr
et al. 2007). Protonal vestiture of semi-appressed, hair-like setae; pronotal disc glabrous,
except for mycangial setae. Basal pronotum with dense patch of short, erect setae
indicating the presense of a pomotal-mesonotal mycangium. Pronotal disc moderately
punctate basally. Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotum
with lateral costa and carina. Procoxae widely separated. Protibiae with 4 socketed teeth
on lateral margin; mesotibiae with 11 socketed teeth; metatibiae with 13 socketed teeth.
Elytra 1.2 times longer than wide; 1.4 times longer than pronotum. Discal striae
punctate; interstriae uniseriate punctate. Declivital face of elytra steep, abruptly
separated from disc. Declivity flattened, lateral margin carinate to 7th interstriae. Five
striae visible on declivity. Striae punctate, without setae. Interstriae granulate.
uniseriate, with erect, hair-like setae. shorter than the width of second declivital
interstriae.

This species is morphologically similar to X morigerus (Fig. 2.35) and X.
terminatus (Fig. 2.41). X ylosandrus derupteterminatus can be distinguished from X

morigerus by the following characters: face of elytral declivity flattened; and 5 striae

121

visible on declivity. Xylosandris derupteterminatus can be distinguished from X
terminatus by the following characters: larger species, 2.0 — 2.3 mm long; declivital striae
with erect, hair-like setae; and declivital interstriae multiseriate granulate.

Distribution. ORIENTAL REGION: Indonesia (Java, Moluccas. Sulawesi).

Hosts. Unknown.

Specimens Examined. (10 Q; 0 6)

Type material: Holotype X yleborus derupteterminatus (Q ;NHMW).

Other material: ORIENTAL REGION: Indonesia: INDONESIA: SULAWESI
UTARA, Dumoga-Bone N.P., Plot A, ca 200 m Lowland forest, May—85, Flight intercept
trap (4 Q; RAB); INDONESIA: SULAWESI UTARA, Dumoga—Bone N.P., Plot A. ca
200 m Lowland forest, Dec. 1985, suspended carrion (1 Q; RAB); INDONESIA:
SULAWESI UTARA, Dumoga-Bone N.P., Plot C, ca 400 m lowland forest, Apr-85.
Flight intercept trap (2 Q; RAB); INDONESIA: SULAWESI UTARA, Dumoga-Bone
N.P., Plot C, ca 400 m lowland forest, Feb-85, Flight intercept trap (1 Q; RAB);
INDONESIA: SULAWESI UTARA, Dumoga—Bone N.P., G.Mogogonipa summit, 1008

m., 22-24.x.85, yellow plate (1 Q; RAB).

X ylosandrus deruptulus (Schedl)
(Fig. 2.19)
Xyleborus deruptulus Schedl, 1942b: 37. Lectotype Q: Java, Mount Dede, 800 m, VIII-
1923, Nr 54, Kalshoven; NHMW; designated by Schedl, 1979a: 78.

Xylosandrus deruptulus (Schedl): Schedl, 1964: 213.

122

Diagnosis. Female 1.8 mm long; 2.2 times longer than wide. Body brown; antennae and
legs light brown. F rons rugose. Antennae with 5 funicular segments. Antennal club
obliquely truncate; first segment forming a circular costa; segment one covering entire
posterior face. Pronotum 0.9 times longer than wide. Dorsal aspect of pronotum rounded
(type 1, Hulcr et al. 2007). Pronotal vestiture of appressed, hair-like setae; setae less
dense on disc. Specimen examined too damaged to determine whether pronotal
mycangial setae are present. Pronotal disc moderately punctate. Lateral aspect of
pronotmn basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa and carina.
Procoxae widely separated. Pro-, meso- and metatibial teeth not visible on specimen
examined. Elytra 1.4 times longer than wide; 1.6 times longer than pronotum. Discal
striae punctate; interstriae multiseriate punctate. Elytral disc gradually curving into
declivity. Declivity convex, lateral margins carinate to 7th interstriae. Six striae visible
on declivity. Striae punctate. Interstriae granulate, uniseriate. Specimen examined too
damaged to determine states of declivital setae and setae not mentioned in original
description (Schedl 1942b).

This species is morphologically similar to X assequens (Fig. 2.10). X ylosandrus
deruptulus can be distinguished from X assequens by the following characters: more
elongate, 2.2 times longer than wide; elytral disc with interstriae more sparsely
punctured, multiseriate only on first interstriae; declivital interstriae uniseraite granulate;
and declivital surface shinning, not matte.

Distribution. ORIENTAL REGION: Indonesia (Java).
Hosts. Unknown.

Specimens Examined. (1 Q; 0 6)

123

Type material: Lectotype X yleborus deruptulus (Q; NHMW).

X ylosandrus discolor (Blandford)
(Fig. 2.20)

Xyleborus discolor Blandford, 1898: 429. Holotype Q: Ceylon, E. E. Green; BMNH.
Xylosandrus discolor (Blandford): Browne, 1963: 55.
Xyleborus posticestriatus Eggers, 1939b: 119. Lectotype Q: Formosa (Taihoku); USNM;
designated by Anderson and Anderson, 1971: 26. Synonymy: Schedl, 195 8: 149.
Xylosandrus posticestriatus (Eggers): Nunberg, 1959: 434.
Diagnosis. Female 1.8 — 2.0 mm long; 1.8 times longer than wide. Body bicolored;
pronotum distinctly lighter than elytra; pronotum light brown and elytra dark brown;
antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.8 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
erect, hair-like setae; setae less dense on disc. Basal pronotum with a dense patch of
short, erect setae, indicating the presense of a pronotal-mesonotal mycangium. Pronotal
disc densely asperate-punctate, with sculpture separated by distance less than or equal to
their size. Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotum with
lateral costa and carina. Procoxae widely separated. Protibiae with 4 — 5 socketed teeth
on lateral margin; meso- and metatibiae with 8 — 9 socketed teeth. Elytra of equal length
and width; 1.2 times longer than pronotum. Discal striae punctate; interstriae multiseriate

punctate. Declivital face of elytra steep and abruptly separated from disc. Declivity

124

flattened, lateral margins carinate to 7th interstriae. Four striae visible on declivity. Striae
coarsely granulate, with appressed hair-like setae, shorter than the width of second
declivital interstriae. Interstriae granulate, multiseriate, with appressed. hair-like setae.
shorter than the width of second declivital interstriae.

This species is one of four bicolored Xylosandrus with the pronotum distinctly
lighter than the elytra: X arquatus (Fig. 2.9), X discolor, X ferinus (Fig. 2.23), and X
mesuae (Fig. 2.30). Xylosandrus discolor can be distinguished from these species by the
following characters: elytral declivity steep and abruptly separated from disc. and
declivital striae and interstriae with a dense vestiture of appressed setae.
Distribution. AUSTRALIAN REGION: Queensland. OCEANIA: Micronesia.
ORIENTAL REGION: Burma, China (F ujian, Guangdong, Sichuan, Yunnan), India
(Andaman Islands, Assam, Tamil Nadu, Uttar Pardesh), Indonesia (Java), Malaysia. Sri
Lanka, Taiwan, Thailand.
Hosts. Ailanthus altissima (Mill.) Swingle, Albizia Benth. sp., “Avocado” (Java).
Bauhinia variegata L., Camellia sinensis Kuntze, Cassia multijuga Rich., Castanopsis
fargesii Franch., Cedrela toona Roxb. ex Rottler and Willd., Chloroxylon swietenia DC..
Coffea L. spp., Coflea Arabica L., C. robusta L. Linden, Cinnamomum camphora (L.) J.
Presl., Eupatorium L. sp., Grevillea robusta A. Cunn., Hevea brasiliensis (Willd. ex A.
J uss) Miill. Arq., “Igall sp.” (Thailand), Juglans nigra L., “Liana” [woody vine] (Sri
Lanka) sp., Machilus indica Kurz., “Mango” (Thailand), Pterospermum acerifolium
Willd., “Rukaththana” (Sri Lanka), Rhus chinensis Mill., Sophorajaponica L., Swietenia
mahagoni (L.) Jacq., Tephrosia candida DC., Terminalia myriocarpa Van Heurck and

Miill. Arq., T. procera Roxb., Theobroma cacao L., Vitis vinifera L.

125

Specimens Examined. (61 Q; 4 6)

Type material: Lectotype Xyleborus posticestriatus: Formosa: Taihoku, 10.VII.1934, C 01.
M. Chujo (Q; NHMW). Cotype Xyleborus posticestriatus: XI.1926 (locality unreadable)
(6; NHMW).

Other material: AUSTRALIAN REGION: Queensland: AUSTRALIA: N. QLD, Iron
Range, 26-31.X.1991, Wood, Dunn, and Hasenpusch (3 Q; RAB); NE.Q: 16°54'S x
145°42'E, Whitefield Range, 550 m, 28 Aug - 19 Oct 1991, Monteith and Janetzki, Pitfall
and Intercept traps (1 Q; RAB). ORIENTAL REGION: Indonesia: Java, 565 m.,
Bandjar, VII.32, L. G. E. Kalshoven (1 Q; NHMW); Java, Boger, VIII-1964, N. L. H.
Krauss, avocado branch (4 Q; USNM); W. Java: 800 m Mount Gedeh, 7.1933, Leg.
Kalshoven, Planta nutrix, Eupartorium (1 Q; USNM). Malaysia: Koror, Palau Is. Apr
1953, J. W. Beardsley (1 Q; USNM). Sri Lanka: Ceylon [Sri Lanka], Hantane,
XII.1962, Dr. D. Calnaido (1 Q; NHMW); Ceylon [Sri Lanka]: Perdadeiya, 11.VII.1914.
A. Rutherford (2 Q; USNM); Ceylon [Sri Lanka], Sabargamuva, 12.Xl.1950, E. Judenko
(1 Q; NHMW); Sri Lanka: Col. Dist., Labugama, 23 June 1975, S. L. Wood, Misc. hosts
(4 Q; USNM); Sri Lanka: Col. Dist., Labugama, 23 June 1975, S. L. Wood, collected
from twigs (1 Q; USNM); Sri Lanka: Mate. Dist., 48 km N Naula, 200 mtrs., 14 June
1975, S. L. Wood, Host: Rukaththana (1 Q; USNM); Sri Lanka: Matte. Dist., 48 km, N
Naula, 200 mtrs., 14 June 1975, S. L. Wood, collected from log (1 Q; USNM); Sri Lanka:
Mon. Dist., 8 km NW Bibile, 50 mtrs, 7 June 1975, S. L. Wood, collected from tree
seedling (13 Q; USNM); Sri Lanka: Mon. Dist., 8 km NW Bibile, 50 mtrs, 7 June 1975.
S. L. Wood, collected from pole (2 Q; USNM); Sri Lanka: Mon Dist. 8 km NW Bibile.

50 mtrs, 7 June 1975, S. L. Wood, colleted from Liana (2 Q; USNM); Sri Lanka: Rat.

126

Dist., Ratnapura, Resthouse, 200-300 ft., 24-X-1976, black light, Collected by: G. F.
Hevel, R. E. Dietz, W. S. Karunaratne, D. W. Balasooriya (1 Q; USNM). Thailand:
THAILAND: Chiang Mai, 2.vii.72, R. A. Beaver, ex Bauhinia variegata (1 Q; RAB);
Thailand: Chiang Mai: Doi Inthanon, 5.viii.02, R. A. Beaver and K. Koivisto (1 Q;
RAB); Thailand: Chiang Mai: Maerim, 13-14.iv.00, R. A. Beaver (1 Q; RAB); Thailand:
Chiang Mai: Maerim, 12.x.93, R. A. Beaver, At light (1 Q; RAB); Thailand: Chiang Mai:
Maerim, 1-3.I.2000, R. A. Beaver, M.T. (1 Q; RAB); Thailand: Chiang Mai: Maerim.
5.v.94, R. A. Beaver, At light (1 Q; RAB); Thailand: Chiang Mai: Maerim, 29-31.x.95,
R. A. Beaver, M.T. (I Q; RAB); Thailand: Chiang Mai: Maerim, 28.xi.02, R. A. Beaver.
FIT (1 Q; RAB); Thailand: Chiang Mai: Maerim, 9.x.96, R. A. Beaver, ex mango twig (2
Q; RAB); Thailand: Chiang Mai: Maerim, 5.x.96, R. A. Beaver, ex mango twig (1 Q, 1
6; RAB); Thailand: Chiang Mai: Maerim, 21 .ix.96, R. A. Beaver, ex mango twig (1 Q;
RAB); Thailand: Chiang Mai: Maerim, 3.xii.96, R. A. Beaver, ex mango twig (1 Q:
RAB); Thailand: Chiang Mai: Maerim, 16.x.96, R. A. Beaver, ex mango twig (1 6;
RAB); Thailand: Chiang Mai: Maerim, 26.viii.99, R. A. Beaver, ex Igall sp. (1 Q, l 6 ;
RAB); Thailand: Chiang Mai: Maerim, 23.vii.03, R. A. Beaver, FIT (1 Q; RAB);
Thailand: Chiang Mai: Maerim, 10.xii.02, R. A. Beaver, FIT (1 Q; RAB); Thailand:
Chiang Mai: Maerim, 18.xi.03, R. A. Beaver, FIT (1 Q; RAB); THAILAND: Chiang Mai
300 m, Maerim, 16.1.92, R. A. Beaver (1 Q; RAB); THAILAND: Chiang Mai 300 m,

Maerim, 15.I.92, R. A. Beaver (1 Q; RAB).

X ylosandrus diversepilosus (Eggers)

(Fig. 2.21)

127

Xyleborus diversepilosus Eggers, 1941b: 224. Holotype Q: China, Prov. Fukien (Kuatun.
2300 m), 1.5.1938, L. J. Klapperich; ZMFK.
Xylosandrus diversepilosus (Eggers): Browne, 1963: 55.
Diagnosis. Female 1.8 — 2.3 mm long; 2.1 times longer than wide. Body light brown:
antennae and legs yellowish brown. F rons not visible on specimen examined. Antennae
with 5 funicular segments. Antennal club obliquely truncate; first segment forming a
circular costa; segment one covering entire posterior face. Pronotum 0.9 times longer
than wide. Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal
vestiture of semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum
with a dense patch of short, erect setae, indicating the presence of a pronotal-mesonotal
mycangium. Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotal disc
densely asperate-granulate, with sculpture separated by distance less than or equal to their
size. Pronotum with lateral costa, carinate. Procoxae not visible on specimen examined.
Pro-, meso- and metatibial teeth not visible on specimen examined. Elytra 1.2 times
longer than wide; 1.3 times longer than pronotum. Discal striae punctate; interstriae
multiseriate punctate. Declivital face steep and abruptly separated from disc. Declivity
flattened, lateral margins carinate to 7th interstriae. Four striae visible on declivity. Striae
coarsely granulate, without setae. Interstriae granulate, biseriate, with erect, hair-like
setae, longer than twice the width of second declivital interstriae.

This species is morphologically similar to X borealis (Fig. 2.11) and X brevis
(Fig. 2.13). Xylosandrus. diversepilosus can be distinguished from these species by the

following characters: declivital striae coarsely granulate, without setae; and interstriae

128

granulate, with erect, hair-like setae, longer than twice the width of second declivital
interstriae.

Distribution. ORIENTAL REGION: China (F ukian).

Hosts. Unknown.

Specimens Examined. (1 Q; 0 6)

Type material: Holotype X yleborus diversepilosus (Q; ZMF K).

X ylosandrus eupatorii (Eggers)
(Fig. 2.22)

Xyleborus eupatorii Eggers, 1940: 140. Holotype Q: Java (Tjibodas, G. Gedeh);
Kalshoven Collection.
Xylosandrus eupatorii (Eggers): Schedl, 1964: 213.
Diagnosis. Female 1.8 — 2.1 mm long; 2.2 times longer than wide. Body brown to dark
brown; antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum of equal length and width. Dorsal
aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-
appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presence of a pronotal-
mesonotal mycangium. Pronotal disc moderately punctate. Lateral aspect of pronotum
rounded (type 1, Hulcr et al. 2007). Pronotum with lateral costa and carina. Procoxae
widely separated. Protibiae with 4 socketed teeth on lateral margin; meso- and metatibial

teeth not visible on specimen examined. Elytra 1.2 times longer than wide; 1.2 times

129

 

 

longer than pronotum. Discal striae punctate; interstriae uniseriate punctate. Elytral disc
gradually curving into declivity. Declivity convex, lateral margins carinate to 7th
interstriae. Six striae visible on declivity. Striae punctate, without setae. Interstriae
granulate, uniseriate, with erect, hair-like setae, longer than twice the width of second
declivital intrstriae.

This species is morphologically similar to X adherescens (Fig. 2.7) and X
germanus (Fig. 2.24). Xylosandrus eupatorii can be distinguished from these species by
the following characters: pronotum of equal length and width; pronotal disc glabrous,
except for mycangial setae; and declivital interstriae uniseriate granulate, with erect, hair-
like setae, longer than twice the width of second declivital interstriae.

Distribution. ORIENTAL REGION: Indonesia (Java).

Hosts. “Eupatorium tjibeureum” (Java).

Specimens Examined. (1 Q; 0 6)

Type material: Cotype Xyleborus eupatorii: Java: Gedeh, 1700 M., V1.32, leg. H. R. A.

Muller, Eupatorium tjibeureum (Q; NHMW).

X ylosandrus ferinus (Schedl)
(Fig. 2.23)
Xyleborusferinus Schedl, 1936: 31. Lectotype Q: India: Travancore Mt. Estate, Tamil
Nadu, VII.1934, S. A. Ran, on red gum; NHMW; designated by Schedl, 1979a: 96.
Xylosandrusferinus (Schedl): Browne, 1963: 55.
Notes. Schedl (1936) failed to designate a holotype in his original description of X

ferinus and subsequently designated a lectotype (Schedl 1979a).

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Diagnosis. Female 1.6 — 1.8 mm long; 2.2 times longer than wide. Body bicolored,
pronotum distinctly lighter than elytra; pronotum light brown and elytra brown; antennae
and legs yellowish brown. F rons punctate. Antennae with 5 funicular segments.
Antennal club obliquely truncate; first segment forming a circular costa; segment one
covering entire posterior face. Pronotum 0.9 times longer than wide. Dorsal aspect of
pronotum rounded (type 1, Hulcr et a1. 2007). Pronotal vestiture of semi-appressed, hair-
like setae; pronotal disc glabrous, except for mycangial setae. Basal pronotum with a
dense patch of short, erect setae, indicating the presence of a pronotal-mesonotal
mycangium. Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type
0, Hulcr et al. 2007). Pronotum with lateral costa, not carinate. Procoxae widely
separated. Protibiae with 5 socketed teeth on lateral margin; mesotibiae with 8 socketed
teeth; metatibial teeth not visible on specimens examined. Elytra 1.4 times longer than
wide; 1.6 times longer than pronotum. Discal striae punctate; insterstriae uniseriate
punctate. Elytral disc gradually curving into declivity. Declivity convex, lateral margins
carinate to 7th interstriae. Five striae visible on declivity. Striae punctate, without setae.
Interstriae granulate, uniseriate, with erect, hair-like setae, shorter than the width of
second declivital interstriae.

This species is one of four bicolored Xylosandrus with the pronotum distinctly
lighter than the elytra: X. arquatus (Fig. 2.9), X discolor (Fig. 2.20), X ferinus, and X
mesuae (Fig. 2.30). Xylosandrusferinus can be distinguished from X discolor by the
following characters: elytral disc gradually curving into declivity. X ylosandrus arquatus
can be distinguished from X mesuae by its larger size of 1.6 — 1.8 mm long. Xylosandrus

ferinus can be distinguished from X arquatus by the following characters: five striae

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visible on the elytral declivity; declivital striae without setae; and lighter color, with
pronotum light brown and elytra brown.

Distribution. ORIENTAL REGION: India (Tamil Nadu).

Hosts. Red gum.

Specimens Examined. (2 Q; 0 6)

Type material: Lectotype X yleborus ferinus (Q; NHMW). Syntype Xyleborusferinus:
India: Travancore Mt. Estate, Tamil Nadu, VII.1934, S. A. Ran, on red gum (Q:

NHMW).

X ylosandrus germanus (Blandford)
(Fig. 2.24)

Xyleborus germanus Blandford, 1894c: 106. Syntypes (sex?); Oyayama, Nikko,
Subashiri, Kiga, Miyanashita: BMNH.
X ylosandrus germanus (Blanford): Hoffrnann, 1941: 38.
Xyleborus orbatus Blandford, 1894c: 123. Holotype Q: Kurigahara, Japan; BMNH.
Synonymy: Choo, 1983: 100.
Notes on types: Browne (1963) wrongly lists X germanus as the type species for
X ylosandrus.
Diagnosis. Female 1.9 — 2.5 mm long; 2.3 times longer than wide. Body brown to dark
brown; antennae and legs yellowish brown. F rons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 1.1 times longer than wide.

Dorsal aspect of pronotum rounded (typel , Hulcr et al. 2007). Pronotal vestiture of semi-

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appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense patch of
short, erect setae, indicating the presence of a pronotal-mesonotal mycangium. Pronotal
disc moderately punctate. Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007).
Pronotum with lateral costa and carina. Procoxae widely separated. Protibiae with 5
socketed teeth on lateral margin; mesotibiae with 11 — 12 socketed teeth; metatibiae with
12 — 13 socketed teeth. Elytra 1.1 times longer than wide; 1.1 times longer than
pronotum. Discal striae punctate; interstriae uniseriate punctate. Elytral disc gradually
curving into declivity. Declivity convex, lateral margins carinate to 7'h interstriae. Six
striae visible on declivity. Striae punctate, without setae. Interstriae granulate,
uniseriate, with erect, hair-like setae, longer than the width of second declivital
interstriae.

This species is morphologically similar to X adherescens (Fig. 2.7) and X
eupatorii (Fig. 2.22). Xylosandrus germanus can be distinguished from these species by
the following characters: pronotum longer than wide; and pronotal disc evenly pubescent.
not glabrous.

Distribution. NEARCTIC REGION (Introduced): Canada (British Columbia.
Ontario, Quebec), United States (Connecticut, Delaware, Florida, Illinois, Indiana.
Kentucky, Maine, Massachusets, New Jersey, New York, North Carolina, Ohio, Oregon.
Pennsylvania, Rhode Island, Virginia, West Virginia). OCEANIA: Hawaiian Islands.
ORIENTAL REGION: China (Anhui, Fujian, Shanxi, Xizang [Tibet], Yunnan), Japan,
Ryukyu Islands, Taiwan, Vietnam. PALEARCTIC REGION (Introduced): Austria.

Belgium, France, Germany, Italy, Korea. Switzerland. Yugoslavia.

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Hosts. Abies fabric (Masters) Craib, A. pectinata Poir., Acer L. spp., A. platanoides L.,
A. pseudoplatanus Falk., Alnus glutinosa (L.) Gaertn., Alnus Mill. sp. . Betula verrucosa
Ehrh., Carpinus betulus L., C. laxiflora (Siebold and Zucc.) Blume, C arya Nutt. sp..
Cassia siamea Lam., Castanea crenata Siebold and Zucc., Castanopsis (D. Don) Spach
sp., Celtis tenuifolia Nutt., Chamaecyparis obtuse Siebold and Zucc., C leyera japonica
Siebold and Zucc., C ornus florida L., Diospyros kaki Thunb., F agus crenata Blume.

F agus multinervis Nakai, F. sylvatica L., F raxinus L. spp., Juglans regia L., Juglans L.
sp., Myrica carolinensis Mill. , Myrica L. sp., Lindera erythrocarpa Makino,
Liriodendron tulipifera L., Machilus Nees sp., Morus L. spp., Nyssa aquatica L., Picea
abies (L.) K. Harst., P. excelsa Link, Pinus densiflora Siebold and Zucc., P. pentaphylla
Carriére, P. strobes L., Pinus L. sp., Prunus avium (L.) L., P. cerasus L., Prunus
scrotina Ehrh., Prunus L. spp., Pyrus scrotina Rehder, Pyrus L. sp., Rhus chinensis Mill.,
Quercus rubra L., Q. sessiliflora Salisb., Quercus L. spp., Robinia pseudoacacia L..
Schima superba Gardn. and Champ, Styrax obassia Siebold and Zucc., S. japonicus
Siebold and Zucc., Taxodium distichum Rich., Ulmus effuse Willd., Ulmus L. spp., Vitis
L. sp., “young chestnut” (Japan), Ziziphusjujuba Mill..

Specimens Examined. (66 Q; 6 6)

Other material: NEARCTIC REGION: United States: Indiana: Turkey Run St. Park,
VI-10-1967, L. and C. W. O’Brien (1 Q; USNM); Indiana: W. Lafayette, McCormick
Woods, V.26—l968, Collectors L. and C. W. O’Brien (1 Q;USNM); IND., W. Lafayette.
Tippercanoe Co., 12 Apr. 1981, M. and N. Deyrup, in pan trap below Malaise trap (2 Q;
RAB); Maryland: Takoma Park, Montgomery Co., 17-IV-1969, W. H. Tyson (1 Q;

USNM); North Carolina: McDowell Co., 10.20.1976, B. C. Weber collector (1 6;

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USNM); North Carolina: McDowell Co., 7-21 1977, B. C. Weber collector (1 Q, 1 6 :
USNM); North Carolina: McDowell Co., 10-19 1976, B. C. Weber collector (2 Q;
USNM); North Carolina: McDowell Co., 10.20.1976, B. C. Weber collector (9 Q;
USNM); North Carolina: Raleigh, 25.6.63, Prunus scrotina (1 Q; NHMW); New Jersey:
Norwood, Apl 11 1944, Schott, from branches of Myrica carol (6 Q; USNM); New
Jersey: Princeton, 2 June 1971, R. J. Gouger, Fagus sylvatica (4 Q; USNM); New York:
Kingston, Jun 8, 1939, Esselbaugh, H. R. Dodge Collection (1 Q; USNM); New York:
New Rochelle, IV.20.1932, L. Lacey collector (3 Q; USNM); Ohio: Chillicothe, 28-VI-
50, Liriodendron tulipifera (3 Q; USNM); Ohio: Moreland Hills, 3.VIII.1981, W. V.
Miller (1 Q; CAS); Virginia: Albemarle Co., Ivy ex Vitis, cane tunnels, 29 Apr 1983, D.
G. Pfeiffer coll. (2 Q; USNM). ORIENTAL REGION: China: S. China: Fukien,
Shaowu, Tachulan, V.14.1942, T. Ma (2 Q; CAS). Japan: Japan: III-1917, Van Dyke
Collection, in trunk of young chestnut (1 Q; CAS); Japan, Hokkaido, Gamushi,
V1.9.1956, Akira Nobuchi (2 Q; USNM); JAPAN: HONSHU, Higashiaraya, Kushibiki-
machi, Yamagata Pref, 13.Vi.1996, K. Domon leg., Host tree: Pyrus scrotina Rehder (1
Q, 2 6; RAB); JAPAN: KYUSHU, Shiiya-touge Pass, Miyazaki-Kumarnoto Pref.,
25.Vi.1994, H. Goto leg. (1 Q; RAB); Japan: Matsukawa, 27-VII-1980, S. L. Wood (1 Q;
USNM); Japan: Tamagowa, 29-VII-1980, S. L. Wood, Fagus cranatus (6 Q; USNM);
Japan: Tokyo, Mt. Takao, IX.22.1950, Akira Nobuchi (2 Q; USNM). Thailand:
Thailand: Chiang Mai: Doi Suthep, 7.Viii.02, Beaver, K. Koivisto (2 Q; RAB).
PALEARCTIC REGION: Germany: Gerrnania (Hessen) 28.8.52, Darmst-Kranichst,
Geishecke, leg. F. Groschike, Quercus (1 6; NHMW, 1 6; BMNH); 1GB — Sengscheid,

SAAR, Wald, 18.6.90, leg. Mosbacher (4 Q; RAB). Switzerland: Helvetia. Arlesheim,

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1988, In traps (2 Q; RAB). OCEANIA: Hawaiian Islands: USA: HI: O’ahu I.. N.
Halawa Valley, NW of Honolulu, 390 m, 21°25'N, 157°51'W, 1 1-29.VI.1991.
FMHD#91-4, Met. Polym.-Psidium-Hibiscus mixed nat./exotic forest, A. Newton and M.

Thayer, #869, window trap (4 Q; FMNH).

X ylosandrus hirsutipennis (Schedl)
(Fig. 2.25)

Xyleborus hirsutipennis Schedl, 1961b: 144. Holotype Q: Madagascar, Perinet. Montague
d’Ambre, Antaniditra: IRSM.
Xylosandrus hirsutipennis (Schedl): Wood and Bright, 1992: 796.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors failed to indicate it as a “new combination.”
Diagnosis. Female 1.9 — 2.2 mm long; 2.3 times longer than wide. Body brown to dark
brown; antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum of equal length and width. Dorsal
aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-
appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presence of a pronotal-
mesonotal mycangium. Pronotal disc moderately punctate. Lateral aspect of pronotum
basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa, not carinate. Procoxae
widely separated. Protibiae with 5 socketed teeth on lateral margin; mesotibiae with 7

socketed teeth; metatibiae with 7 socketed teeth. Elytra 1.3 times longer than wide: 1.3

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times longer than pronotum. Discal striae punctate; interstriae uniseriate punctate.

Elytral disc gradually curving into declivity. Declivity convex. lateral margins carinate to
7th interstriae. Five striae visible on declivity. Striae granulate, with semi-appressed.
hair-like setae, longer than the width of second declivital interstriae. Interstriae
granulate, multiseriate, with semi-appressed, hair-like setae. longer than the width of
second declivital interstriae.

This species is morphologically similar to X crassiusculus (Fig. 2.16). Both
species have a declivital face that is matte in appearance due to densely, finely, and
confusedly granulate striae and interstriae. X ylosandrus hirsutipennis can be
distinguished from X crassiusculus by the following characters: frons punctate;
pronotum with a lateral carina; pronotal disc glabrous, except for mycangial setae; elytral
disc uniseriate punctate; five striae visible on elytral declivity; and declivital striae and
interstriae with semi-appressed, hair-like setae, longer than the width of second declivital
interstriae.

Distribution. AFROTROPICAL REGION: Madagascar.

Hosts. Cassipourea Aubl. sp., Ficus soroceoides Baker var. macrophlebia H. Perrier,
Harungana madagascariensis Poir., Psychotria L. sp., Urophyllum lyallii Baker,
Vernonia Schreb. sp.

Specimens Examined. (3 Q; 1 6)

Type material: Paratypes Xyleborus hirsutipennis: Madagascar, Perinet, 21.XI.1952, Dr.
K. E. Schedl (2 Q; NHMW); Madagascar, Montagne d’Ambre, 7.XII.1952, Dr. K. E.
Schedl (1 6; NHMW); Madagascar, Montagne d’Ambre, 5.XII.1952, Dr. K. E. Schedl

(1 Q;NHMW).

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Xylosandrus hulcri Dole and Cognato, new species
(Fig. 2.26)

Description. Female (Fig. 2.26): Body oval, 2.4 - 2.7 mm long, 2.0 times longer than
wide, bicolored, with pronotum and elytral apices dark brown, dorsal 1/3 and lateral 2/3
of elytra yellowish brown, ventral side and appendages dark brown. Frons convex.
coriaceous, sparsely punctuate between eyes, long erect hair-like setae originating from
punctures, weak ridge at middle of frons just above epistoma. Epistoma with dense row
of long and short hair-like setae along lower margin. Eyes emarginate. Antennal funicle
5-segmented, scape and funicle with spare, short, hair-like setae; club obliquely truncate,
first segment sclerotized, forming a circular costa (type I, Hulcr et al. 2007), circular
costa closed posteriad, oblique part of club densely pubescent, second segment not
comeous; posterior face of club covered entirely by first segment (type 1, Hulcr et al.
2007). Pronotum 0.9 times longer than wide, rounded dorsally (type 1, Hulcr et al.
2007), widest about one-half pronotal length from base, anterior half broadly rounded and
then tapering slightly toward apex, basal angles rounded, anterior margin with 4-6
asperities; anterior slope densely aspirate, aperities smallest at summit and increasing in
size toward anterior margin; disc moderately punctate, lacking a patch of denser
punctures medially at base, background sculpture finely granulate; lateral aspect of
pronotum rounded (type 1, Hulcr et al. 2007), lateral costa extending approximately two-
thirds pronotal length, lateral carina absent; pronotal vestiture of long and short,
moderately appressed, hair-like setae. Scutellum triangular, yellowish-brown to dark

brown, flush with surface of elytra. Elytra 1.2 times longer than wide. 1.4 times as long

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as pronotum, shining, parallel-sided on basal two-thirds and then broadly rounded toward
apex. Striae not impressed, shallowly and regularly punctuate, long erect hair-like setae
originating from punctures. Interstriae twice the width of striae, finely punctuate,
confused, long (1.5 - 2 times width of interstria) and short (less than width of interstria)
erect hair-like setae originating from punctures. Declivity commencing behind mid-point
of elytra, gradually separating from disc; lateral margins carinate to 7th interstriae.
Procoxae widely separated. Protibiae with 5 socketed teeth on lateral margin; mesotibiae
with 7 - 8 socketed teeth; metatibiae with 8 socketed teeth. Abdominal ventrites evenly
punctured, punctures with long, erect, hair-like setae.
Specimens Examined. (4 Q; 0 6)
Type Material. Holotype Q: PAPUA NEW GUINEA, Chimbu, Mu Village, March
2006, 1600 m asl. Hulcr and Coganto coll. Ficus mollior. Vila 1462. In MSU. Paratypes
Q: PAPUA NEW GUINEA, Chimbu, Mu Village, March 2006, 1600 m asl. Hulcr and
Coganto coll. Ficus mollior. Vila 1462. One paratype in F ICB. Three paratypes in MSU.
One paratype female was used for DNA extraction, and remains are vouchered at MSU.
Male: Unknown.
Etymology. This species is named for Jiri Hulcr (Michigan State University), who
collected the type series and whose collecting efforts have contributed greatly to our
knowledge of tropical scolytine fauna.
Diagnosis. This species can be readily distinguished from all other known Xylosandrus
species by its distinct color pattern, with a testaceous patches on the elytra basally and
laterally.

Distribution. AUSTRALIAN REGION: Papua New Guinea.

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Hosts. Ficus mollior Benth.

Discussion. This species is known from a single collecting event. Specimens were
collected at the same locality, but from galleries in several host trees. This species is
mycocleptes and has been found in association with and stealing ambrosial fungus from
Xylosandrus rotundicollis. Phylogenetic analysis places it as sister to the Australian

species, X queenslandi.

Xylosandrusjaintianus (Schedl)
(Fig. 2.27)

Xyleborusjaintianus Schedl, 1967: 161. Holotype Q: Shillong, Assam, C. F. C. Beeson,
22.V.1925; NHMW.
Xylosandrusjaintianus (Schedl): Wood and Bright, 1992: 796.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992). but
the authors failed to indicate it as a “new combination.”
Diagnosis. Female 3.0 mm long; 2.0 times longer than wide. Body brown; antennae
and legs same color as body. Frons rugose, with distinct median keel between eyes.
Antennae with 5 funicular segments. Antennal club obliquely truncate; first segment
forming a circular costa; segment one covering entire posterior face. Pronotum 0.9 times
longer than wide. Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007).
Pronotal vestiture of semi-appressed, hair-like setae; setae less dense on disc. Basal
pronotum with a dense patch of short, erect setae, indicating the presense of a pronotal-
mestonotal mycangium. Pronotal disc densely asperate-granulate, with sculpture

separated by distance less than or equal to their width. Lateral aspect of pronotum basic

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(type 0, Hulcr et al. 2007). Pronotum with lateral costa, not carinate. Procoxae widely
separated. Protibiae with 5 socketed teeth on lateral margin; mesotibiae with 11 socketed
teeth; metatibiae with 11 — 12 socketed teeth. Elytra 1.1 times longer than wide; 1.1
times longer than pronotum. Discal striae punctate; interstriae multiseriate punctate.
Declivital face of elytra steep and abruptly separated from disc. Declivity with depressed
areas on each side of raise interstriae, lateral margins carinate to 7th interstriae. Four
striae visible on declivity. Striae coarsely granulate, with appressed, hair-like setae,
shorter than the width of second declivital interstriae. Interstriae granulate, multiseriate.
with appressed, hair-like setae, shorter than the width of second declivital interstrie.

This species is morphologically similar to X beesoni (Illustrated in Saha et al.
1992), X discolor (Fig. 2.20), and X subsimiliformis (Fig. 2.40). Xylosandrusjaintianus
can be distinguished from these species by the following characters: first and second
declivital interstriae elevated toward apex, with depressed areas on each side of raised
interstriae; frons rugose with a distinct median keel.
Distribution. ORIENTAL REGION: Burma, India (Assam). PALEARCTIC
REGION: Nepal.
Hosts. Unknown.
Specimens Examined. (4 Q; 0 6)
Type material: Holotype X yleborus jaintianus (Q; NHMW). Paratype Xyleborus
jaintinaus: N. E. Burma: Kambaiti, 7000ft., 23/5/1934, R. Malaise (Q; BMNH).
Other material: PALEARCTIC REGION: Nepal: NEPAL-HIMALAYA Annapurna

mts. N-Pokhara, Madi-Khola-Tal, 1850m, 2.v.1996. leg. O. Jager (1 Q; RAB); NEPAL.

141

Kali Gandaki Tai 2 km SO Narcheng oberth.Rele Khola, HO-Hang, 2300 m, N28°30'40".

E83°41'33", 25.v.2001, leg. o. Jager (1 9; RAB).

Xylosandrus mancus (Blandford)
(Fig. 2.28)

Xyleborus mancus Blandford, 1898: 428. Holotype Q: Ceylon. E. E. Green; BMNH.
Apoxyleborus mancus (Blandford): Wood 1980: 90.
Xylosandrus mancus (Blandford): Wood 1984: 229.
Xyleborus abruptus Sampson, 1914: 388. Syntypes Q: Seychelles, Mahe: high forest of
Morne Blanc, and Cascade Estate; BMNH. Synonymy: Schedl, 1951: 51.
Xyleborus mancusformosanus Eggers, 1930: 186. Holotype Q: Formosa: Taihoku; FRI.
Synonymy: Schedl, 1952: 61.
Notes. This species was first included in Xylosandrus, when Wood (1984) synonymized
Apoxyleborus with X ylosandrus.
Diagnosis. Femle 2.9 — 3.3 mm long; 1.2 — 1.4 times longer than wide. Body yellowish
brown to brown; elytra darker brown at apex and declivity; legs and antennae the same
color as body. Frons punctate. Antennae with 5 funicular segments. Antennal club
obliquely truncate; first segment forming circular costa; segment one covering entire
posterior face. Of approximately equal length and width. Dorsal aspect of pronotum
rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-appressed, hair-like setae;
setae less dense on disc. Basal pronotum with a dense patch of short, erect setae,
indicating the presence of a pronotal-mesonotal mycangium. Pronotal disc moderately

punctate. Lateral aspect of pronotum basic (type 0, Hulcr et a1. 2007). Pronotum with

142

lateral costa, not carinate. Procoxae widely separated. Protibiae with 5 socketed teeth on
lateral margin; mesotibiae with 11 socketed teeth; metatibiae with 12 socketed teeth.
Elytra 1.2 times longer than wide; 1.2 times longer than pronotum. Discal striae
punctate; interstriae multiseriate punctate. Declivital face of elytra steep and abruptly
separated from disc. Declivity flattened, lateral margins carinate, with carina extending
beyond 7th interstriae, forming a circumdeclivital ring. Four striae visible on declivity.
Straie punctate, without setae. Interstriae granulate, without setae.

This species is one of two Xylosandrus with the margin of the elytral declivity
with a carina or rim of granules that extends beyond the 7‘h interstriae, forming a
circumdeclivital ring: X amputatus (Fig. 2.8) and X mancus. Xylosandrus mancus can
be distinguished from X amputatus by the following characters: 2.9 — 3.3 mm long; 1.2
— 1.4 times as long as wide; declivtial striae with a row of large, shallow punctures,
arranged in a somewhat wavy line; and declivital interstriae shining, not densely
granulate.
Distribution. AF ROTROPICAL REGION: Madagascar, Mauritania, Seychelles
Islands, Tanzania. ORIENTAL REGION: China (Gansu, Xizang [Tibet]),
Maharashtra, Inidia (Tamil Nadu), Indonesia (Java, Sumatra), Japan, Malaya, Philipine
Islands, Sri Lanka, Taiwan, Vietnam.
Hosts. Adenanthera pavonina L., A lbizzia Benth. sp., Anacardium occidentale L.,
Aphanamixis rohituka Pierre, Artocarpus dadak Miq., Brackenridgea hookeri (Planch.)
A. Gray, Buchanania lanzan Sprenq., B. latifolia Roxb., Buteafrondosa Wall.,
Calophyllum inophyllum L., Cassia fistula L., Cordia dichotoma G. Forst, C ordia myxa

L., Dalbergia latifolia Roxb., Drypbalanops aromatica Gaertn., D. oblongifolia Dyer,

143

Gomphia serrata (Gaertn.) Kanis, Grewia paniculata Roxb., Hibiscus macrophyllus
Roxb., Hopea beccariana Burck, H. ferrea Heim, Hullettia dumosa King, Khaya
senegalensis Juss., Litsea megacarpa Gamble, Magnififera indica L., Melanorrhoea Wall.
sp., Nephelium lappaceum L., Palaquium gutta Burk., Pometia pinnata J. R. Forst. and
G. Forst, Quercus L. sp., Shorea bracteolate Dyer, S. leprosula Miq., S. macroptera
Dyer, S. sumatrana (Slooten) Desch, Styrax benzoin Dryand., Swietenia macrophylla
King, S. mahagoni (L.) Jacq., T ectona grandis L., Theobroma cacao L., Toonia sureni
Merrill., Tristania whitiana Griff, Vateria copallifera (Retz.) Alston, Vitex pubescens
Vahl.

Specimens Examined. (69 Q; 2 6)

Type material: Holotype X yleborus mancus (Q; BMNH).

Other material: AF ROTROPICAL REGION: D. O. Africa (1 Q; NHMW).
Madagascar: Madagascar, Ambila, 28.XI.1952, K. E. Schedl (1 Q. 1 6; NHMW).
Seychelles: Seychelles: P. R. Dupont, 1915, attacking Cashew (3 Q; BMNH).
ORIENTAL REGION: China: Salagar, Kafang, ex Clenderai, 5. XII.1948, F. G.
Browne (3 Q; USNM). Indonesia: Java: Kediri Forest, 11.1925, L. G. E. Kalshoven (2
Q; USNM); Java: Kediri, 111.1925, L. G. E Kalshoven (1 Q; NHMW). Japan: Japan:
Kagoshima Pref., Tarumizuy, Oonohara, Broaddleaf forest, 425 m, 14 May 2001.
Yoshikazu Sato coll., ex. ETOH-baited trap (2 Q; MSU); Japan: Kagoshima Pref.,
Tarumizuy, Oonohara, Broaddleaf forest, 425 m, 3 July 2001, Yoshikazu Sato coll.. ex.
ETOH-baited trap (1 Q; MSU). Malaysia: Sarawak, Bako National Park, 29 Oct - 2
Nov 1998, 50 m, B. Jordal coll. (1 Q; MSU); Malaya, Selangor, Kepong, 8.1V.1934,

Selenager Mus., ex. Styrax benzoin (1 Q; NHMW). Philipine Islands: Manila, Philipine

144

Islands, Colln PC, McGregow (1 Q; USNM); Philipine Islands: Puerto Princesa. Palawan
Is., sea level, 2nd growth forest, IV.29.47, H. Hoogstraal leg. (1 Q; FMNH). Singapore:
Singapore, Bukit Timah, 50 m, 25—27 Oct 1998, B. H. Jordal coll., C innamomum (1 Q:
MSU). Sri Lanka: Sri Lanka: Bad. Dist., Buttala, 5 June 1975, S. L. Wood, collected
from Liana (5 Q; USNM); Sri Lanka: Bad. Dist., Butatala, 5 June 1975, S. L. Wood.
collected from pole (8 Q, l 6; USNM); Sri Lanka: Bat. Dist., Unnichchai, 9 June 1975,
S. L. Wood, collected from Liana (2 Q; USNM); Ceylon [Sri Lanka]: Col. Dist.
Tunmodera, 200 ft, l7-XI-1970, O. S. Flint, Jr. (2 Q; UISNM); Sri Lanka: Gal. Dist.,
Karmeliya, 250 mtrs., 23 May 1975, S. L. Wood, Collected from pole (9 Q; USNM);
Ceylon [Sri Lanka]: Hantane, XII.1962, D. Calnaido (1 Q; NHMW); Ceylon [Sri Lanka]:
Kan. Dist., Kandy, 1-15 March 1971, Piyadasa and Somapala (4 Q; USNM); Ceylon [Sri
Lanka]: Kan. Dist., 5 mi NW Mahiyangana, 30 Mar-9 Apr 1971, P. and P. Spangler (2 Q;
USNM); Sri Lanka: Keg. Dist., Kitulgala, 200 m, 30 May 1975, S. L. Wood, host:
Osbeckia aspera (1 Q; USNM); Sri Lanka: Matte. Dist., 5 km SE Naula, 200 mtrs, 14
June 1975, S. L. Wood, misc. hosts (4 Q; USNM); Sri Lanka: Matte. Dist., 5 km SE
Naula, 200 mtrs, 14 June 1975, S. L. Wood, collected from Liana (1 Q; USNM); Sri
Lanka: Mon. Dist. 8 km NW Bibile, 50 mtrs, 7 June 1975, S. L. Wood, collected from
pole (5 Q; USNM); Sri Lanka: Mon. Dist., Buttala, 50 mtrs, 6 June 1975, S. L. Wood.
collected from pole (1 Q; USNM); Sri Lanka: Pol. Dist. 32 km N Polonnaruwa, 11 June
1975, S. L. Wood, collected from pole (1 Q; USNM); Sri Lanka: Rat. Dist. Gilimale, 17
May 1975, S. L. Wood, host: Myristica dactyloides (1 Q: USNM); Ceylon [Sri Lanka]:

Rat. Dist., Uggaalkaltota 350 ft., irrigation bungalow. 31 Jan — 8 Feb 1970, Davis and

145

Rowe (1 Q; USNM); Ceylon [Sri Lanka], Sabagamuva, Millawitiya Est, 3-10.Vii.1956.

E. Judenko (1 Q; NHMW).

Xylosandrus mediocris (Schedl)
(Fig. 2.29)

Xyleborus mediocris Schedl, 1942a: 185. Lectotype Q: Malaya. N.S. Pasoh Forest
Reserve, 12-11-193 8, ex Shorea dasyphylla; NHMW.
Xylosandrus mediocris (Schedl): Browne, 1963: 55.
Diagnosis. Female 1.4 mm long; 2.3 times longer than wide. Body brown; antennae and
legs yellowish brown. Frons not visible on specimen examined. Antennae with 5
funicular segments. Antennal club obliquely truncate; first segment forming a circular
costa; segment one covering entire posterior face. Pronotum of equal length and width.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense
patch of short, erect setae, indicating the presence of a pornotal-mesonotal mycangium.
Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type 0, Hulcr et al.
2007). Pronotum with lateral costa and carina. Procoxae not visible on specimen
examined. Pro-, meso- and metatibial teeth not visible on specimen examined. Elytra 1.3
times longer than wide; 1.3 times longer than pronotum. Discal striae punctate;
interstriae uniseritae punctate. Elytral disc gradually curving into declivity. Declivity
convex, lateral margins carinate to 7'h interstriae. Six striae visible on declivity. Striae
punctate, without setae. Interstriae granulate, uniseriate, with erect. hair-like setae.

longer than the width of second declivital interstriae.

146

This species is morphologically similar to X adherescens (Fig. 2.7), X eupatorii
(Fig. 2.22), and X germanus (Fig. 2.24). Xylosandrus mediocris can be distinguished
from these species by the following characters: 1.4 mm long; pronotum of equal length
and width; and declivital interstriae uniseriate granulate, with erect, hair-like setae. longer
than the width of second declivital interstriae.
Distribution. ORIENTAL REGION: Malaysia.
Hosts. Dipterocarpus cornutus Dyer, Shorea dasyphylla Foxw..
Specimens Examined. (1 Q; 0 6)

Type material: Lectotype Xyleborus mediocris (Q; NHMW).

Xylosandrus mesuae (Eggers)

(Fig. 2.30)
Xyleboruss mesuae Eggers, 1930: 182. Holotype Q: Bengal (Kalimpong), Aug. 1910 on
Mesuaferra; FRI.
Xylosandrus mesuae (Eggers): Browne, 1963: 55.
Diagnosis. Female 1.1 - 1.3 mm long; 2.2 times longer than wide. Body bicolored,
pronotum distinctly lighter than elytra; pronotum light brown and elytra dark brown;
antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.8 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense

patch of short, erect setae, indicating the presence of a pronotal-mesonotal mycangium.

147

Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type 0, Hulcr et al.
2007). Pronotum with lateral costa, not carinate. Procoxae widely separated. Protibiae
with 4 socketed teeth on lateral margin; mesotibiae with 6 socketed teeth; metatibiae with
8 socketed teeth. Elytra 1.3 times longer than wide; 1.6 times longer than pronotum.
Discal striae punctate; interstriae uniseriate, very finely punctate. Elytral disc gradually
curving into declivity. Declivity convex, carinate to 7’h interstriae. Six striae visible on
declivity. Striae punctate, without setae. Interstriae granulate, uniseriate. with erect,
hair-like setae, longer than the width of second declivital interstriae.

This species is one of four bicolored Xylosandrus with the pronotum distinctly
lighter than the elytra: X arquatus (Fig. 2.9), X discolor (Fig. 2.20), X ferinus (Fig.
2.23), and X mesuae. Xylosandrus mesuae can be distinguished from X discolor by the
following characters: elytral disc gradually curving into declivity. Xylosandrus mesuae
can be distinguished from X arquatus and X ferinus by the following characters: 1.1 —
1.3 mm long; declivital striae with setae; declivital interstriae with erect, hair-like setae.
longer than the width of second declivital interstriae.

Distribution. AUSTRALIAN REGION: Papua New Guinea. ORIENTAL REGION:
India (Bengal, Uttar Pradesh), Sri Lanka.

Hosts. Diptericarpus zeylanicus Thwaites., Macaranga Thou. sp., Mesuaferrea L.,
Osbeckia aspera Benth., Shorea robusta Roth.

Specimens Examined. (102 Q; 1 6)

Type material: Cotype X yleborus mesuae: Bengal, Kalimpong on Mesuaferra (Q;

NHMW).

148

Other material: AUSTRALIAN REGION: Papua New Guinea: Papua New Guinea,
Lae, 15 km S Lae, 100 m, 13.2.2003, B. Jordal and A. Sequeira leg. ex Macaranga
petoles (3 Q; RAB). ORIENTAL REGION: Sri Lanka: Ceylon [Sri Lanka]: 10-
16.VII.1956, Sabargamuva, Millawitiya Est., E. Judenko (1 Q; NHMW); Sri Lanka: C 01
Dist. Labugama, 23 June 1975, S. L. Wood, collected from legume tree (2 Q; USNM);
Sri Lanka: Col. Dist. Labugama, 23 June 1975, S. L. Wood, collected from twigs (64 Q;
USNM); Sri Lanka: Col. Dist. Labugama, 23 June 1975, S. L. Wood, host: Dicterocarpus
zeylanicus (25 Q, 1 6; USNM); Sri Lanka: Keg. Dis. Kitulgala, 200 m, 30 May 1975, S.

L. Wood, host: Osbeckia aspera (6 Q; USNM).

X ylosandrus metagermanus (Schedl)
(Fig. 2.31)

Xyleborus metagermanus (Schedl), 1951: 58. Holotype Q: U. Dihing Res., Lakhimpur
Assam, 6. VIII. 1931, ex Gmelina arborea, C. F. C. Beeson; NHMW.
Xylosandrus metagermanus (Schedl): Browne, 1963: 55.
Diagnosis. Female 1.8 mm long; 2.0 times longer than wide. Body light brown to
brown; antennae and legs same color as body. Frons rugose. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.8 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presense of a pronotal-

mesonotal mycangium. Pronotal disc moderately punctate. Lateral aspect of pronotum

149

basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa, not carinate. Procoxae
widely separated. Protibiae with 5 socketed teeth on lateral margin; meso- and metatibial
teeth not visible on specimen examined. Elytra 1.1 times longer than wide; 1.2 times
longer than pronotum. Discal striae punctate; interstriae uniseriate punctate. Elytral disc
gradually curving into declivity. Declivity convex, lateral margins carinate to 7‘h
interstriae. Six striae visible on declivity. Striae punctate, without setae. Interstriae
granulate, uniseriate, with erect, hair-like setae, longer than the width of second declivital
interstriae.

This species is morphologically similar to X mixtus (Fig. 2.32). Xylosandrus
metagermanus can be distinguished from X mixtus by the following characters: 1.8 mm
long; six striae visible on declivity; declivital striae without setae; declivital interstriae
uniseriate granulate.

Distribution. ORIENTAL REGION: India (Assam).
Hosts. Gmelina arborea Roxb.
Specimens Examined. (1 Q; 0 6)

Type material: Holotype X yleborus metagermanus (Q; NHMW).

Xylosandrus mixtus (Schedl), new combination
(Fig. 2.32)
Xyleborus mixtus Schedl, 1979b: 108. Holotype Q: Papua, Butolo, Morobe Distr., Upper
Monki, L. A., 16.3.1973, sticky trap Nr. 1604, F. R. Wylie and P. Shanahan; NHMW.

Amasa mixtus (Schedl): Wood and Bright, 1992: 683.

150

Notes. This species was first included in Amasa by Wood and Bright (1992). but the
authors failed to indicate it as a “new combination.”
Diagnosis. Female 2.6 — 2.7 mm long; 2.4 times longer than wide. Body light brown;
antennae and legs same color as body. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa:
segment one covering entire posterior face. Pronotum 0.9 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense
patch of short, erect setae, indicating the presence of a pronotal-mesonotal mycangium.
Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type 0, Hulcr et al.
2007). Pronotum with lateral costa, not carinate. Procoxae narrowly, but completely
separated. Protibiae with 4 socketed teeth on lateral margin; mesotibiae with 9 socketed
teeth; metatibiae with 10 socketed teeth. Elytra 1.4 times longer than wide; 1.6 times
longer than pronotum. Discal striae punctate; interstriae mutliseriate punctate. Elytral
disc gradually curving into declivity. Declivity convex, carinate to 7th interstriae. Five
striae visible on declivity. Striae punctate, with semi-appressed, hair-like setae, shorter
than the width of second declivital interstriae. Interstriae granulate, mutiseriatae, with
erect, hair-like setae, more than twice as long as width of second declivital interstriae.
This species is morphologically similar to X metagermanus (Fig. 2.31).
Xylosandrus mixtus can be distinguished from X metagermanus by the following
characters: 2.6 — 2.7 mm long; five striae visible on elytral declivity; declivital striae with
semi-appressed, hair-like setae; declivital interstriae multiseriate granulate.

Distribution. AUSTRALIAN REGION: New Guinea.

151

Hosts. Unknown.
Specimens Examined. (1 Q; 0 6)

Type material: Holotype Xyleborus mixtus (Q; NHMW).

X ylosandrus monteithi Dole and Beaver
(Fig.2.33 and 2.34)

Xylosandrus monteithi Dole and Beaver, in press. Holotype Q: AUSTRALIA,
Queensland, Palmerston, Henrietta Cr., 550m, ex unknown tree, 22.1.2000 (B. Jordal). In
QMB (Accession # T144402).
Diagnosis. Female 3.0 — 3.4 mm long; 2.1. times longer than wide. Body dark brown or
blackish, base of pronotum and elytra lighter brown, ventral side and appendages
yellowish brown. Frons retiuculate with coarse punctures. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.8 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of erect
and semi-appresssed long hair-like setae; pronotal disc densely setose, setae as dense as
on anterior pronotum. Pronotum lacking a dense patch of setae at base of pronotum.
Pronotal disc moderately punctate. Lateral aspect of pronotum rounded (type 1, Hulcr et
al. 2007). Pronotum with lateral costa, not carinate. Procoxae widely separated.
Protibiae with 4 socketed teeth on lateral margin, meso- and metatibiae with 7 — 9
socketed teeth. Elytra 1.2 times longer than wide; 1.6 times longer than pronotum.
Discal striae punctate; interstraie mutliseriate punctate, becoming granulate toward

declivity. Elytral disc gradually curving into declivity. Declivity convex, lateral margins

152

rounded, without a carina or a row of tubercles or serrations. Six striae visible on
declivity. Striae punctate, with semi-appressed, hair-like setae, longer than the width of
second declivital interstriae. Insterstriae granulate, multiseriate, with semi-appressed,
hair-like setae, longer than the width of second declivital interstriae.

This species is one of two Xylosandrus with the margin of the elytral declivity
rounded: X monteithi and X rotundicollis (Fig. 2.39). Xylosandrus monteithi can be
distinguished from X rotudndicollis by the following characters: 3.0 — 3.4 mm long;
basal pronotum lacking a dense patch of setae; declivital striae with semi-appressed, hair-
like setae, longer than the width of second declivital interstriae; and declivital interstriae
muliseriate granulate, with semi-appressed, hair-like setae, longer than the width of
second declivital interstriae. Xylosandrus moteithi may also be confused with X woodi.
but the later species has small tubercles marking the lateral declivital margin.
Distribution. AUSTRALIAN REGION: Queensland.

Hosts. Unknown.

Specimens Examined. (1 Q; 1 6)

Type material: Paratype Xylosandrus monteithi: Auatralia, Queensland, Palmerston,
Watchua Falls, Jan. 2000, 550 m, ex unknown tree, 24.1 B. Jordal leg (Q; MSU).
Allotype Xylosandrus monteithi: Auatralia, Queensland, Palmerston, Watchua Falls. Jan.

2000, 550 m, ex unknown tree, 24.1 B. Jordal leg. (6; MSU).

X ylosandrus morigerus (Blandford)

(Fig. 2.35)

Xyleborus morigerus Blandford, 1894a: 264. Syntypes Q: probably New Guinea: BMNH.

153

Xylosandrus morigerus (Blandford): Reitter, 1913: 83.

Xyleborus coffiaae Wurth, 1908: 199. Syntypes Q: Java; type location unknown.
Synonymy: Strohmeyer, 1910: 86.

Xyleborus luzonicus Eggers, 1923: 174. Lectotype Q: Mt. Makiling, lnsel Luzon.
Philippinen; USNM.

Xylosandrus luzonicus (Eggers): Browne, 1963: 55. Synonymy: Wood, 1974: 287.
Xyleborus diflicilis Eggers, 1923: 174. Lectotype Q: Java, Hagedom coll., 1915; USNM.
Xylosandrus difficilis (Eggers): Browne, 1963: 55. Synonymy: Synonymy Bright and
Skidmore 1997: 4, 169.

Xyleborus abruptoides Schedl, 1955: 298. Holotype Q: Fiji: Viti Levu, Navai Mill, near
Nandarivatu, 2500 ft., 15.IX.1938, beating shrubbery; BPBM.

X ylosandrus abruptoides (Schedl): Browne, 1963: 55. Synonymy (= X ylosandrus
diflicilis) Beaver 1995: 17.

Diagnosis. Female 1.5 — 2.0 mm long; 2.1 times longer than wide. Body light to dark
brown; antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa:
segment one covering entire posterior face. Pronotum 0.9 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense
patch of short, erect setae, indicating the presence of a pronotal-mesonotal mycangium.
Pronotal disc moderately punctate basally. Lateral aspect of pronotum rounded (type 1,
Hulcr et al. 2007). Pronotum with a lateral costa and carina. Procoxae widely separated.

Protibiae with 4 socketed teeth on lateral margin; mesotibiae with 8 — 10 socketed teeth;

154

metatibiae with 10 socketed teeth. Elytra 1.2 times longer than wide; 1.4 times longer
than pronotum. Discal striae punctate; interstriae uniseriate punctate. Declivital face
steep and abruptly separated from disc. Declivity convex, lateral margins carinate to 7lh
interstriae. Six striae visible on declivity. Striae punctate, with erect hair-like setae.
shorter than the width of second declivital interstriae. Interstriae punctate, uniseriate.
with erect, hair-like setae, longer than twice the width of second declivital interstriae.
This species is morphologically similar to X derupteterminatus (Fig. 2.18) and X
terminatus (Fig. 2.41). Xylosandrus morigerus can be distinguished from these species
by the following characters: elytral declivity convex; six striae visible on declivity:
declivital striae with setae; and declivital interstriae uniseriate punctate, with erect, hair-
like setae, longer than twice the width of second declivital interstriae.
Distribution. AF ROTROPICAL REGION: Gabon, Madagascar, Mauritius Islands,
Zaire. AUSTRALIAN REGION: New Britain Island, New Guinea, Solomon Islands
Queensland. NEOTROPICAL REGION: Brazil, Colombia, Costa Rica, Ecuador
(Santa Cruz in Galapagos Islands), Honduras, Mexico (Campeche, Chiapas. Oaxaca.
Tabasco, Varacruz), Panama, Puerto Rico, Tobago, Venezuela. OCEANIA: Fiji Islands,
Hawaiian Islands, Micronesia (Caroline Islands, Timor in Mariana Islands), Samoan
Islands, Tonga (Vava’u). ORIENTAL REGION: India (Bengal, Tamil Nadu),
Indonesia (Borneo, Celebes, Java, Sumatra), Malaysia (Sarawak), Philippine Islands, Sri
Lanka, Taiwan, Vietnam (Tonkin Island, Vietnam). PALEARCTIC REGION
(Introduced): Austria, Czechoslovakia, England, France, Italy, Jordan, Lebanon.
Hosts. Acacia gaumeri S. F. Blake, Acalypha L. sp., A ctinophorafragrans Wall.,

Adenanthera pavonina L., Albizziafalcate (L.) Backer, A. glauca Benth., A. procera

155

Benth, Albizia Durazz. sp., Alseis yucatanensis Standl., Altingia excelsa Noronha.
Arthrophyllum diversifolium Blume, Amomum L. sp., Aspidosperma Mart and Zucc. sp.,
Astronium graveolens Jacq., Bixa orellana L., Boehmeria Jacq. sp., Bridelia Willd. sp..
Brosimum alicastrum Sw., Bursera simaruba (L.) Sarq., Butea monosperma Kuntze,
Calamus L. sp., C. caesius Blume, Calophyllum brasiliense Camb., Camellia sinensis
Kuntze, C. thea Link, C. theifera Dyer, Cassia multijuga Rich., Castanea argentea
Blume, Castanopsis (D. Don.) Sprach. sp., Cattleya Lind. sp., Cedrela odorata L.,
Cedrela P. Borwne sp., Cecropia obtusifolia Bertol., Ceiba pentandra (L.) Gaertn. ,
Centrosema plumieri Benth., Cecropia Loefl. sp., Chrysophyllum cainito L., C inchona L.
sp., Claoxylon polot Merr., Clidemia hirta Don., Cocos nucifera L., C oflea arabica L.,
Coffea excelsa Cheval., “C. hybrida” (Java: Schedl 1962) , C. liberica Bull. ex Schum..
C. robusta L. Linden, Cola acuminate Schott and Endl., Cordia dodecandra DC..
Crotalaria L. sp., C. anagyroides Kunth, C. usaramoensis Baker , Dalbergia latifolia
Roxb., Dendrobium SW. sp., D. phalaenopsis Fitzq., D. superbum Rchb., D. veratrifolium
Lindl., Derris microphylla (Miq.) B. D. Jacks, Didymopanax Decne. and Planch. sp.,
Dryobalanops oblongifolia Dyer, Endospermum malaccense Muell.-Arg., Epidendrum
stamfordianum Bateman , Erythrina lithosperma Miq. Var, inermis de. and Val.,
Erythroxylon novogranatense Hieron., Esenbeckia pentaphylla Griseb., Eugenia
polyantha Phil., Eupatorium pallescens DC., Eusideroxylon zwageri Teijsm. and Binn.,
Ficus L. sp., Ficus ampelas Burm., F issistigma elegans Merr., F lemingia strobilifera
(L.) W. T. Aiton, Freycinetia hombroni Martelli, Fuchsia L. sp., Glochidion J. R. Forst.
and G. Forst. sp., Grewia laevigata Vahl, Gynotroches onillaris Blume, Hevea

brasiliensis Muell.-Arg., Intsia palembanica Miq., “Laurel roja” (Venezuela). Lecythis

156

Loefl. sp., Leucaena glauca Benth., Licania hypoleuca Benth., Lonicera caprijolium L.,
Macaranga Thou. sp., Machaerium cirrhiferum Pittier, “Mahogany” (Fiji), Marumia
muscosa Blume, Melia azedarach L., Miconia trinervia Coqn., Ochroma lagopus Sw..
“Palito negro” (Venezuela), Persea gratissima Gaertn., Phalaenopsis Blume sp., Pometia
pinnata Forst., Pouteria sapota (Jacq.) H. E. Moore and Steam, Quararibea Aubl. sp..
Renathera storiei Rchb., Sambucusjavanica Reinw. ex Blume, Schizolobium parahyba
(Vell.) Blake, Schleichera oleosa Merr., Serjania Mill. sp., Shorea Roxb. ex C. F. Gaertn
sp.. S. leprosula Mil., Spondias mombin L., Swietenia macrophylla King, S. mahagoni
(L.) J acq., Tabebuia rosea DC ., Tarenna incerta Koord. and Valeton, Tectona grandis L.,
Tephrosia Pers. sp., T. maxima Pers., T. vogelii Hook, Thea sinensis L., Theobroma
cacao L., Terminalia amazonica (J. F. Gmel.) Exell, Trema micrantha (L.) Blume, T.
orientale Blume, Vanda Jones ex. R.Br. sp., V. coerulea Griff. ex Lindl., V. teres Lindl.,
V. tricolor Lindl., Vitis L. sp.

Specimens Examined. (197 Q; 13 6)

Type material: Lectotype Xyleborus difficilis (Q; USNM). Allotype X ylosandrus
difiicilis: Java, Bnadya, VII-33, L. G. E. Kalshoven (6; NHMW). Cotype Xyleborus
difficilis: Java: Coll. Hagedom, 1915 (Q; NHMW). Lectotype Xyleborus luzonicus: Mt.
Makiling, Luzon, Baker, Eggers collection 1948 (Q; USNM). Paratype Xyleborus
apruptoides: Fiji: Viti Levu, Navai Mill. Nr. Nandarivatu, 2500 ft., 15.IX.1938, beating
shrubbery (Q; NHMW).

Other material: AUSTRALIAN REGION: Australia: In Phalaenopsis sp. VI-l7-35
(17 Q, l 6; USNM); Australia, on Vanda coerulea, VI-25-1938 (3 Q; USNM); Australia:

In Vanda coerulea, VII-7-36 (9 Q, 3 6 ; USNM); Australia, F ullaway, Nov. 26 1934,

157

Dendrobium phalaenopsis (1 Q; USNM); Australia: Brisbane, S. F., 3/1/49. In Quar.,
Dendrobium phalenopsis host, collector Art Retan (4 Q, 1 6; CAS). Soloman Islands:
Solomon Islands: Guadalcanel, Mt. Auston. 11 Jan 1984, M. Bigger, boring in Pometia
pinnata midrib (1 Q; BMNH). NEARCTIC REGION: United States: New Jersey,
Bound Brook, Aug 14/16, B. Weiss Harry colr., Cattleya (6 Q, 3 6; USNM).
NEOTROPICAL REGION: Colombia: Colombia: 24 km E Barbosa, VII-18-70.
Antioquia, e1. 1200 m, S. L. Wood, Lauraceae (1 Q; USNM); Colombia: 24 km E
Barbosa, VII-18-70, Antioquia, el. 1200 m, S. L. Wood, Xelopia sp. (2 Q; USNM);
Colombia: 8 km S Colonia V. dc Cauca, VII-9-70, e1. 30 m, S. L. Wood, Aspidosperma
sp. (5 Q; USNM); Colombia: 8 km S Colonia V. de Cauca, VII-9-70, e1. 30 m, S. L.
Wood, Lecythis sp. (1 Q; USNM); Colombia: 8 km S Colonia V. de Cauca, VII-9-70, e1.
30 m, S. L. Wood, Chrysophyllum catmito (1 Q; USNM). Costa Rica: Costa Rica:
Pandora lim., 150 ft., VIII-23-l963, S. L. Wood, Cecropia sp. leaf petioles (1 Q:
USNM); Costa Rica: San Jose, Santa Ana, 4000 ft., X 4—1963, S. L. Wood, unknown
limb (1 Q; USNM). Ecuador: Ecuador: Galap: St. Cruz, 1.7 km N St, Rosa, l-30.V.91.
550 In, Scalesia, mal-F IT, S. and J. Peck (2 Q; CAS); Ecuador, Napo Prov., Rees.
Ethnica Waorani, 1 km S. Onkone Gare Camp, Trans. Ent. 00°39’10”S, 076°26’W, 220
m. elev., July 1995, T. L. Erwin, et a1 collectors (1 Q; USNM); Ecuador, Napo Prov.,
Rees. Ethnica Waorani, 1 km S. Onkone Gare Camp, Trans. Ent. 00°39’10”S, 076°26’W.
220 m. elev., January 1996, T. L. Erwin, et al collectors (1 Q; USNM); Ecuador, Napo
Prov., Rees. Ethnica Waorani, 1 km S. Onkone Gare Camp, Trans. Ent. 00°39’10”S.
076°26’W, 220 m. elev., July 1996, T. L. Erwin, et al collectors (3 Q; USNM); Ecuador,

Napo Prov., Tiputini Biodiversity Station, 220-250 m. October 1998, 00°37‘55”S.

158

076°08’39”W, T. L. Erwin, et al collectors (3 Q; USNM). Honduras: Honduras, at
Tampa, 28-III-1968, J. Jordan, in orchid plnt. stem (1 Q; USNM); Honduras: Zamorano,
Moraz, 2200 ft., IV-18-1946, S. L. Wood, Serjania (2 Q; USNM). Mexico: Mexico:
OAX, Romero 18 mi N, 400 ft, VI-29-l967, S. L. Wood, unknown twig (1 Q; USNM);
Mexico: OAX, Romero 18 mi N, 400 ft, VI—29-1967, S. L. Wood, unknown branch (1 Q:
USNM); Mexico: OAX, Romero 23 mi N, VI-29-l967, 300 ft., S. L. Wood, unknown
branch (1 Q; USNM); Mexico: VC, Coatzocoalcos, 18 mi E, VI-26-l967, e1. 10011., S. L.
Wood, unknown log (2 Q; USNM); Mexico, Vera Cruz, and Oaxaca, 24-VIII-66, Cool.
G. Schwenke, Orchids and bromeliads (8 Q; USNM). Nicaragua: F inca San Pedro,
Matagalpa, 10-IV-81, en tallo de Cacao, Coll. J. A. Estrada (4 Q; USNM). Panama:
Ancon, C. 1., l8.VII-45, Epidendrum stamfordianum Broelle (1 Q; USNM), Barro
Colorado, CZ Panama, 16, 19, 21/5 1986, Henk Wolda (2 Q; UCDC); Barro Colorado.
CZ Panama, 25, 28, 30. IV.1986, Henk Wolda (2 Q; UCDC); Barro Colorado. CZ
Panama, 22, 24, 26. IX. 1986, Henk Wolda (6 Q; UCDC); Barro Colorado, CZ Panama,
28, 31/3. 2/4. 1986, Henk Wolda (2 Q; UCDC); Barro Colorado, CZ Panama, 20-
24.X.1986, Henk Wolda (4 Q; UCDC); Barro Colorado, CZ Panama, 25-29.V.1987,
Henk Wolda (2 Q; UCDC); Barro Colorado, CZ Panama, l-5.VI.1987, Henk Wolda(1
Q; UCDC); Barro Colorado, CZ Panama. l-5.XII.1986, Henk Wolda (1 Q; UCDC);
Barro Colorado, CZ Panama, 23-27.II.1987, Henk Wolda (1 Q; UCDC); Barro Colorado.
CZ Panama, 15, 17, 19. IX.1986, Henk Wolda (3 Q; UCDC); Barro Colorado, CZ
Panama, 4, 8, 9. IV. 1986, Henk Wolda (2 Q; UCDC); Barro Colorado, CZ Panama, 14,
17, 19, .III. 1986, Henk Wolda (1 Q; UCDC); Barro Colorado, CZ Panama, 9, 12, 14. V.

1986, Henk Wolda (2 Q; UCDC); Barro Colorado, CZ Panama. 1, 3. 5.IX.1986, Henk

159

Wolda (13 Q; UCDC); Barro Colorado, CZ Panama, 25, 27. VIII. 1986, Henk Wolda (1
Q; UCDC); Barro Colorado, CZ Panama, 18, 21, 23. IV. 1986, Henk Wolda (2 Q;
UCDC); Barro Colorado, CZ Panama, 29.1X/1,3.X.1986, Henk Wolda (6 Q; UCDC);
Barro Colorado, CZ Panama, 8-12.XII.1986, Henk Wolda (1 Q; UCDC); Barro Colorado.
CZ Panama, 11, 14, 16/4.1986, Henk Wolda (3 Q; UCDC); Barro Colorado, CZ Panama.
2, 5, 7/5. 1986, Henk Wolda (2 Q; UCDC); Barro Colorado, CZ Panama, 31-1/3-II.1986,
Henk Wolda (1 Q; UCDC); Barro Colorado, CZ Panama, 6, 9, ll.VI.l986, Henk Wolda
(1 Q; UCDC); Barro Colorado, CZ Panama, 7, 9, 11.VII.1986, Henk Wolda (1 Q;
UCDC); Barro Colorado, CZ Panama, 18, 20, 22.VIII. 1986, Henk Wolda (1 Q; UCDC):
Barro Colorado, CZ Panama, 13, 15, l7.X.1986, Henk Wolda (1 Q; UCDC); Barro
Colorado, CZ Panama, 21, 23, 25.VII.1986, Henk Wolda (1 Q; UCDC); Panama: Barro
Colo Is. OZ, I-II-45, J. Zetek (3 Q; USNM); Baro C010. 15., CZ, Mar. Apr. 49, Zetek (1
Q; USNM); Barro Colorado 151., CZ, XII-46-II-47 J Zetek collector (1 Q; USNM):
Panama: Barro Colorado Island, CZ, VII-23-1966, S. L. Wood, unknown log (1 Q;
USNM); CZ Panama, Lion Hill, VI 21 1982, R. B. Kimsey Col. (1 Q; UCDC); Panama:
CZ, Ft Amador, VlI-27-66, S. L. Wood, unknown twigs (4 Q; USNM); Panama: CZ.
Gatun Darn, 40 ft., XII-31-1963, S. L. Wood, Cecropia sp. leaf petioles (2 Q; USNM);
Panama: Panama Prov., 6-8 km N El Llano on El Llano-Carti Road, VI-6-1994, F.
Andrews and A. Gilbert (1 Q; CSCA); Panama: Panama Prov. 9 km N E] Llano, V-21-
1993, F. Andrews and A. Gilbert (1 Q; CSCA). Venezuela: Venezuela 9 km S of
Barrancas, Barinas, 150 m, XI-5-69, S. L.‘Wood, Spondias mombin (2 Q; USNM);
Venezuela: 9 km S of Barrancas, Barinas, 150 m, XI-5-69, S. L. Wood, Inga (1 Q;

USNM); Venezuela: 8 km SW Bumbum, Barinas, II-11-1970, 150 m. S. L. Wood.

160

Cucurbitaceae (1 Q; USNM); Venezuela: 20 km SE El Vigia Merida, XII-10 69, e1. 50 m.
S. L. Wood, unknown vine (1 Q; USNM); Venezuela: F inca Monasterios, Cacaugua,
Mir., 1971, Theobroma cacao (1 Q; USNM). Sanat Domingo, 25-1-1980, Coffee. J.
Esenbar (1 Q; USNM); Venezuela: 40km SE Socopo, Barinas, I-25-1970, 150 m, S. L.
Wood, Palito negro (1 Q; USNM); Venezuela: 40 km SE Socopo, Barinas, I-25-1970,
150 m, S. L. Wood, Laurel roja(1 Q; USNM). OCEANIA: Fiji Islands: Fiji, Viti Levu,
Nadarivatu, VIII.1955, B. A. O’Connor, young mahogany (2 Q; NHMW). Samoan
Islands: Samoa: Afiamalu, Upolu, VII-10-40, beating dead branches, F. C. Zimmerman
collector (1 Q; NHMW). ORIENTAL REGION: Indonesia: Java, Bnadya, VII-33, L.
G. E. Kalshoven (2 Q; NHMW). India: Coffee Res Sub-station, Chethalli, Kamataka.
Sp. 70. on Coffea robusta (1 Q; BMNH); Java: W Bandjar, 1933, leg. Kalshoven,
Tectona (2 Q; USNM). La Reunion, Saint Pierre, l7.X.1989, Orchidee, S. Quilici (3 Q;
USNM). Malaysia: Malaya: Kelantan, I:VII:1947, F. G. Browne (3 6; BMNH).
Philipine Islands: Philipines, X-23-63, E. Shiroma and E. Davidson, in orchid sp. (1 Q:
USNM); Philipine 15., Apr. 2, 1940, In Ranathera storiei (1 Q; USNM); Philipine Is.,
Dendrobium superbum, E. Arbios, Sept.19.l933 (1 Q; USNM). Sri Lanka: Ceylon [Sri
Lanka]: Peradeniya, 29.VII.1914, A. Rutherford (1 Q; NHMW); Sri Lanka: Kal. Dist..
Morapitiya, 250 mtrs., 27 May 1975, S. L. Wood (1 Q; USNM). PALEARCTIC
REGION: England: England, St. Albans, Dendrobium phalenopsis, IV-25-30, E.

Rannells, intercepted Washington DC. (3 Q, 4 6; USNM).

Xylosandrus nanus (Blandford), nomen dubium

(Fig. 2.36)

161

Xyleborus nanus Blandford, 1896b: 242. Holotype Q: Noumea (Delauney); Location of
holotyope unknown.

X ylosandrus nanus (Blandford): Browne, 1963: 55.

Notes. Xylosandrus nanus was described from a single specimen by Blandford in 1896.
When Browne (1963) transferred the species to Xylosandrus, he noted that it was
“probably at most a variation of X morigerus.” Furthermore, Wood and Bright (1992)
incorrectly cite the holotype as being housed in the BMNH. Since Blandford did not
indicate where the holotype was deposited in his original description, the location of the
holotype is unknown. Therefore, the authors were unable to examine any specimens of
X nanus. Blandford separted it from X morigerus by the characters of minute granules
and setae on the elytral declivity. In several hundred specimens examined from New
Caledonia, not a single one corresponded to X nanus (Beaver per comm). Given that
after over 100 years, X nanus is still only known from the holotype, the validity of this
species is doubtful.

Distribution. OCEANIA: New Caledonia.

Hosts. Unknown.

Specimens Examined. Unable to examine any specimens (see notes above).

Xylosandrus pusillus (Schedl)
(Fig. 2.36)
Xyleborus pusillus Schedl, 1961a: 91. Holotype Q: Luzon, Rizal, Mt. Irid; NHMW.

Xylosandrus pusillus (Schedl): Schedl. 1964: 213.

162

Diagnosis. Female 1.5 — 1.7 mm long; 1.9 times longer than wide. Body light brown;
antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum 0.7 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense
patch of short, erect setae, indicating the presence of a pronotal-mesonotal mycangium.
Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type 0, Hulcr et al.
2007). Pronotum with a lateral costa and carina. Procoxae widely separated. Protibiae
with 4 socketed teeth on lateral margin; mesotibial teeth not visible on specimen
examined; metatibiae with 10 socketed teeth. Elytra 1.2 times longer than wide; 1.4
times longer than pronotum. Discal striae punctate; interstriae uniseriate punctate.
Elytral disc gradually curving into declivity. Declivity convex, lateral margins carinate to
7th interstriae. Six striae visible on declivity. Striae punctate, with semi-appressed, hair-
like setae, shorter than the width of second declivital interstriae. Interstriae granulate,
uniseriate, with erect, hair-like setae, longer than twice the width of second declivital
interstriae.

This species is morphologically similar to X compactus (Fig. 2.14) and X
curtulus (Fig. 2.17). It can be distinguished from X curtulus by a pronotal disc that is
evenly pubescent, rather than mostly glabrous. X ylosandrus pusillus is nearly
morphologically identical to X compactus. The only character distinguishing the two

species is the degree of body stoutness, with X pusillus being 1.9 times as long as wide

163

and X compactus being 2.3 times as long as wide. However, this is too large a disparity
to warrant synonymizing the two species without further investigation.

Distribution. ORIENTAL REGION: Philippine Islands (Luzon).

Hosts. Unknown.

Specimens Examined. (1 Q; 0 6)

Type material: Holotype X yleborus pusillus (Q; NHMW).

X ylosandrus pygmaeus (Eggers)
(Fig. 2.3 8)

Xyleborus pygmaeus Eggers, 1940: 142. Holotype Q: Ost-Java (Alas Tbedek) leg.
Bedemann; NHMW.
Xylosandrus pygmaeus (Eggers): Browne, 1963: 55.
Diagnosis. Female 1.3 — 1.4 mm long; 2.3 times longer than wide. Body light brown to
brown; antennae and legs yellowish brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa;
segment one covering entire posterior face. Pronotum of equal length and width. Dorsal
aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-
appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presence of a pronotal-
mesonotal mycangium. Pronotal disc moderately punctate basally. Lateral aspect of
pronotum basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa and carina.
Procoxae widely separated. Protibiae with 4 socketed teeth on lateral margin; mesotibiae

with 6 socketed teeth; metatibiae with 5 socketed teeth. Elytra times 1.2 longer than

164

wide; 1.4 times longer than pronotum. Discal striae punctate; interstriae uniseriate
punctate. Declivital face steep and abruptly separated from disc. Declivity flattened.
lateral margins carinate to 7th interstriae. Five striae visible on declivity. Striae punctate,
without setae. Interstriae granulate, uniseriate, with erect, hair-like setae, longer than the
width of second declivital interstriae.

This species is morphologically similar to X boreensis (Fig. 2.12). X ylosandrus
pygmaeus can be distinguished from X boreensis by the following characters: elyral
declivity with striae less impressed; five striae visible on declivity; declivital striae with
erect or semi-erect, hair-like setae, longer than the width of the second declivital
interstriae; and pronotum with a lateral costa, but not carinate.

Distribution. ORIENTAL REGION: Indonesia (Java), Malaysia, Sri Lanka.

Hosts. Litsea amara Blume, Vitex pubescens Vahl.

Specimens Examined. (11 Q; 0 6)

Type material: Holotype Xyleborus pygmaeus (1 Q; NHMW).

Other material: ORIENTAL REGION: Indonesia: E. Java: Bangelan, 14.5.1932 (1 Q:
USNM); INDONESIA: SULAWESI UTARA, Dumoga-Bone N.P., Plot B, ca 300 m
Lowland Forest, Mar-85, Flight intercept trap (1 Q; RAB). Malaysia: BRUNEI:
Temburong: Nr. K. Belalong Field, Study Centre 150 m, 4°33'N 155°09'E, 21 .ii.1992, R.
A. Beaver, RGS/UBD Exped. (I Q; RAB); MALAYSIA: Sabah Sipitang, Mendolong.
T6/R, l4.iii.l989, leg. S. Abdebratt (1 Q; RAB); MALAYSIA: Sabah Sipitang,
Mendolong, T5/R, 10.iii.1989, leg. S. Abdebratt (1 Q; RAB); MALAYSIA: Sabah
Sipitang, Mendolong, T6/R, l4.iii.l989, leg. S. Abdebratt (1 Q; RAB); MALAYSIA:

Sabah Sipitang, Mendolong. T6/R, 11.v.1988, leg. S. Abdebratt (1 Q; RAB);

165

MALAYSIA: Sabah Sipitang: Mendolong, T5/R, 28.iv.1988, leg. S. Abdebratt, comps.
Sp'm det. F. G. Browne (1 Q; RAB). Sri Lanka: Ceylon [Sri Lanka]: W. Prov.,
Labugama, 24 miles ESE Colombo, 21.1.62, in sweep net (1 Q; NHMW). OTHER
(uncertain): Selega (?), Kefang (?), 25. 1. 1949, F. G. Browne, ex Vitex pubescens (1 Q:

BMNH).

Xylosandrus queenslandi Dole and Beaver
(Fig. 2.3 8)

Xylosandrus queenslandi Dole and Beaver, in press. Holotype Q: AUSTRALIA,
Queensland, Bunya Mountain NP, 1100m, ex Leguminosae tree, 19.i.2000 (B. Jordal and
A. Sequeira); QMB.
Diagnosis. Female 1.6 — 1.9 mm long; 2.2 times longer than wide. Body light brown to
brown; elytra slightly darker than pronotum; antennae and appendages light brown.
F rons retilculate and sparsely punctate. Antennae with 5 funicular segments. Antennal
club obliquely truncate; first segment forming a circular costa; segment one covering
entire posterior face. Pronotum 0.9 times longer than wide. Dorsal aspect of pronotum
rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi—appressed, hair-like setae;
setae less dense on disc. Basal pronotum with a dense patch of short, erect setae,
indicating the presense of a pronotal-mesonotal mycangium. Pronotal disc moderately
punctate. Lateral aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotum with
lateral costa, not carinate. Procoxae widely separated. Protibiae with 4 socketed teeth on
lateral margin, meso- and metatibiae with 7 —— 9 socketed teeth. Elytra 1.2 times longer

than wide; 1.4 times longer than pronotum. Discal striae punctate; interstraie uniseriate

166

punctate. Declivtal face of elytra steep and abruptly separated from disc. Declivity
flattened, lateral margin marked by a row of coarse, closely placed serrations. Six striae
visible on declivity. Striae coarsely granulate, with erect, tapered hair-like setae, shorter
than the width of second declivital interstriae. Interstriae granulate, uniseriate, with erect.
hair-like setae, longer than the width of second declivital interstriae.

This species is one of three Xylosandrus with lateral declivital marigins that are
marked by coarse serrations: X abruptulus (Fig. 2.6), X corthyloides (Fig. 2.15), and X
queenslandi. Xylosandrus queenslandi can be distinguished from X corthyloides by the
presense of a dense patch of mycangial setae on the basal pronotum. X ylosandrus
queenslandi can be distinguished from X abruptulus by the following characters:
declivital face steep and abruptly separated from disc; declivity matte; and declivital
striae with erect, acutely tapering, hair-like setae, shorter than the width of the second
declivital interstriae.

Distribution. AUSTRALIAN REGION: Queensland.

Hosts. Argyrodendron actinophyllum (Bailey) Edlin.

Specimens Examined. (2 Q; 0 6)

Type material: Paratypes X ylosandrus queenslandi: Australia: Queensland, Bunya
mountains National Park, Jan. 2000, 1100 m, ex. Leguminosae tree, 19.1 B. Jordal and A.

Sequeira leg. (2 Q; MSU).

Xylosandrus rotundicollis (Browne), new combination

(Fig. 2.39)

167

Xyleborus rotundicollis Browne, 1984: 73. Holotype Q: New Guinea: Morobe District,
Mount Kaindi, 2350 m, 4.XI.1972; BMNH.
Diagnosis. Female 3.7 — 4.1 mm long; 2.2 — 2.6 times longer than wide. Body dark
brown; antennae and legs light brown. Frons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa:
segment one covering entire posterior face. Pronotum 0.8 — 1.0 times longer than wide.
Dorsal aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; setae less dense on disc. Basal pronotum with a dense
patch of short, erect setae, indicating the presence of a pronotal-mesonotal mycangium.
Pronotal disc moderately punctate. Lateral aspect of pronotum basic (type 0. Hulcr et al.
2007). Pronotum with lateral costa, not carinate. Procoxae narrowly, but completely
separated. Protibiae with 4 — 5 socketed teeth on lateral margin; mesotibiae with 10
socketed teeth; metatibiae with 11 socketed teeth. Elytra 1.2 — 1.7 times longer than
wide; 1.2 — 1.8 times longer than pronotum. Discal striae punctate; interstriae
multiseriate punctate. Elytral disc gradually curving into declivity. Declivity convex.
lateral margins rounded, without a carina or a row of tubercles or serrations. Six striae
visible on declivity. Striae punctate, with semi-appresssed, hair-like setae, shorter than
the width of second declivital interstriae. Interstriae granulate, uniseriate, with erect.
hair-like setae, longer than twice the width of second declivital interstriae.

This species is morphologically similar to X monteithi (Fig. 2.33). Xylosandrus
rotundicollis can be distinguished from X monteithi by the following characters: 3.7 —
4.1 mm long; basal pronotum with a dense patch of short, erect setae, indicating the

presence of a pronotal-mesonotal mycangium; declivital striae with semi-appressed. hair-

168

like setae, shorter than the width of the second declivital interstriae; and declivital
interstriae uniseriate granulate, with erect, hair-like setae, longer than twice the width of
the second declivital interstriae.

Distribution. AUSTRALIAN REGION: Papua New Guinea.

Hosts. Ficus L. sp., F. mollior Benth., Gordonia Ellis sp., Meliosma Blume sp.,
Schefllera sp. J. R. Forst and G. F orst.

Specimens Examined. (56 Q; 0 6)

Type material: Holotype Xyleborus rotundicollis (Q; BMN H).

Other material: AUSTRALIAN REGION: Papua New Guinea: Papua New Guinea.
Mu village, March 2006, 1600 m asl, Hulcr and Cognato coll. (54 Q; MSU). Papua New

Guinea, Kupa Range, Morobe, 2000 m., glue trap, Roberts coll. (1 Q; FICB).

Xylosandrus subsimiliformis (Eggers)
(Fig. 2.41)

Xyleborus subsimiliformis Eggers, 1939a: 11. Holotype Q: Nordostbirma (Kaim, 7000
Fuss) 17.V.l934; NHR (Eggers Cotype in NHMW).
Xylosandrus subsimiliformis (Eggers): Wood and Bright, 1992: 800.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors failed to indicate it as a “new combination.”
Diagnosis. Female 2.8 mm long; 2.1 times longer than wide. Body brown; antennae and
legs same color as body. Frons punctate. Antennae with 5 funicular segments. Antennal

club obliquely truncate; first segment forming a circular costa; segment one covering

169

entire posterior face. Pronotum 0.9 times longer than wide. Dorsal aspect of pronotum
rounded (type 1, Hulcr et a1. 2007). Pronotal vestiture or semi-appressed, hair-like setae;
setae less dense on disc. Basal pronotum with a dense patch of short, erect setae.
indicating the presence of a pronotal-mesonotal mycangium. Pronotal disc densely
aspirate-granulate, with sculpture separated by distance less than or equal to their width.
Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa.
not carinate. Procoxae widely separated. Protibiae with 4 socketed teeth on lateral
margin; mesotibiae with 10 socketed teeth; metatibiae with 10 socketed teeth. Elytra 1.2
times longer than wide; 1.3 times longer than pronotum. Discal striae punctate;
interstriae multiseriate punctate. Declivital face of elytra steep and abruptly separated
from disc. Declivity flattened, lateral margins carinate to 7th interstriae. Four striae
visible on declivity. Striae granulate, with appressed, hair-like setae, shorter than the
width of second declivital interstriae. Interstriae finely granulate, multiseriate, with
appressed, hair-like setae, shorter than the width of second declivital interstriae.

This species is morphologically similar to X beesoni (Illustrated in Saha et al.
1992), X discolor (Fig. 2.20), and X jaintianus (Fig.2.27). X subsimiliformis can be
distinguished from these species by the following characters: declivity flattened;
pronotum with a conspicuous summit on basal third; four striae visible on elytral
declivity; and frons punctate, without a distinct median keel.
Distribution. ORIENTAL REGION: Bumia.
Hosts. Unknown.

Specimens Examined. (I Q; 0 6)

170

Type material: Cotype Xyleborus subsimiliformis: N. E. Burma, Nambaiti. 7000 ft.,

12.5.1924, R. Malaise (Q; NHMW).

X ylosandrus subsimilis (Eggers)
(Fig. 2.41)

Xyleborus subsimilis Eggers, 1930: 186. Holotype Q: Assam (Shillong, 6000 ft): FRI.
Xylosandrus subsimilis (Eggers): Wood and Bright, 1992: 800.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992). but
the authors failed to indicate it as a “new combination.”
Diagnosis. Female 2.6 mm long; 2.0 times longer than wide. Body brown; antennae and
legs same color as body. F rons rugose. Antennae with 5 funicular segments. Antennal
club obliquely truncate; first segment forming a circular costa; segment one covering
entire posterior face. Pronotum 0.9 times longer than wide. Dorsal aspect of pronotum
rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi- appressed, hair-like setae;
setae less dense on disc. Basal pronotum with a dense patch of short, erect setae,
indicating the presence of a pronotal-mesonotal mycangium. Pronotal disc densely
asperate-granulate, with sculpture separated by distance less than or equal to their width.
Lateral aspect of pronotum basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa
and carina. Procoxae widely separated. Protibiae with 4 - 5 socketed teeth on lateral
margin; mesotibiae with 9 — 11 socketed teeth; metatibiae with 10 socketed teeth. Elytra
1.1 times longer than wide; 1.2 times longer than pronotum. Discal striae punctate;
interstraie multiseriate punctate. Declivital face steep and abruptly separated from disc.

Declivity flattened, lateral margins carinate to 7‘h interstriae. Four striae visible on

171

declivity. Striae coarsely granulate, with appressed, scale-like setae, shorter than the
width of second declivital interstriae. Interstriae granulate, multiseriate, with appressed.
scale-like setae shorter than the width of second declivital interstriae.
This species is morphologically similar to X beesoni (Illustrated in Saha et al.
1992), X discolor (Fig. 2.20), X jaintianus (Fig. 2.27), and X subsimiliformis (Fig.
2.41). However, X subsimilis can easily be distinguished from these species by its
declivital vestiture of dense, flattened, scale-like setae.
Distribution. ORIENTAL REGION: China, India (Assam, Bengal), Thailand.
Hosts. Cinnamomum obtusifolium Nees., Tectona grandis L., Terminalia myriocarpa
Van Heurck and Mfill. Arq..
Specimens Examined. (8 Q; 1 6)
Type material: Cotype Xyleborus subsimilis: Shillong, 6000 it, C. F. C. Beeson.
14.V.l925, ex. unknown wood (1 Q; NHMW).
Other material: ORIENTAL REGION: China: China, 1962.VII (2 Q; USNM). India:
Samsingh 1800, Kalimpong, Bengal, 4.X.33, C. F. C. Beeson. ex. Cinnamomum
obtusifolium branches (2 Q; USNM). Thailand: Thailand: Chiang Mai: Doi Pui, 16.1.05.
R. A. Beaver (1 Q; RAB); Thailand: Chiang Mai. Doi Suthep, c.1400m, 18.x.04, R. A.
Beaver (2 Q. l 6; RAB).
X ylosandrus terminatis (Eggers)
(Fig. 2.42)
Xyleborus terminatis Eggers, 1930: 182. Holotype Q: Coorg (Virojapet, Sidapur); FRI.

Xylosandrus terminatis (Eggers): Browne, 1963: 55.

Diagnosis. Female 1.5 — 1.9 mm long; 2.1 times longer than wide. Body dark brown to
black; antennae and legs yellowish brown. F rons punctate. Antennae with 5 funicular
segments. Antennal club obliquely truncate; first segment forming a circular costa:
segment one covering entire posterior face. Pronotum 0.8 times longer than wide.

Dorsal aspect of pronotum rounded (type 1, Hulcr et a1. 2007). Pronotal vestiture of
semi-appressed, hair-like setae; pronotal disc glabrous, except for mycangial setae. Basal
pronotum with a dense patch of short, erect setae, indicating the presence of a pomotal-
mesonotal mycangium. Pronotal disc moderately punctate. Lateral aspect of pronotum
basic (type 0, Hulcr et al. 2007). Pronotum with lateral costa and carina. Procoxae
widely separated. Protibiae with 4 socketed teeth on lateral margin; meso- and metatibial
teeth not visible on specimens examined. Elytra 1.2 times longer than wide; 1.4 times
longer than pronotum. Discal striae punctate; interstriae uniseriate punctate. Declivital
face of elytra steep and abruptly separated from disc. Declivity flattened, lateral margins
carinate to 7th interstriae. Striae punctate, without setae. Interstriae punctate, uniseriate.
with erect, hair-like setae, longer than the width of second declivital interstriae.

This species is morphologically similar to X derupteterminatus (Fig. 2.18) and X
morigerus (Fig. 2.35). Xylosandrus terminatus can be distinguished from X morigerus
by the following characters: elytral declivity flattened; and fiver striae visible on
declivity. Xylosandrus terminatus can be distinguished from X derupteterminatus by the
following characters: smaller species, 1.5 - 1.9 mm long; declivital striae without setae;
and declivital interstriae uniseriate punctate.

Distribution. ORIENTAL REGION: India (Kamataka. Maharashtra, Tamil Nadu).

Hosts. Holigarna arnottiana Hook, Lantana L. sp., Swietenia macrophylla King.

173

Specimens Examined. (3 Q; 0 6)

Type material: Cotype Xyleborus terminatis: Coorg, Virojapet, Sidapur, Y. R. Rao coll..
9. XI.1917, boring into twigs of Lantana (Q; NHMW).

Other material: ORIENTAL REGION: Inida: Chandanthode, Wynadd, Madras.

Research Forester., 8.XII.1938, ex Swietenia macrophylla stump (2 Q; USNM).

X ylosandrus woodi Dole and Beaver
(Fig.2.42)

Xylosandrus woodi Dole and Beaver, in press. Holotype Q: [AUSTRALIA],
NEQ[ueensland], 16°30'S x 145°19'E, Mt Demi summit, 1100 m, flight intercept,
17.xii.1995-22.i.l996 (Monteith, Thompson and Ford). In QMB (Accession # T144404).
Diagnosis. Female 2.3 — 2.4 mm long; 2.1 times longer than wide. Body dark brown;
antennae and legs light brown. F rons punctate. Antennae with 5 funicular segments.
Antennal club obliquely truncate; first segment forming a circular costa; segment one
covering entire posterior face. Pronotum 0.8 times longer than wide. Dorsal aspect of
pronotum rounded (type 1, Hulcr et al. 2007). Pronotal vestiture of semi-appressed hair—
like setae; pronotal disc densely setose, with setae as dense as on anterior pronotum.
Lacking a dense patch of setae at base of pronotum. Pronotal disc densely asperate-
granulate. Lateral aspect of pronotum rounded (type 1, Hulcr et al. 2007). Pronotum
with lateral costa, not carinate. Procoxae widely separated. Protibiae with 4-5 socketed
teeth on lateral margin; meso- and metatibiae with 7-8 socketed teeth. Elytra 1.3 times
longer than wide; 1.6 times longer than pronotum. Discal striae punctate; interstriae

multiseriate punctate. Elytral disc gradually curving into declivity. Declivity convex.

174

lateral margin with discontinuous row of small tubercles, some of these towards apex
with carinate tip. Five striae visible on declivity. Straie punctate, with appressed, hair-
like setae, longer than the width of second declivital interstriae. Interstriae granulate.
multiseriate. with appressed hair-like setae, longer than the width of second declivital
interstriae.

This species can be distinguished from all other known X ylosandrus by its lateral
declivital margin which is marked by a discontinuous row of small tubercles.
X ylosandrus woodi may be confused with X monteithi (Fig. 2.33) and X rotundicollis
(Fig. 2.39), but these two species have lateral declivital margins that are rounded and
without a row of tubercles.
Distribution. AUSTRALIAN REGION: Queensland.
Hosts. Unknown.
Specimens Examined. (1 Q; 0 6)
Type material: Paratype Xylosandrus woodi: Australia: NEQ, 19°07‘S, 146°23‘E, Mt
Halifax summit, 1050m, heath, rainforest, pitfalls, 19-21.iii.1991, G. Monteith and D.

Cook (1 Q; MSU).

New Combinations, Amasa
Amasa cylindrotomicus (Schedl), new combination
Pseudoxyleborus cylindrotomicus Schedl, 1939: 40. Lectotype Q: Sumatra, Benkoelen.
23-6-31, leg. Shuller; NHMW; designated by Schedl, 1979a: 74.

Xylosandrus cylindrotomicus (Schedl): Wood and Bright, 1992: 793.

175

Xyleborus semitruncatus Schedl, 1942b: 35. Lectotype Q: Sumatra, Manna, 15-lX-34:
NHMW; designated by Schedl, 1979a: 224. Synonymy: Wood, 1989: 177.

Xyleborus truncatellus Schedl, 1951: 79. Lectotype Q: Z. Sumatra, Poelau Pisang, and
Manna; NHMW; designated by Schedl 1979a: 256. Synonymy: Kalshoven, 1959: 95.
Xyleborusjucundus Schedl, 1954: 138. Lectotype Q: Z. Sumatra, Poelau Pisang, and
Manna; NHMW; designated by Schedl, 1979a: 256. Synonymy: Kalshoven, 1959: 95.
Xyleborus ramulorum Schedl, 1957: 115. Holotype Q: Congo Belge: Yangambi; MRC B.
Synonymy: Wood, 1989: 177.

Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.

Distribution. AF ROTROPICAL REGION: Zaire. ORIENTAL REGION: Indonesia
(Sumatra).

Specimens Examined. (4 Q; 1 6)

Type material: Lectotype Pseudoxyleborus cylindrotomicus (Q; NHMW). Lectotype

X yleborus semitruncatus (Q; NHMW). Lectotype Xyleborus truncatellus (Q; NHMW).
Allotype Xyleborus truncatellus: Sumatra, Manna, 25-IX-1934 (6; NHMW).

Other material: ORIENTAL REGION: Indonesia: Sumatra, Benkoelen. 23-6—31, leg.
Shuller (1 Q; NHMW).

Discussion. Based on the contiguous procoxae, antennal club with 3 segments visible on
the anterior and posterior face, the declivital face that is steep and abruptly separated
from the elytral disc and the circular costa forming a complete circumdeclivital ring, this

species is here transferred to A masa.

176

Amasa omissus (Schedl), new combination
X yleborus omissus Schedl, 1961b: 153. Holotype Q: Madagascar. Antaniditra pres
Perinet, 18 November 1952, K. E. Schedl; IRSM.
Xylosandrus omissus (Schedl): Wood and Bright, 1992: 799.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992). but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. AF ROTROPICAL REGION: Madagascar.
Specimens Examined. (2 Q; 0 6)
Type material: Paratypes Xyleborus omissus: Madagascar. Perinet, 17.XI.1952, K. E.
Schedl (2 Q; NHMW)
Discussion. Based on the contiguous procoxae, antennal club with 3 segments visible on
the anterior and posterior face, the declivital face that is steep and abruptly separated
from the elytral disc and the circular costa forming a complete circumdeclivital ring. this

species is here transferred to Amasa.

Amasa oralis (Schedl), new combination
Xyleborus oralis Schedl, 1961b: 154. Holotype Q: Madagascar, Antaniditra pres Perinet.
18 November 1952, K. E. Schedl; IRSM.

X ylosandrus oralis (Schedl): Wood and Bright, 1992: 799.

177

Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.

Distribution. AF ROTROPICAL REGION: Madagascar.

Specimens Examined. (1 Q; 0 6)

Type material: Paratype Xyleborus oralis: Madagascar, Perinet, 17.XI.1952, K. E.
Schedl (9; NHMW).

Discussion. Based on the contiguous procoxae, antennal club with 3 segments visible on
the anterior and posterior face, and the declivital face that is steep and abruptly separated

from the elytral disc, this species is here transferred to Amasa.

New Combinations, Anisandrus
Anisandrus butamali (Beeson), new combination

Xyleborus butamali Beeson, 1930: 40. Syntypes Q: Bombay: Agsur, South Kanara, and
Dandeli, North Kanara, B. M. Bhatia, Nov.; FRI.
Xylosandrus butamali (Beeson): Wood and Bright, 1992: 788.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. ORIENTAL REGION: India (Maharaashtra. Tamil Nadu).
Specimens Examined. (7 Q; 0 6)
Type material: Paratypes Xyleborus butamali: Bombay, Agsur, S. Kanara Div., l3.XI.29.

B. M. Bhatia, ex. Dillenia pentagyna (2 Q; NHMW).

178

Other material: ORIENTAL REGION: India: S. India: Anamalis Hills, C inohona,
3500 ft, April 1956, P. S. Nathan (1 Q; USNM); Kattiyur, Wynaad, Madras, F. R. J.
Project, 25.12.1945 (3 Q; USNM); Manantoddy. N. Malabar, G. C. Robinson. 8.XI.1930.
ex Tectona grandis (1 Q; USNM).

Discussion. Based on the contiguous procoxae, obliquely truncate antennal club with
first segment forming a circular costa and segment one covering the entire posterior face.
protibiae with 6 socketed teeth. and rounded lateral pronotal margins this species is here

transferred to Anisandrus.

Anisandrus ursa (Eggers), new combination
Xyleborus ursa Eggers, 1923: 172. Lectotype Q: Haveri, N. Guinea, S. E., Havari. Loria.
VII-XI-93; USNM; designated by Anderson and Anderson, 1971: 35.
X ylosandrus ursa (Eggers): Wood and Bright, 1992: 801.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. AUSTRALIAN REGION: New Guinea, Solomon Islands.ORIENTAL
REGION: Malaya, Indonesia (Celebes, Sumatra), Philipine Islands (Luzon).
Specimens Examined. (8 Q; 0 6)
Type material: Lectotype Xyleborus ursa (Q; USNM).
Other material: ORIENTAL REGION: Indonesia: INDONESIA. Irian Jaya: PT.,
Freeport Concession Wapoga, camp. 03.14°S 136.57°E, 3600 ft., 19-29 April 1998, R. R.

Snelling, Malaise trap montane primary rainforest (1 Q; RAB); INDONESIA:

179

SULAWESI UTARA, Dumoga-Bone NP, 1008 m G. mogongonipa summit, Aug-85,
Malaise Trap (1 Q; RAB); INDONESIA: SULAWESI UTARA, Dumoga-Bone N.P.
G.Mogogonipa summit, 1008 m., Sep-85, Pitfall trap (1 Q; RAB); INDONESIA:
SULAWESI UTARA, Dumoga-Bone N.P. G.Mogogonipa summit, 1008 m., May-85.
Malaise Trap (2 Q; RAB). Philipine Islands: Luzon, P. I. Baguio Mt. Sto, Tomas W.
Schultze (2 Q; NHMW).

Discussion. Based on the contiguous procoxae, obliquely truncate antennal club with
first segment forming a circular costa and segment one covering the entire posterior face,
protibiae with 7 socketed teeth, and rounded lateral pronotal margins this species is here

transferred to Anisandrus.

Anisandrus ursinus (Hagedom), new combination
Xyleborus ursinus Hagedom, 1908: 381. Holotype Q: Sumatra, Si-Ramve; MNB.
Xylosandrus ursinus (Hagedom): Wood and Bright, 1992: 801.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. ORIENTAL REGION: Indonesia (Boreo/Soemba Island. Sumatra),
Philipine Islands (Mindoro).
Specimens Examined. (9 Q; 0 6)
Type material: Unable to examine type material.
Other material: ORIENTAL REGION: Indonesia: Dammerman, N. O. Soemba,

Kambera, 111.1925 (1 Q; NHMW); Sumatra: Solok, Coll. F. Schneiber, 1911 (1 Q;

180

MTD). Philipine Islands: Philipine Islands: Glog Riv., Mt. Apo, Mindanao, IX.29,30.
Altitude 6000 ft., C011. by C. F. Clagg (2 Q; FMNH); Philipine Islands: La Lun Mts.
Davao Prov., Mindanao, 1.1.31, C011. by C. F. Clagg, at light (1 Q; FMNH); Mindoro.
Port Galera, Mc Gregor (2 Q; USNM); Mt. Makiling, Laguna P. 1., 7-21-22, F. C.
Hagedom Collection (1 Q; USNM); Philipine Islands: Seliban Riv., Mt. Apo, Mindanao.
VII.30.30, Altitude 6000 ft., Coll. by C. F. Clagg (1 Q; FMNH). I

Discussion. Based on the contiguous procoxae, obliquely truncate antennal club with
first segment forming a circular costa and segment one covering the entire posterior face,
protibiae with 7 socketed teeth, and rounded lateral pronotal margins this species is here

transferred to Anisandrus.

Anisandrus ursulus (Eggers), new combination
Xyleborus ursulus Eggers, 1923: 173. Holotype Q: Ost Bali, Kintamani, E. Sundainseln.
1913; MTD.
Xylosandrus ursulus (Eggers): Wood and Bright, 1992: 801.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. ORIENTAL REGION: China (F ujian), India (Bengal, Nicobar Islands),
Malaya, Thailand, Indonesia (Bali, Borneo, Java, Sumatra, Sunda Island), Philipine
Islands (Luzon).

Specimens Examined. (29 Q; 1 6)

181

Type material: Holotype Xyleborus ursulus (Q; MTD). Cotype Xyleborus ursulus: Java:
Batoerranden, G. Slamet, F. C. Drescher, 19.VII.1930 (6; NHMW).

Other material: ORIENTAL REGION: India: Samasingh, Kalimpong, Bengal.
111.1934, Mohal Lall. (3 Q; USNM); Kamsingh, 1800, Kalimpong, Bengal, 22.111.34, C.
F. C. Beeson, ex Sterculia colorata or Sapium eugeniaefolium (2 Q; USNM). Indonesia:
Java: Batoerraden, G. Slamet, F. C. Drescher, 22.111.1930 (5 Q; USNM); Java:
Batoerranden, G. Slamet, F. C. Drescher, 10.IV.1930 (2 Q; NHMW); Java: Mount Gede.
1900 m, XI-1932, L. G. E. Kalshoven (1 Q; USNM); INDONESIA: SULAWESI
UTARA, Dumoga-Bone N.P., Plot B, ca 300 m Lowland Forest, Apr-85. Flight intercept
trap (2 Q; RAB); INDONESIA: SULAWESI UTARA, Dumoga-Bone N.P.,
G.Mogogonipa summit, 1008 m, May-85, Malaise Trap (1 Q; RAB); INDONESIA:
SULAWESI UTARA, Dumoga-Bone N.P., Oct-85 (1 Q; RAB); INDONESIA:
SULAWESI UTARA, Dumoga-Bone N.P., Plot C, ca 400 m lowland forest, Apr-85,
Flight intercept trap (2 Q; RAB); INDONESIA: SULAWESI UTARA, Dumoga-Bone
N.P., Plot C, ca 400 m lowland forest. May-85, Flight intercept trap (1 Q; RAB).
Malaysia: MALAYA: Pahang: Palau, Tioman: Kampong Tekek, to Kampong J uara.
20.iii.1962, K. J. Kuncheria Collector BISHOP, In Jungle (1 Q; RAB); Mlay Penin:
Pabang. F. M. S., Fraseas [‘?] Hill, 4200 ft, 28-6-1931 (1 Q; NHMW); MALAYSIA:
Penang, Pencing Hill. 1701, 10.viii.77, R. A. Beaver, Ex. Pitfall trap (1 Q; RAB);
SABAH: Poring Spring, Xanthophyllum affine, Lower montane, 650 m, Mixed
dipterocarp Fst., 20. Vi. 1992, A. Floren, Fog Za4/F1 (1 Q; RAB). Thailand: Thailand:
Chiang Mai, Doi Chiang Dao, 12-13.vii.02, A. Cognato, ex ETOH trap on dead tree (1 Q;

RAB); Thailand: Kanchanaburi, 14.70N 98.87E, l7.vii.02, A. Cognato (3 Q; RAB).

182

Discussion. Based on the contiguous procoxae, obliquely truncate antennal club with
first segment forming a circular costa and segment one covering the entire posterior face.
protibiae with 7 socketed teeth, and rounded lateral pronotal margins this species is here

transferred to Anisandrus.

New Combinations, Cnestus

Cnestus ater (Eggers), new combination
Xyleborus ater Eggers, 1923: 210. Holotype Q: Batoe Insel (Tanah Masa); Natura Artis
Magistra.
Xylosandrus ater (Eggers): Wood and Bright, 1992: 787.
Xyleborus retusiformis Schedl, 1.936: 31. Holotype Q: Borneo; NHMW. Synonymy:
Wood, 1989: 177.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. ORIENTAL REGION: China (Fujian), Malaya, Indonesia (Batoe
Island, Borneo).
Specimens Examined. (6 Q; 0 6)
Type material: Holotype Xyleborus retusiformis (Q; NHMW).
Other material: ORIENTAL REGION: Brunei: BRUNEI: Temburong: Nr. K.
Belalong Field, Study Centre 300m, 4°33'N 155°09'E, 10.ii.1992, R. A. Beaver,
RGS/UBD Exped. Ex. Rattan Calamus Daeninorops (2 Q; RAB); BRUNEI: Temburong:

Nr. K. Belalong Field, Study Centre 300m, 4°33'N 155°09'E, 10.ii.1992, R. A. Beaver.

183

RGS/UBD Exped. Ex. Calamus Daeninorops sp. (2 Q; RAB). Indonesia: Padang. ex.
Ljengkeh [?], 14.Xii.35, Kalshoven (1 Q; NHMW).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most of the posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, the protibiae with 6 socketed teeth, and

the elytra that are wider than long, this species is here transferred to C nestus.

Cnestusfijianus (Schedl), new combination
Xyleborusfijianus Schedl, 1938: 50. Lectotype Q: Fiji Islands, Taveme Quilai, 800 feet,
October 18, 1924, H. S. Evans; NHMW.
Xylosandrusfijianus (Schedl): Wood and Bright, 1992: 794-795.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. OCEANIA: Fiji Islands.
Specimens Examined. (9 Q; 2 6)
Type material: Lectotype Xyleborusfijianus (Q; NHMW).
Other material: OCEANIA: Fiji: Fiji, Lami Quarry, Nr. Suva, VII-24-38, E. C.
Zimerman collector, beating shrubs, 10’, 250’ (1 Q; NHMW)I Fijji. Viti Levu, Suva,
22.xi.88, R. A. Beaver, ex Swietenia macrophylla (1 Q; RAB); FIJI, Viti Levu, Savura
Creek, 29.iii.83, R. A. Beaver, ex Serianthea melanesica (1 Q; RAB); FIJI, Viti Levu,

Namosi 10km, 19-20. V. 85, R. A. Beaver (2 Q; RAB); FIJI, Viti Levu, Savura C reek.

184

19.iii.83, R. A. Beaver, ex Canarium vitiense (2 Q; RAB)1; FIJI, Viti Levu, Savura
Creek, v.83, R. A. Beaver, ex Canarium vitiense (1 6; RAB); Fiji, Suva, ex. Swietenia
macrophylla (1 6; BMNH). Fiji, Colo-I-Suva, Static Trap 71, Mahongany plantation.
Cpt 23, 10/10/2006, K. Wotherspoon.

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle,
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most of the posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, the protibiae with 7 socketed teeth. and

the elytra that are wider than long, this species is here transferred to C nestus.

Cnestus gravidus (Blandford), new combination
Xyleborus gravidus Blandford, 1898: 427. Holotype Q: Chittagong Hills [Bangladesh];
BMNH.
X ylosandrus gravidus (Blandford): Wood and Bright, 1992: 796.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. ORIENTAL REGION: Bangladesh, Burma, India (Assam, Bengal),
Laos, Sri Lanka, Thailand, Vietnam (Tonkin). PALEARCTIC REGION: China
(Xizang [Tibet]).
Specimens Examined. (2 Q; 0 6)

Type material: Unable to examine type material.

185

Other material: ORIENTAL REGION: Vietnam: Nord Vietman, L. Thainguyen.
VII.1969, leg. Le. Van. Nong (1 Q; NHMW); Phovi, S 07 (1 Q; NHMW).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most of the posterior face, the presence of a coarse median pair of asperities
on the anterior margin of the pronotum, the protibiae with 7 socketed teeth. and the elytra

that are wider than long, this species is here transferred to Cnestus.

Cnestus improcerus (Sampson), new combination
Xyleborus improcerus Sampson, 1921: 33. Holotype Q: Siam; BMNH.
Xylosandrus improcerus (Sampson): Beaver, 1998: 183.
Distribution. ORIENTAL REGION: Brunei, India, Indonesia (Borneo). Malaya,
Thailand.
Specimens Examined. (7 Q; 0 6)
Type material: Unable to examine type material.
Other material: ORIENTAL REGION: Brunei. BRUNEI: Temburong: Nr. K.
Belalong Field, Study Centre 250 m, 4°33'N 155°09'E, 7.ii.1992, R. A. Beaver,
RGS/UBD Exped. (3 Q; RAB); BRUNEI: Temburong: Nr. K. Belalong Field, Study
Centre 250 m, 4°33'N 155°09'E, 21.ii.1992, R. A. Beaver, RGS/UBD Exped. (1 Q:
RAB); Lagleari [?] BC. Sarak, 20:X:l948, F. G. Browne (1 Q; USNM); Malaya,
Kelantan, E. G. B., 6.10.1946 (1 Q; USNM); MALAYA, Teneguann, Besut Dist.. V-

1958, R. Traub, 400 ft. (1 Q; USNM).

186

Discussion. Based on the 4-segmented antennal funicle, obliquely truncate antennal club
with first segment forming a circular costa and segment one covering the entire posterior
face, the presence of a coarse median pair of asperities on the produced anterior margin
of the pronotum, the protibiae with 6 socketed teeth, and the elytra that are wider than

long, this species is here transferred to Cnestus.

C nestus laticeps (Wood), new combination
Xyleborus laticeps Wood. 1977: 219. Holotype Q: 20 km SW El Vigia. Merida,
Venezuela; USNM.
Xylosandrus laticeps (Wood): Wood and Bright, 1992: 796.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. NEOTROPICAL REGION: Venezuela.
Specimens Examined. (51 Q; 12 6)
Type material: Xyleborus laticeps (Q; USNM).
Other material: NEOTROPICAL REGION: Venezuela: Venezuela: 9 km S
Barrancas, 150 m, X-l-69, S. L. Wood, Protium tenuifolium (13 Q; USNM); Venezuela:
9 km S of Barrancas, Barinas, 150 m, X-5-69, S. L. Wood (2 Q; USNM); Venezuela: 9
.km S. of Barrancas, Barinas, 150 m, XI-5-69, S. L. Wood, Spondias mombin (5 Q:
USNM); Venezuela: 9 km S. of Barrancas, Barinas, 150 m, Xl-5-69, S. L. Wood,
Dendropanax arboreum (1 Q; USNM); Venezuela: 9 km S Barrancas, Barinas, XII-2-69,

e1. 150 m, S. L. Wood, Melicoccous bijugata (1 Q; USNM); Venezuela: 8 km SW

187

Bumbum, Barinas, II-11-1970, 150 m, S. L. Wood, T artaguito sp. (2 Q; USNM);
Venezuela: 40 km, E Canton, Barinas, III-8-1970, 70 m, S. L. Wood, Protium tenui/olium
(1 Q; USNM); Venezuela: 40 km E Canton, Barinas, III-8—1970, 70 m, S. L. Wood.
Palito negro (1 Q; USNM); Venezuela: 40 km E Canton, Barinas, III-8-1970, 70 m, S. L.
Wood, tree seedling (1 Q; USNM); Venezuela: 40 km E Canton, Barinas, III-8-1970, 70
m, S. L. Wood, Pouteria anibaefolia (1 Q; USNM); Venezuela: 5 km W El Pino, 10 m,
Zulia, X-20-69, S. L. Wood, unknown tree (2 Q; USNM); Venezuela: 20 km SW El
Vigio, Merida, XI-21-69, e1. 50 m, S. L. Wood, Jacaranda sp. (10 Q; USNM);
Venezuela: Merida, 1700 m, IX-22-69, S. L. Wood (4 Q, 12 6; USNM); Venezuela: 40
km SE Socopo, Barinas, I-25-l970, 150 m, S. L. Wood, Protium sp. (1 Q; USNM);
Venezuela: 17 km SE of Miri, Barinas, XII-l7-69, 150 m, S. L. Wood, Protium sp. (1 Q;
USNM); Venezuela: 10 km SE of Miri, Barinas, II-8-1970, 150 m, S. L. Wood, Inga sp.
(1 Q; USNM); Venezuela: 7 km NW Socopo, Barinas, II-l3-1970, 200 m, S. L. Wood.
Protium sp. (1 Q; USNM); Venezuela: Valle de Choroni, IV-3-1964, J. L. Saunders,

T heobroma cacao (3 Q; USNM).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle,
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering the entire posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, the protibiae with 6 socketed teeth, and

the elytra that are wider than long, this species is here transferred to Cnestus.

Cnestus mutilatus (Blandford), new combination

Xyleborus mutilatus Blandford, 1894c: 103. Holotype Q: Japan; BMNH.

188

Xylosandrus mutilatus (Blandford): Wood and Bright, 1992: 799.

Xyleborus sampsoni Eggers, 1930: 184. Holotype Q: Assam (Haflong, Cachar); FRI.
Synonymy: Wood, 1989: 177.

Xyleborus banjoewangi Schedl, 1939: 41. Lectotype Q: Banjoewangi, 270 m.
Tjoerahlele, 25-11-36; NHMW. Synonymy: Kalshoven, 1960: 63.

Xyleborus taitonus Eggers 1939b: 118. Holotype Q: Formosa, Taito; Chujo Collection.
Synonmy: Wood and Bright, 1992: 799.

Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer. Xyleborus taitonus was also listed as a synonym by Wood and Bright (1992).
but was not indicated as a “new synonymy.” Wood and Bright (1992) indicate that
“Specimens in the FRI, Dehra Dun labeled by Eggers as taitonus are mutilatus;
synonymy needs confirmation”, but this author was unable to examine the specimens
referred to.

Distribution. AUSTRALIAN REGION: New Guinea. NEARCTIC REGION:
United States (Florida, Mississippi, Texas). ORIENTAL REGION: Burma, China
(Anhei, Sichuan, Yunnan, Zhejiang), India (Andaman Islands, Assam), Indonesia (Batoe.
Boreneo, Java, Sumatra), Japan, Malaya, Sri Lanka, Taiwan, Thailand. PALEARCTIC
REGION: Korea.

Specimens Examined. (34 Q; 3 6)

Type material: Paratype Xyleborus mutilatus: Malaya, Salagor, 16:XI:1948, F.G.

Browne ex wood of Adenanthera tauonina (6; BMNH). Lectotype X yleborus

189

banjoewangi (Q; NHMW). Allotype Xyleborus banjoewangi: Banjoewangi, II-l936. leg.
Baschwesen [?] (1 6; NHMW).

Other material: ORIENTAL REGION: Japan: JAPAN: Okinawa 1., Mt. Oppadake.
20.Vi.95, H. Goto, ex Rhus succedanea (2 Q, l 6; RAB); JAPAN: RYUKYUS. Mt.
Oppadake, Okinawa Is., emerged from the logs, 19.vii.1995, H. Goto leg., Host tree:
Rhus succedanea L. (1 Q, 1 6; RAB). NEARCTIC REGION: United States:
Mississippi, Oktibbeha Co., 3 mi W. of Adaton, 33°29’00”N 88°58’13”W, 23-26 April
2004, T. L. Schiefer coll (39 Q; MSU); Miss., Oktibbeha Co., 3 mi W. of Adaton,
33°29’00”N 88°58’13”W, 17-19 April 2002, T. L. Schiefer, in Lindgren funnel trap (1 Q:
MSU).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most of the posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, the protibiae with 7 socketed teeth. and

the elytra that are wider than long, this species is here transferred to C nestus.

Cnestus orbiculatus (Schedl), new combination
Xyleborus orbiculatus Schedl, 1942a: 186. Holotype Q: Borneo; NHMW.
Xylosandrus orbiculatus (Schedl): Wood and Bright, 1992: 800.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.

Distribution. ORIENTAL REGION: Indonesia (Borneo).

190

Specimens Examined. (2 Q; 0 6)

Type material: Holotype Xyleborus orbiculatus (Q; NHMW).

Other material: ORIENTAL REGION: Borneo: BRITISH N. BORNEO, Tenompok.
15.11.1959, T. C. Maa Collector (1 Q; RAB).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle,
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most the entire posterior face, the presence of a coarse median pair of
asperities on the produced anterior margin of the pronotum, and the elytra that are wider

than long, this species is here transferred to Cnestus.

C nestus peruanus (Wood), new combination
X ylosandrus peruanus Wood, 2007: 468. Holotype Q: Satipo [Junin], Peru, V-VI.1942,
Paprzycki; USNM.
Distribution. NEOTROPICAL REGION: Peru (Junin).
Specimens Examined. (1 Q; 0 6)
Type material: Holotype Xylosandrus peruanus (Q; USNM).
Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most the entire posterior face, the presence of a coarse median pair of
asperities on the produced anterior margin of the pronotum, and the protibiae with 6

socketed teeth this species is here transferred to C nestus.

Cnestus retifer (Wood), new combination

191

Xylosandrus retifer Wood, 2007: 468. Holotype Q: Fazende Laminit. Intinga do
Maranhio, Brazil; MZUSP.

Distribution. NEOTROPICAL REGION: Brazil.

Specimens Examined. (1 Q; 0 6)

Type material: Paratype Xylosandrus retifer BR-MA-Itinga do Maranhao, Fazenda
Laminit, 6-yr old Schizolobium nicum stand, ethanol baited FIT, Ataide, J. A. col.
VII/2002 (9; USNM).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering the entire posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, and the elytra that are wider than long,

this species is here transferred to C nestus.

C nestus retusus (Eichhofl), new combination
Xyleborus retusus Eichhoff, 1868: 151. Syntypes Q: N. Freiburg [Brazil]; Hamburg
Museum, lost.
Xylosandrus retusus (Eichhoff): Wood and Bright, 1992: 800.
Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.
Distribution. NEOTROPICAL REGION: Argentina. Brazil.
Specimens Examined. (120 Q; 0 6)

Type material: Unable to examine type specimens.

192

Other material: NEOTROPICAL REGION: Brazil: Brazil: Nova Teutonia, Sta. C at..
X-56, Plaumann (19 Q; USNM); Brazil: Nova Teutonia, 27°11’8”.52°23‘1”, F.
Plaumann, 300-500 m, XI.1956 (1 Q; USNM); Brazil: Nova Teutonia, 27°] 1’8"
52°23’1”, F. Plaumann, 300-500 m, I-l.l957 (1 Q; USNM); Brazil: Nova Teutonia.
1944, F. Plaumann coll. (5 Q; USNM); Brasil, Nova Teutonia, Santa Catarina. 1X-18-
1944, 27°] l’La., 52°23’Lo., Fritz Plaumann collector (I Q; AMNH); Brazilen: Nova
Teutonia, 28 Oct 1951, F. Plaumann coll. (10 Q; FMNH); Brazilen: Nova Teutonia, 25
Oct 1951, F. Plaumann coll. (7 Q; FMNH); Brazilien, Nova Teutonia, 27°11" B. 52°23“
L., 500m, XI.1947, Fritz Plaumann (8 Q; FMNH); Brazil, Nova Teutonia, Sta. Catharina.
VII.14.1944 (1 Q; F MNH); Brasilien: Nova Teutonia, 1944, F. Plaumann coll. (3 Q;
NHMW); Brasil, Rio Claro-S. Paulo, Mat no. 5, III-947 [?] (1 Q; USNM)1 Brasil. Rio
Claro—S. Paulo, Mat no. 2 III-946[?] (1 Q; USNM); Brazil: Santa Catharina, Nova
Teutonia, 300-500 m, 27°11’-52°23’, XI-1947, leg. F. Plaumann (60 Q; FMNH); Brasil.
Varginha, M Gerais, M. Alvarenga leg., 111.1972 (1 Q; NHMW); RS/RGS Exp. Brazil.
12049’S 51046'W, 29.xi.1968, R. A. Beaver (1 Q; RAB).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering the entire posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, the protibiae with 6 socketed teeth. and

the elytra that are wider than long, this species is here transferred to C nestus.

Cnestus testudo (Eggers), new combination

193

Xyleborus testudo Eggers, 1939b: 116. Lectotype Q: Formosa, Trichu, XI.1930.. col. T.
Mitono; USNM; designated by Anderson and Anderson, 1971: 34.

Xylosandrus testudo (Eggers): Wood and Bright, 1992: 801.

Notes. This species was first included in Xylosandrus by Wood and Bright (1992), but
the authors did not indicate it as a “new combination” or cite any characters justifying its
transfer.

Distribution. ORIENTAL REGION: Taiwan, Thailand, Vietnam.

Specimens Examined. (3 Q; 0 6)

Type material: Lectotype X yleborus testudo (Q; USNM).

Other material: ORIENTAL REGION: Thailand: THAILAND: Chiang Mai 600 m.
Fang, 17.iii.74, R. A. Beaver, comp. Paralectotyps ex. TARI. (1 Q; RAB). Vietnam:
Tonkin, Reg De Moa binn, A. De Looman, 1929 (1 Q; NHMW).

Discussion. Based on the subcontiguous procoxae, the 4-segmented antennal funicle.
obliquely truncate antennal club with first segment forming a circular costa and segment
one covering most of the posterior face, the presence of a coarse median pair of asperities
on the produced anterior margin of the pronotum, the protibiae with 7 socketed teeth. and

the elytra that are wider than long, this species is here transferred to C nestus.

New Combinations, C yclorhipidion
C yclorhipidion squamulatus (Beaver and Loyttyniemi), new combination
Apoxyleborus squamulatus Beaver and Loyttyniemi, 1985: 69. Holotype Q: Zambia:
Lusaka, 21 .i.1980, light trap. R. A. Beaver coll; BMNH.

X ylosandrus squamulatus (Beaver and Loyttyniemi): Wood, 1984: 229.

194

Notes. Beaver and Loyttyniemi (1985) described this species in the genus Apo.x;1.'lehorus.
However, Wood (1984) had synonymized Apoxyleborus with X ylosandrus. The species
was listed in X ylosandrus by Wood and Bright (1992).

Distribution. AFROTROPICAL REGION: Zambia.

Hosts. Unknown.

Specimens Examined. (2 Q; 0 6)

Type material: Holotype Apoxyleborus squamulatus (Q; BMNH). Paratype
Apoxyleborus squamulatus: ZAMBIA, Lusaka, 2052, 8-9.v.80, R. A. Beaver, trap light
(9; RAB)-

Discussion. Based on the contiguous procoxae, antennal club with segment one not
forming a circular costa (type 3, Hulcr et a1. 2007), and arched protibiae with 8 socketed
teeth, this species is herin transferred to C yclorhipidion. This species was compared with
the holotype of the type species of C yclorhipidion: C. pelliculosus. The inclusion this
species, along with C ylcorhipidion pelliculosus Eggers, in the phylogenetic analysis of
morphological data caused the strict consensus tree to become completely unresolved.
Since C yclorhipidion has not historically been confused with X ylosandrus, the genus was

not included in the phylogenetic analysis.

195

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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Specres ‘13 Z

X. abruptulus o

X adherescens I

X amputatus n. comb. O

X arquatus O

X assequens O

X beesoni O

X. borealis O O

X boreensis n. sp. 0

X brevis o I

X compactus I A I I I

X. corthyloides O

X crassiusculus I I A o I I A

X curtulus I I

X derupteterminatus O

X deruptulus O

X discolor o I o

X. diversepilosus o

X eupatorii I

X ferinus I

X germanus A O O A

X hirsutipennis O

X hulcri n. sp. 0

X ’aintianus O O

X mancus O O

X mediocris O

X mesuae O O

X metagermanus 0

X mixtus n. comb. I

X monteithi o

X morigerus I I I I o A

X pusillus O

X pygmeaus O

X. queenslandi O

X rotundicollis n. comb. O

X subsimiliformis I

X subsimilis O

X terminatus o

X woodi o

 

 

 

 

 

 

 

 

 

 

Table 2.1: Biogeographic distribution of Xylosandrus species: 0 = present: A = present
as introduced species.

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Characters

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43

33 34 35 36 37 38 39 4O 41 42

28 29 30 3] 32

26 27

25

22 23 24

Species

 

A masa anomalus
A. bicostatus

(‘1

A. cylindriform is

A. cylindrotomicus

A. fulgens

A. omissus
A. oralis

A. striatotruncatus

A. umbratulus
A. versicolor

Anisandrus butamali

A. dispar
A. obesus

A. sayi

A. ursa

N
O

A. ursinus
A. ursulus

Materials

1n

described

11‘ scores

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ies

f 43 characters for 73 spec

1X0

Morphological character matr

Table 2.2 (cont)
and Methods.

Characters

2]

 

42 43

34 35 36 37 38 39 40 4]

33

32

29 30 31

28

27

26

22 23 24 25

Species

 

Cnestus ater

C. bimaculatus

C. fijianus

(\l

("l

(‘1

C. gravidus

C. improcerus
C. laticeps

C. mutilatus

C. orbiculatus
C. peruanus

C. pseudosolidus

C. pseudosuturalis

C. retifer

C. retusus
C. solidus
C. testudo

202

C. triangularis

C occotrypes dactyliperda
X yleborus aflim’s
X. califomicus

Materials

1n

described

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ies

f 43 characters for 73 spec

1X0

Morphological character matr

Table 2.2 (cont)
and Methods.

Characters

2]

 

42 43

36 37 38 39 40 41

32 33 34 35

26 27 28 29 30 31

25

22 23 24

§pecies

 

0

Xylosandrus adherescens

X. amputatus
X. arquatus

(\l

X. assequens
X. beesoni

X. borealis

X. borneensis

X. brevis

X. compact us
X. corthyloides

X. crassiusculus
X. curtulus

X. derupteterminatus

X. deruptul us
X. discolor

N
O
b)

X. diversepilosus
X. eupatorii
X. ferinus

X. germanus

Materials

1n

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ies

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Morphological character matr

Table 2.2 (cont)
and Methods.

Characters

21

 

42 43

32 33 34 35 36 37 38 39 40 41

27 28 29 30 3|

26

25

24

22 23

Species

 

X. hirsutipennis

X. hulcri

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(\l

X. jaintianus
X. mancus

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X. mediocris
X. mesuae

X. metagermanus
X. mixtus

X. montethi

X. morigerus
X. pusillus

X. pygmaeus

X. queenslandi
X. rotundicollis

X. subsimilis
X. subsimiliformis

X. terminatis
X. woodi

204

(‘1

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Gene

 

 

 

Partition
Ef-1a COI ArgK CAD 28$ Morphology Combined
node

1 3.00 2.00 5.00 8.00 0.00 5.00 23
2 3.00 2.00 5.00 8.00 0.00 5.00 23
3 1.00 4.00 2.00 0.00 1.00 -1.00 7
4 0.00 -2.00 2.00 4.00 -2.00 0.00 2
5 8.00 -1.50 7.50 11.00 -1.00 -2.00 22.00
6 3.00 0.00 -1.00 0.00 4.00 0.00 6.00
7 2.00 0.00 -4.00 -2.00 36.00 -4.00 28.00
8 0.00 5.00 0.00 11.00 25.00 5.00 46.00
9 0.00 2.00 -3.00 -4.00 15.00 -4.00 6.00
10 0.00 2.00 -3.00 -4.00 15.00 -4.00 6.00
11 5.00 6.00 2.00 -1.00 63.00 -4.00 71.00
12 2.00 -1.00 -1.00 -1.00 2.00 0.00 1.00
13 5.00 0.00 15.00 10.00 11.00 5.00 46.00
14 3.00 -2.00 1.00 -2.00 2.00 1.00 3.00
15 0.00 1.00 10.00 15.00 8.00 1.00 35.00
16 13.00 5.67 29.67 13.33 8.33 7.00 77.00
17 3.00 2.00 5.00 8.00 0.00 5.00 23.00
18 -1.00 -3.00 -3.00 0.00 26.00 0.00 19.00
19 2.00 0.00 1.00 0.00 5.00 1.00 9.00
20 0.00 -0.50 4.00 5.00 2.00 2.50 13.00
21 2.00 -2.00 -1.00 2.00 5.00 0.00 6.00
22 3.00 -2.00 0.00 10.00 3.00 0.00 14.00
23 8.50 2.50 -0.50 18.50 3.00 11.00 43.00
24 0.00 11.00 7.00 7.00 2.00 -3.00 24.00
Total 65.50 31.17 79.67 116.83 233.33 26.50 442.00

 

Table 2.5: Partition branch support for the most parsimonious tree found by the analysis
of the combined data set. Node numbers refer to Figure 2.2.

210

dact li erda
ailing

   
  
 
   

Amasa cylindrotomicus n. comb.
masa omissus n. comb.

Amasa oralis n. comb.

Amasa anomalus

Amasa bicostatus .

Amasa cylmdrofonms

Amasa fulgens

Amasa striatotruncatus

Amasa umbratulus

Amasa verswolor

Figure 2.1: Strict consensus of 10,000 most parsimonious trees found in parsimony
analysis of the morphological data martrix. Bootstrap values are given below the nodes
for clades with support 2 90%.

211

Amasa bicostatus

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

10343 24 »— Amasa striatotruncatus
1%“ 4 1000-4 — Amasa versicolor
.2— Xyleborus califomicus
2 Xyleborus afiinis
92/23 I r—— Xylosandrus mancus
92/23 Xylosandrus discolor
I5 Cnestus bimaculatus
100/ 35 Cnestus pseudosuturalis
101377 13 Cnestus ater n. comb.
17 14 100/46 Cnestus mutilatus n. comb.
97/23 Cnestus improcerus n. comb.
Xylosandrus crassiusculus
18 Anisandrus dispar
97/19 H Anisandrus obesus
100/71 9 i___— Anisandrus ursa n. comb.
lO 8
19 I2 100/461——— Anisandrus ursinus n. comb.
Anisandrus sayi
7 Xylosandrus moteithi
100/ 28 Xylosandrus hulcri n. sp.
20 Xylosandrus rotundicollis n. comb.
Xylosandrus compactus
1005/22 A Xylosandrus germanus
6 4 Xylosandrus borneensis n. sp.
% Xylosandrus morigerus
Xylosandrus queenslandi
Coccotrypes dactyliperda
—— 50 changes

Figure 2.2: Most parsimonious tree found in the analysis of the combined data matrix.
Node numbers given above branches and bootstrap support/Bremer support values given
below.

212

 

Figure 2.3: Lateral habitus of Amasa cylindrotomicus n. comb.. female (A) and Amasa
bicostatus, female (B).

213

 

Figure 2.4: Lateral habitus of Anisandrus ursulus n. comb., female (A) and Anisandrus
sayi, female (B).

214

 

Figure 2.5: Lateral habitus of Cnestus improcerus n. comb.. female (A) and Cnestus
pseudosuturalis, female (B).

215

 

Figure 2.6: Lateral (A) and dorsal (B) views of Xylosandrus abruptulus, 1.9 — 2.1 mm,
female lectotype.

216

 

Figure 2.7: Lateral (A) and dorsal (B) views of X ylosandrus adherescens, 2.0 mm,
female holotype.

217

 

Figure 2.8: Lateral (A) and dorsal (B) views of Xylosandrus amputatus n. comb., 2.7 —
2.9 mm, female.

218

 

Figure 2.9: Lateral (A) and dorsal (B) views of X ylosandrus arquatus, 2.3 — 2.5 mm,
female.

219

 

Figure 2.10: Lateral (A) and dorsal (B) views of Xylosandrus assequens. 1.6 — 2.3 mm,
female, holotype.

220

‘4 3., _, g
.7 ”NHL {twice-9% — a

 

Figure 2.11: Lateral (A) and dorsal (B) views of Xylosandrus borealis, 2.0 — 2.1 mm.
female.

221

 

B

Figure 2.12: Lateral (A) and dorsal (B) views of Xylosandrus borneensis n. sp., 1.5 — 1.6
mm, female holotype.

222

 

B

Figure 2.13: Lateral (A) and dorsal (B) views of Xylosandrus brevis, 2.5 — 2.8 mm.
female.

223

 

Figure 2.14: Lateral (A) and dorsal (B) views of Xylosandrus compactus. 1.4 — 1.9 mm,
female.

224

 

Figure 2.15: Lateral (A) and dorsal (B) views of Xylosandrus corthyloides, 2.7 — 3.0
mm, female lectotype.

225

 

Figure 2.16: Lateral (A) and dorsal (B) views of Xylosandrus crassiusculus, 1.7 - 2.9
mm, female.

226

 

Figure 2.17: Lateral (A) and dorsal (B) views of Xylosandrus curtulus, 1.3 — 1.5 mm,
female holotype of synonym X. biseriatus.

227

 

Figure 2.18: Lateral (A) and dorsal (B) views of Xylosandrus derupteterminatus, 2.0 —
2.3 mm, female holotype.

228

 

Figure 2.19: Lateral (A) and dorsal (B) views of Xylosandrus deruptulus. 1.8 mm,
female lectotype.

229

 

Figure 2.20: Lateral (A) and dorsal (B) views of Xylosandrus discolor, 1.8 — 2.0 mm,
female.

230

 

Figure 2.21: Lateral (A) and dorsal (B) views of Xylosandrus diversepilosus, 1.8 — 2.3
mm, female holotype.

231

 

Figure 2.22: Lateral (A) and dorsal (B) views of Xylosandrus eupatorii, 1.8 — 2.1 mm,
female cotype.

232

 

Figure 2.23: Lateral (A) and dorsal (B) views of Xylosandrus ferinus, 1.6 — 1.8 mm,
female lectotype.

233

 

Figure 2.24: Lateral (A) and dorsal (B) views of Xylosandrus germanus, 1.9 — 2.5 mm.
female.

234

 

Figure 2.25: Lateral (A) and dorsal (B) views of Xylosandrus hirsutipennis, 1.9 — 2.2
mm, female paratype.

235

 

Figure 2.26: Lateral (A) and dorsal (B) views of Xylosandrus hulcri n. sp., 2.4 — 2.7mm.
female holotype.

236

 

Figure 2.27: Lateral (A) and dorsal (B) views of Xylosandrus jaintianus, 3.0 mm, female
holotype.

237

 

Figure 2.28: Lateral (A) and dorsal (B) views of Xylosandrus mancus, 2.9 — 3.3 mm,
female.

238

 

Figure 2.29: Lateral (A) and dorsal (B) views of Xylosandrus mediocris, 1.4 mm, female.

239

 

Figure 2.30: Lateral (A) and dorsal (B) views of Xylosandrus mesuae, 1.1 — 1.3 mm,
female.

240

 

Figure 2.31: Lateral (A) and dorsal (B) views of Xylosandrus metagermanus, 1.8 mm.
female holotype.

241

 

Figure 2.32: Lateral (A) and dorsal (B) views of Xylosandrus mixtus n. comb., 2.6 — 2.7
mm, female holotype.

242

 

Figure 2.33: Lateral (A) and dorsal (B) views of Xylosandrus monteithi, 3.0 — 3.4 mm,
female paratype.

243

 

Figure 2.34: Lateral (C) and dorsal (D) views of Xylosandrus monteithi, 2.5 mm, male
allotype.

244

 

Figure 2.35: Lateral (A) and dorsal (B) views of Xylosandrus morigerus, 1.5 — 2.0 mm,
female.

245

 

Figure 2.36: Lateral (A) and dorsal (B) views of Xylosandrus pusillus, 1.5 — 1.7 mm
female holotype.

,

246

 

Figure 2.37: Lateral (A) and dorsal (B) views of Xylosandrus pygmaeus, 1.3 — 1.4 mm,
female holotype.

247

 

Figure 2.38: Lateral (A) and dorsal (B) views of Xylosandrus queenslandi, 1.6 — 1.9 mm.
female paratype.

248

 

Figure 2.39: Lateral (A) and dorsal (B) views of Xylosandrus rotundicollis n. comb., 3.7
— 4.1 mm, female holotype.

249

 

Figure 2.40: Lateral (A) and dorsal (B) views of Xylosandrus subsimiliformis. 2.8 mm,
female cotype.

250

 

Figure 2.41: Lateral (A) and dorsal (B) views of Xylosandrus subsimilis, 2.6 mm, female
cotype.

251

 

Figure 2.42: Lateral (A) and dorsal (B) views of Xylosandrus terminatus, 1.5 — 1.9 mm,
female cotype.

252

 

  

B

Figure 2.43: Lateral (A) and dorsal (B) views of Xylosandrus woodi, 2.3 — 2.4 mm,
female paratype.

253

CHAPTER 3

Diversity of Scolytinae (Coleoptera: Curculionidae) in the Ecuadorian rain forest
canopy

254

ABSTRACT

Canopy fogging was used to sample the diversity of bark and ambrosia beetles
(Coleoptera: Curculionidae: Scolytinae) at two Western Amazonian rain forest sites in
Ecuador. Sampling spanning from 1994-2006 yielded 1,158 samples containing 2.500
scolytine specimens representing more than 400 morphospecies. Here, we analyze a
subset of this data representing two ecological groups: true bark beetles (52
morphospecies) and ambrosia beetles (69 morphospecies). A high percentage of these
taxa occurred as singletons and doubletons and their species accumulation curves did not
reach an asymptote. Diversity estimates placed the total scolytine species richness for
this taxon subset present at the two sites between 260 and 323 species. High levels of (x-
and B-diversity were discovered. However, high B-diversity was found to be an artifact
of undersampling and does not appear to be biologically significant. This study
demonstrates the utility of canopy fogging methods for the sampling of scolytine richness

and for the discovery of new scolytine taxa.

255

INTRODUCTION

Studies of tropical canopy ecosystems have documented the richness of this
habitat and its potential for the discovery of new arthropod species (Erwin 1982, 1983a
1983b, 1988; Lowman and Witman 1996; Erwin et al. 2005). Erwin’s studies of
arthropod diversity in tropical forest canopies resulted in the controversial estimate of 30-
50 million arthropod species (Erwin 1982). Subsequently, several authors were critical of
this estimate (May 1988; Adis 1990; Gaston 1991), which was actually intended to spark
discussion of the use of hypothesis driven data analysis for global species estimation and
the adequacy of current sampling methods, rather than be an authoritative estimation of
global biodiversity (Erwin 1991). Nevertheless, Erwin’s estimate highlighted the under-
explored biodiversity contained in the canopy ecosystem and the need to document and
describe canopy species in light of tropical deforestation.

The subfamily Scolytinae (Coleoptera: Curculionidae) is comprised of
approximately 225 genera containing 6,000 species worldwide (Wood and Bright 1992).
Scolytine species feed sub-cortically on a wide variety of woody and herbaceous plants.
The typical life cycle of scolytines consists of a brief dispersal flight period after adult
emergence, followed by colonization of new hosts. Once a suitable host has been
located, adults bore galleries into the host tissue, where eggs are laid, and larvae complete
their development into the next generation of adults.

Scolytine beetles can be divided into two ecological groups: bark beetles and
ambrosia beetles. The degree to which scolytines specialize on specific hosts varies
considerably depending on these ecological groups. Bark beetles bore into the phloem of

trees, feed on tree tissues, and tend to have more specialized host preferences. Ambrosia

256

beetles bore into the xylem, feed on species of symbiotic fungi which grow along the
walls of their galleries, and tend to have more generalized tree host preferences (Beaver
1979; Kirkendall 1993). The majority of scolytine diversity occurs in tropical regions of
the world, with much of this species richness believed to be undescribed.

First pioneered by Southwood (1961) in temperate forests and then later adapted
for tropical research by numerous others (Lowman and Witman 1996), canopy fogging
methods use insecticide to collect arthropods from the upper architecture of the forest
habitat. These methods have become widely used in surveying arthropod diversity,
particularly in lowland rain forests (Basset 2001). For a more than a decade, Erwin and
colleagues have sampled arthropod diversity in the Ecuadorian Amazon rain forest
canopy at two lowland rain forest sites, separated by 21km of contiguous primary forest.
using a standardized insecticidal fogging protocol (Erwin 1983a, 1983b; Erwin et al.
2005). Sampling has taken place during dry, wet, and transitional seasons to measure
temporal turnover in species composition. Tree data were also recorded for fogging
stations within the two study sites (Pitman et a1. 2001). Canopy fogging samples
collected in these surveys offer a potentially rich source of new scolytine specimens
because collecting scolytines directly from their hosts can require great effort and can
also be limited by the abundance and accessibility of colonized hosts. Thus canopy
fogging may provide taxonomists access to species which complement those collected
using traditional, ground-based, hand-collecting methods.

Remarkably few studies have looked at the spatial and temporal turnover of
scolytine species (Ii-diversity) (Deyrup and Atkinson 1987; Thunes 1988; Peltonen et a1.

1998; Hulcr et al. 2007, in press). The majority of studies have focused on the

257

composition of the scolytine fauna of temperate regions and examined the distributions of
a few economically important species (Deyrup and Atkinson 1987; Jakus 1998; Peltonen
et al., 1998; Oliver and Mannion 2001; Gaylord et a1. 2006). Several studies in the
tropics have attempted to determine the effects of seasonal changes in rainfall and
temperature to the composition of scolytine communities (Beaver and Loyttyniemi 1991;
Madoffe and Bakke 1995; Morales et al. 2000; F lecthmann et a1. 2001; Hulcr et al. in
press). Studies in Malaysian forests have examined the spatial distribution of scolytines
across horizontal (Maeto et al. 1999; Chung 2004) and vertical gradients (Maeto and
Fukuyama 2003; Simon et al. 2003). Recent studies in Papua New Guinea (Hulcr et al.
2007) and Thailand (Hulcr et al. in press) have used quantitative sampling schemes to
examine scolytine community composition and attempt to determine the proximate
causes of the distribution of species in tropical habitats. Scolytine communities in
lowland Papua New Guinean rain forests have been found to have low B-diversity, thus
contradicting the expected trend of high B-diversity in the tropics. Novotny et a1. (2007)
similarly demonstrated low species turnover for insect communities when examined on a
large scale across New Guinean forests. In Thailand, two study sites separated by only 5
km were found to have significantly different scolytine species composition (Hulcr et al.
in press). Correspondence analysis found that differences in mean annual temperature
and humidity, rather than seasonal fluctuations. explained the differing species
composition at the two sites. However, the Thailand study sites differed from each other
greatly, one being a mixed evergreen forest located at 1,410 meters altitude and the other
a deciduous dicterocarp forest at the edge of the city of Chiang Mai at 350 meters

altitude. Such habitat variation did not exist in the study of lowland New Guinean rain

258

forests or the two Amazonian study sites examined herein. Thus, it is uncertain whether
Amazonian forests will produce the same trend of low B-diversity seen in Papua New
Guinea or high B-diversity, as was observed between different Thai forests.

In tropical ecosystems, an increase in host specificity combined with an increase
in plant diversity is often used to explain high levels of species diversity. However, bark
and ambrosia beetles show a reverse trend, with a decrease in host specificity in tropical
regions. This is largely due to the greater abundance of ambrosial feeding scolytines
which, as discussed above, tend to be relative host generalists (Beaver 1979).
Nevertheless, most scolytine groups have higher species diversity in tropical regions than
they do in temperate ones (Beaver 1979; Deyrup and Atkinson 1987; Wood and Bright
1992). It is therefore expected that canopy fogging will yield a high number of scolytine
species, but that the B-diversity of scolytines across the Amazonian canopy may not
necessarily be high.

In his 2007 taxonomic monograph of the bark and ambrosia beetles of South
America, Wood lists less than 50 scolytine species records from Ecuador (Wood 2007).
Most scolytine taxonomists would doubt that this paucity of species records represents
the true scolytine diversity contained within the rich tropical ecosystems of the
. Ecuadorian Amazon. What, then, is the true extent of scolytine diversity in Ecuador and
what sampling methods can be utilized to uncover the uncollected and undescribed
diversity that remains?

In this study, we use a subset of scolytine subtribes representing both the bark
beetle (Hylesinini: Bothrostemina, Phloeosinina, Phloeotribina, Phrixosomina) and

ambrosia beetle (Xyleborina, Premnobina) ecological groups to assess the diversity of

259

scolytines at two Western Amazonian forest study sites. We use this data to assess the
value of canopy fogging as a source of scolytine specimens, determine the spatial
turnover of scolytine species, and use the data collected from these foggings to predict
the scolytine species richness at the two sites. Alpha taxonomy for newly discovered
genera and species is a longer-term component of this research and is being undertaken in

conjunction with this work as well (Dole and Cognato 2007).

MATERIALS AND METHODS

Field Sites and Sampling

The two study sites in this investigation were typical lowland rain forest habitats
in the western Amazon Basin at the margin of Yasuni National Park, separated by 21 km
of contiguous primary forest: Onkone Gare Station (cited as “Pirafia” in Pitman et al.
2001 and Erwin et al. 2005) (0° 39’ 25.685” S, 76° 217’ 10.813” W; 216 m) and Tiputini
Biodiversity Station (00 37’ 55.397” S, 76° 08’ 39.205” W; 216 m). The Onkone Gare
and Tiputini study plots were established in 1994 and 1997, respectively. The study sites
receive an average of 2.7 m of rainfall per year (Erwin et al. 2005). Precipitation at the
two sites is seasonal, with the wet season occurring from May to October and the dry
season occurring from November to April.

Tree data were recorded for collecting stations within Erwin”s canopy fogging
study transects. Trees with a diameter at breast height (diameter measured at 1.33 m
from tree base) greater than 10 cm that had at least part of a branch hanging over the

collecting sheet were tagged by Erwin’s team and subsequently identified by Pitman and

260

colleagues (Pitman et al. 2001). In terms of trees, the Onkone Gare and Tiputini sites are
very different at the species level (73% occurring at only one site), moderately dissimilar
at the generic level (53% occurring at only one site), and fairly similar at the family level
(26% occurring at only one site). With approximately 250 species each, the two study
sites represent individually 21.26% (Onkone Gare) and 21.42% (Tiputini) and
collectively 34% of the regional tree diversity (Erwin et al. 2005).

The fogging protocol used was one that has been developed by Erwin (Erwin
1983a, 1983b). The study plot area at each site (100 m x 1000 m) was divided into 10
transects (10 m x 100 m). Each transect consisted of 10 collecting stations (3 m x 3 m).
which were randomly arrayed on both sides of the transect centerline. Each station is
constructed of a sampling sheet tied and suspended 1 m off the ground, on which fogged
arthropods fall from the canopy, and a collecting jar attached to the center of the sheet.
The total area of these collecting stations was 9250 m2, which represented just 1.11% of
the entire transect at each site.

Fogging took place three times per year: January/February (dry), J une/J uly (wet),
and October (transitional). Fogging took place at Onkone Gare from 1994-1996 and
from 2005-2006. Fogging took place at Tiputini from 1998-2002 (intermittent research
funding and social unrest in Ecuador interrupted the research at Tiputini after the plot had
been established). The pyrethroid insecticide resmythrim was fogged for 60 seconds in a
column from just above the sheet to a height that was then recorded for each fogging
event (for details on fogging techniques and equipment used see Lucky et al. 2002).
Foggings took place at 0345-0500 hr in order to minimize insecticidal drift outside of the

column due to air currents. Previous studies had demonstrated that arthropod

261

repopulation occurs within 10 days after fogging (Lucky et al. 2002). Hence, the same
stations could be resampled at each seasonal collecting event without an effect on
sampling.

The samples examined herein represent only a subset (1,158 samples) of the total
(1,400+ samples) taken during the decade of canopy fogging conducted. Samples from
several collecting expeditions have yet to be exported from Ecuador and were therefore
not available for this study. Samples were sorted and 14 target taxa for the larger canopy
arthropod diversity study were extracted, including adult Coleoptera. Scolytines were
extracted from the adult Coleoptera samples, sorted to morphospecies. and identified

using published keys (Wood 2007).

Statistical Analysis

A goal of the statistical analyses was to obtain results that were comparable to
those of other analyses conducted on samples from the same foggings and to similar
studies of scolytine B-diversity (Hulcr et al. 2007, in press). For this reason, statistics
used were determined by previous studies (Lucky et al. 2002; Erwin et al. 2005; Hulcr et
al. 2007, in press) and by a selection of the most robust methods currently available in B-

diversity research.

Species accumulation curves and richness estimators
EstimateS Version 7.5 software (Colwell 2004) was used to calculate species
accumulation curves and richness estimators for the fogging data. For all analyses, the

term “singletons” is used in reference to species represented by a single specimen from a

262

sample, while “doubletons” are defined as species represented by two specimens from a
sample.

Species accumulation curves and richness estimators were used to assess the
completeness of the sampling effort and to predict the overall diversity of scolytines
occurring in the canopy habitat. Richness estimators employ the same principles as
mark-release-recapture techniques used to estimate animal populations. Just as large
populations are indicated by capturing new individuals, rather than the same ones over
and over again, species diversity is indicated by capturing new species rather than
individuals of the same species over and over again. Hence, richness estimators
recognize that different information about diversity is contained within the rare units of
the sample (singletons and doubletons) than in the most abundant units (species
represented by multiple specimens). However, these estimators also recognize that rare
species can skew estimates of diversity and, therefore, several richness estimators treat
the rarer species differently in their calculations than they do the more abundant ones.

There are two categories of richness estimators: abundance-based and incidence-
based. The two abundance-based richness estimators used were Chaol (Chao 1984) and
the Abundance-based Coverage Estimator (ACE) (Chao and Lee 1992; Chao et al. 1993).
The two incidence-based richness estimators used were Chao2 (Chao 1987) and the
Incidence-based Coverage Estimator (ICE) (Lee and Chao 1994). ACE and ICE consider
two classes of species for their calculations: those that are rare and those that are not rare.
For analytical purposes, ACE and ICE consider species with fewer than 10 individuals in
the sample to be rare. The choice of the number 10 is arbitrary and is merely the value

suggested by EstimateS (Colwell 2004). Chaol and ChaoZ treat all species (rare and not

263

 

rare) the same in their calculations. These estimates also attempt to account for the effect
of unseen rare species (rare species that were not sampled). The default bias-corrected
option was used to calculate Chaol and Chao2 in EstimateS (Colwell 2004). This
analysis estimated the critical value for the abundance distribution to be > 0.5 for all
subsets of the data analyzed. In cases such as these, Chao recommends that the Chaol .
Chao2, ACE, and ICE be recalculated using the Classic option instead. Thus, the
richness estimators reported herein were calculated using the Classic option under
diversity settings in EstimateS (Colwell 2004).

In addition to the above richness estimators, a second-order jackknife (Burnham
and Overton 1978, 1979) was calculated using EstimateS (Colwell 2004). This measure
is based on the number of species that occur in one sample, as well as those that occur in
exactly two samples. Colwell and Coddington (1994) have shown the second-order

jackknife to be one of the more reliable predictors of species diversity.

Similarity Estimators

Several measures of diversity were used to analyze these samples to determine the
distinctness of the species composition in each sample. The simplest of these measures is
the Complementarity Index (CI) (Colwell and Coddington 1994) that expresses the
distinctness or dissimilarity between two samples. This calculation is based on the
observed species for each sample, the total species richness, and the number of species
unique to either sample. Complementarity Index values range from 0 to 1.0, where C I =
0 means that the compared samples have all species in common while a CI = 1.0 means

that they do not share any species. Complementary indicies were calculated for each of

264

 

the subtribes examined herein, as well as for the Xyleborina + Premnobina, Hylesinini,
and for the total sample (Hylesinini and Xyleborina + Premnobina) for Onkone Gare and
Tiputini.

More sophisticated measures of shared species attempt to estimate the similarity
of species composition of the samples based on the observed data. These measures
include the classic J accard and Sorensen indicies (Chao et al. 2005, 2006). However,
these measures are notoriously sensitive to sample size, especially when rare species are
present in the samples. They are also based solely on incidence (presence-absence) data
and cannot account for unseen shared species. The more accurate Chao-Sorensen
abundance-based and incidence-based estimators correct for this by incorporating the
effect of unseen shared species (Chao et al. 2005, 2006).

The Onkone Gare study site was sampled more than twice as much as the Tiputini
site. To correct for this, the Onkone Gare sample was rarified to contain the same total
number of individuals and a similar distribution of abundances as the Tiputini site
[Tiputini mean (SD) = 1.93 (12.21); rarified Onkone Gare mean (SD) = 1.93 (12.70)].
The total number of individual specimens for both the Tiputini and the rarified Onkone
Gare samples equaled 230. Two subsequent statistical analyses (Monte Carlo and Chao-
Sorensen analyses) used this rarified data.

We performed a Monte Carlo analysis on the total (Hylesinini and Xyleborina +
Premnobina) dataset to test the hypothesis that the observed similarity is not statistically
significant, but rather a result of random sampling of individuals from similar or identical

scolytine communities. In each replicate of this test individuals of each species were

265

 

randomly distributed between the two sites. This randomized dataset was then used to

calculate the Chao-Sorensen similarity index and the procedure was repeated 100 times.

RESULTS AND DISCUSSION

A total of 1,158 canopy fogging samples were analyzed from the Ecuadorian
Amazon study transects (Table 3.1). Of these, 965 were from the Onkone Gare study site
and 293 were from the Tiputini study site. These samples contained a total of 2,500
individual scolytines, representing more than 400 morphospecies. Scolytines were found
in 60% of the Onkone Gare samples, 75% of the Tiputini samples, and 69% of the total
samples for both sites. The canopy fogging samples contained a large number of rare
species occurring as singletons and doubletons. The subset of taxa examined here

represented 688 individuals and 121 species from this larger dataset (Tables 3.2 and 3.3).

Bark beetles

Bark beetles belonging to the tribe Hylesinini were represented by four subtribes
in the canopy fogging samples: Bothrostemina, Phloeosinina, Phloeotribina, and
Phrixosomina. A total of 183 specimens representing 9 genera and 52 morphospecies of
hylesines were collected (Tables 3.2 and 3.4). In his monograph of South American
Scolytinae Wood (2007) records 13 genera and 207 species of Hylesinini from the region.
Assuming that none of the genera or species collected were previously undiscovered,
canopy fogging yielded 69% of the known genera and 25% of the known species from

South America. However, at least one genus found by the foggings was previously

266

undiscovered: Akrobothrus Dole and Cognato (2007) (Bothrostemina) and several
species have been identified as undescribed. These discoveries suggest that canopy
fogging is not only a useful method for the collection of known scolytine fauna, but also

an effective means for discovering new taxa.

Ambrosia beetles

Ambrosia beetles were represented in our analyses by two subtribes: Xyleborina
and Premnobina. Xyleborines exhibit highly skewed sex ratios (Bright 1968; Kirkendall
1993), with males rarely being collected, and the taxonomic classification of the subtribe
is based on female morphology. Although 18 specimens of xyleborine males were
collected by this study, it was impossible to associate these with their female counterparts
with any certainty. Therefore, these individuals were not included in the analysis. A
total of 504 specimens representing 8 genera and 68 species of Xyleborina were collected
via canopy fogging (Tables 3.3 and 3.4). Wood (2007) records 11 genera and 233
species of Xyleborina from South America. Thus, with the exception of any undescribed
genera or species, fogging yielded 73% of the known genera and 29% of the known
xyleborina species from the region. However, as with the subtribes of Hylesinini, several
morphospecies of Xyleborina appear to represent previously undiscovered taxa.

A single species belonging to the xyleborine genus Amasa was collected via the
canopy foggings. Wood (2007) noted that the existence of Amasa in South America had
been based on an erroneous record but still included the genus in his key to xyleborine
genera of the region. Therefore, this collection represents the first confirmed record of

A masa from South America. Comparison of this species with known Amasa from other

267

 

regions will be necessary to assess whether it is likely a native or introduced species, as
invasive xyleborines have been known to inhabit primary Neotropical forests (Kirkendall
and Qdegaard 2007). A single genus and species of Premnobina was collected:
Premnobius clavipennis Eichhoff. Premnobina is a monogeneric subtribe containing

three South America species (Wood 2007).

Species Accumulation Curves and Richness Estimators

Species accumulation curves for both sites combined and for each site
individually have yet to asymptote at the current sampling levels (Figs. 3.1-3.3). The
steadily increasing species accumulation curves are correlated with a continued
occurrence of rare species (singletons and doubletons) in the samples. This indicates that
our sampling was not adequate to sample the entire scolytine species composition of the
two study sites.

Estimates of true species richness differed between abundance-based and
incidence-based statistics (Table 3.1; Figs. 3.4-3.6). For the two sites combined, the
abundance-based ACE and Chao] estimated a total scolytine species richness of 301 and
308 species, respectively. The incidence-based ICE and Chao2 gave the higher estimates
of 323 and 311 species. Abundance-based and incidence-based estimates of species
richness for the Onkone Gare overlapped slightly. For Onkone Gare, ACE estimated the
total scolytine fauna to be 275 species and Chaol estimated 296 species. Incidence-based
estimates were slightly higher, with ICE estimating 292 species and Chao2 estimating
309 species. Abundance-based and incidence-based statistics arrived at similar estimates

for the Tiputini site. ACE estimated the total richness to be 90 species and Chaol

268

estimated 85 species. The incidence—based statistics, ICE and Chao2 estimated greater
species diversity, with a total of 96 and 90 species, respectively.

The second-order Jackknife estimated a scolytine species richness of 260 species
for both study sites (Table 3.1; Figs. 3.4-3.6). For Onkone Gare, the second-order
J ackknife estimated 217 species and for Tiputini it estimated 95 species.
Shared Species Estimators

Simple Complementarity Indicies found that the composition of the scolytine
fauna was markedly different for the two study sites. Onkone Gare and Tiputini had a C1
= 0.81 for the total analyzed subset of Scolytine subtribes (Table 3.1). Complementarity
Indices for various subtribes of Scolytinae ranged from 0.73 to 1.00 (Table 3.5). In the
samples, at least half of all the species occurring at either site were unique to that site:
76% of species at Onkone Gare and 50% or species at Tiputini (Table 3.1). However, the
Monte Carlo test concluded that the observed difference between the two sites was due to
stochastic sampling error and does not appear to be biologically significant. The Monte
Carlo radonmization test returned a probability of 0.374 [median modeled similarity L' =
0.77 (lower and upper 2.5% quantiles = 0.634 and 0.836)]. These results conclude that it
would be impossible to assess whether the actual dissimilarity between Onkone Gare and
Tiputini is biologically significant without further sampling. Since the species
accumulation curves for Onkone Gare and Tiputini have not assymptoted, any
comparison of the two study sites must be considered preliminary and cannot be seen as
statistically valid result.

Our results indicate that even a large scale, long-terrn sampling effort, such as this

one, was not adequate for uncovering the true scolytine species richness contained in the

269

 

 

rain forest canopy at two Western Amazonian sites. Despite over 1,100 individual
fogging events (representing 14 fogging expeditions), which collected 688 individual
beetles representing 121 species, the species accumulation curves did not reach an
asymptote (Figs. 3.1-3.3). Likewise, the occurrence of rare species in the form of
singletons and doubletons did not decline. These phenomena were observed from the
individual data collected for each study transect and for the collective species data for the
two sites.

Since the levels of scolytine B-diversity observed herein were found to not be
biologically significant and species accumulation curves have not assymptoted, it would
be invalid to compare our results with those of other studies. The different levels of [3-
diversity found by studies of Papua New Guinean and Thai scolytine faunas suggests that
patterns in scolytine B-diversity may vary regionally within tropical forest ecosystems
(Hulcr et. al 2007, in press). Previous canopy fogging studies have found significant
differences in the species composition of different forest types. In Manaus, Brazil, 83%
of beetle species in canopy fogging samples were found in only one type of forest (Erwin
1983a). Of the four forest types characterized and studied by Erwin, “mixed-water”
forests were found to be the most species rich. However, “terre firme” (non—floodplain)
forests had the second highest number of species and the highest number of restricted
species (IS-diversity). Both sites sampled in this study were “terre firme” forest. Erwin
also found that “terre firme” forests carry a high load of smaller insect species (1 mm
class, as defined for his study). This offers another predictor of the true scolytine
diversity in this habitat, as most scolytine species are within the 1-3 mm range. If

scolytnes follow the same general patterns found for all Amazonian insects, we can

270

 

predict that further sampling would uncover an even richer scolytine fauna with possibly
many species not found in other forest types.

One goal of this study was to assess the value of canopy fogging as a source of
scolytine specimens. Prior to this work, there were less than 50 scolytine species records
from Ecuador (Wood 2007). Our entire collection of scolytines from the foggings
represented over 400 morphospecies, the inclusion of these species into the fauna] record
will dramatically increase the number of species that are known from Ecuador and the
two Amazonian sites used in this study represent only one of the many habitat types
known to occur in the region. Although extensive floral data exists for the two study
sites (Pitman et al. 2001), caution must be taken when making associations of scolytine
species with particular hosts based on tree data from the sampling sites. It is unlikely that
scolytines would actually be extracted from their galleries by the insecticide used in
fogging. The typical scolytine reaction to disturbance is to move deeper into the gallery.
rather than to vacate it. Therefore, it is most likely that scolytines collected by canopy
fogging are those in flight or on the surfaces of trees and other canopy vegetation. It
would be erroneous to assume that scolytines fogged from a particular tree utilize that
tree as their host. However, general correlations between tree and beetle diversity could
be made using the data collected via canopy fogging. Therefore, despite the fact that
some data critical to understanding of scolytine ecology is lost with the use of canopy
fogging methods, we have demonstrated that fogging offers a method for sampling
scolytine diversity that can uncover many species missed by traditional hand-collecting
methods. For large-scale sampling of scolytine diversity, canopy fogging offers a

quantitative method that can be repeated multiple times at a single site.

271

Large-scale sampling of tropical habitats offers a rich source of previously
unknown species, but as this study demonstrates, even the most ambitious sampling
schemes may not be enough to adequately assess the true species richness of hyper-
diverse groups. In addition to this, the proper application of statistical tools is necessary
to avoid erroneous conclusions and over estimates of B-diversity based on stochastic
sampling (Hulcr et al. 2007, Lewinsohn and Roslin 2008). Scolytines demonstrate that a
single taxon can show multiple patterns of diversity, even within tropical ecosystems.
Based on the sampling of a single Amazonian habitat, we have uncovered a level of
scolytine a-diversity that increases the known fauna of Ecuador nearly ten-fold.
Ultimately relating this figure to the B-diversity of the region could help reveal how the

compostition of Neotropical scolytine communities are affected by habitat changes.

272

 

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273

 

Subtribe Genus Species Locality(ies) No.§pecimens

 

 

Bothrostemina Akrobothrus ecuadoriensis Onkone Care 3
n. sp.. nr. Onkone
Bothrostemina Bothrostemus truncatus Gare/Tiputini 8
Bothrostemina Bothrosternus sp. Tiputini 1
Bothrostemina Bothrostemus sp. Onkone Gare l
Bothrostemina Bothrosternus sp. Onkone Gare l
Bothrostemina Bothrostemus sp. Onkone Gare I
Bothrostemina Cnesinus sp. Onkone Gare 4
Bothrostemina Cnesinus sp. Onkone Gare 6
Bothrostemina Cnesinus sp. Tiputini l
Bothrostemina Cnesinus sp. Tiputini 2
Bothrostemina C nesinus sp. Tiputini 2
Onkone
Bothrostemina Cnesinus sp. Gare/Tiputini 4
Onkone
Bothrostemina Cnesinus sp. Gare/Tiputini 4
Bothrostemina C nesinus sp. Onkone Gare . l . .
Onkone _
Bothrostemina Cnesinus sp. Gare/Tiputini ' i i ' 5" i '
Bothrostemina Cnesinus sp. Onkone Gare l
Onkone
Bothrostemina Eupagiocerus sp.. Gare/Tiputini 2
Onkone
Bothrostemina Pagiocerus sp. Gare/Tiputini 4
Bothrostemina Stemobothrus sp. Tiputini 2
Bothrostemina Sternobothrus sp. Onkone Care I
Bothrostemina Stemobothrus sp. Onkone Gare 1
Bothrostemina Sternobothrus sp. ' Onkone Gare l
Phloeosinina Chramesus sp. Onkone Gare 1. -' '
Phloeosinina Chramesus sp. Tiputini l
Phloeosinina Chramesus sp. Onkone Gare l
Phloeosinina Chramesus sp. Tiputini l
Phloeosinina Chramesus sp. Onkone Gare l
Phloeosinina Chramesus sp. Onkone Gare l

 

Table 3.2 (cont. on next page): Morphospecies of bark beetles (Hylesinini) found in the
canopy fogging samples.

274

 

Subtribe

Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phloeotribina
Phrixosimina
Phrixosimina
Phrixosimina

Table 3.2 (cont): Morphospecies of bark beetles (Hylesinini) found in the canopy

Genus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phloeotriubus
Phrixosoma
Phrixosoma
Phrixosoma

fogging samples.

Species
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp.
sp..
sp.
sp.

275

Locality(ies)

Onkone Gare/Tiputini

Onkone Gare

Onkone Gare/Tiputini
Onkone Gare/Tiputini

Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare
Tiputini

Onkone Gare
Tiputini

Onkone Gare/Tiputini

Onkone Gare
Onkone Gare
Tiputini

Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare
Onkone Gare

No. Specimens

31
20

—' IQ

 

 

 

 

Subtribe Genus Species Locality(ies) No. Specimens
Xyleborina Amasa sp. Onkone Gare l
Xyleborina A mbrosiodmus sp. Onkone Gare 1
Xyleborina C optoborus sp. Onkone Gare/Tiputini 6
Xyleborina C optoborus sp. Onkone Gare I
Xyleborina Coptoborus sp. Onkone Care 2
Xyleborina Coptoborus sp. Onkone Gare/Tiputini I6
Xyleborina Coptoborus sp. Onkone Gare/Tiputini I I
Xyleborina C oproborus sp. Onkone Gare/Tiputini 2
Xyleborina C oploborus sp. Tiputini I
Xyleborina Coptoborus sp. Onkone Gare/Tiputini 2
Xyleborina Coptoborus sp. Onkone Gare/Tiputini 5
Xyleborina Coptoborus sp. Onkone Gare/Tiputini 6
Xyleborina C optoborus sp. Onkone Care I
Xyleborina Coptoborus sp. Onkone Gare I
Xyleborina Coploborus sp. Onkone Gare l
Xyleborina C optoborus sp. Onkone Gare l
Xyleborina Coptoborus sp. Onkone Gare l
Xyleborina Coptoborus sp. Onkone Gare l
Xyleborina Coptoborus sp. Onkone Gare 3
Xyleborina C optoborus sp. Onkone Gare I
Xyleborina Coptoborus sp. Onkone Gare I
Xyleborina C optoborus sp. Onkone Gare I
Xyleborina C optoborus sp. Onkone Gare l
Xyleborina C optoborus sp. Tiputini I
Xyleborina Coptoborus sp. Onkone Gare |
Xyleborina Coptoborus sp. Onkone Gare I
Xyleborina C optoborus sp. Onkone Gare 2
Xyleborina C optoborus vespatorius Onkone Gare l
Xyleborina Dryocoetoides sp. Onkone Gare 6
Xyleborina Dryocoetoides sp. Onkone Gare l
Xyleborina Dlyocoetoides sp. Onkone Gare I
Xyleborina Dryocoetoia’es sp. Onkone Gare l
Xyleborina Daocoetoides sp. Onkone Gare 2

Table 3.3 (cont. on next page): Morphospecies of ambrosia beetles (Xyleborina +
Premnobina) found in the canopy fogging samples.

276

 

 

 

Subtribe Genus Species Locality(ies) No. Specimens
Xyleborina Theoborus sp. Tiputini I
Xyleborina Theoborus sp. Tiputini 3
Xyleborina Theoborus sp. Onkone Gare/Tiputini 5
Xyleborina Theoborus nr. micarius Onkone Gare I
Xyleborina Theoborus sp. Onkone Gare 3
Xyleborina Theoborus sp. Tiputini I
Xyleborina Theoborus sp. Onkone Gare/Tiputini 8
Xyleborina Theoborus sp. Tiputini l
Xyleborina Theoborus sp. Onkone Gare I
Xyleborina Theoborus sp. Onkone Gare I
Xyleborina Theoborus sp. Onkone Gare l
Xyleborina X yleborinus sp. Tiputini 2
Xyleborina Xyleborus sp. Tiputini 2 I
Xyleborina X yleborus .spathipenm’s Onkone Gare/Tiputini 2
Xyleborina X yleborus aflim‘s Onkone Gare/Tiputini 3 I 7
Xyleborina X yleborus sp. Tiputini I
Xyleborina X yleborus nr. ferrugineus Onkone Gare/Tiputini l2
Xyleborina X yleborus sp. Onkone Gare 2
Xyleborina Xyleborus sp. Tiputini 8
Xyleborina X yleborus sp. Tiputini I
Xyleborina Xyleborus sp. Onkone Gare I
Xyleborina X y/eborus sp. Tiputini l
Xyleborina X yleborus sp. Tiputini I
Xyleborina Xyleborus sp. Onkone Gare 1
Xyleborina , yleborus sp. Onkone Gare 2
Xyleborina Xyleborus sp. Tiputini l
Xyleborina Xyleborus sp. Onkone Gare I
Xyleborina Xyleborus sp. Onkone Gare l
Xyleborina Xyleborus sp. Onkone Gare 2
Xyleborina Xyleborus sp. Onkone Gare I
Xyleborina Xyleborus sp. Onkone Gare l
Xyleborina X yleborus sp. Onkone Gare I
Xyleborina Xyleborus sp. Onkone Gare I
Xyleborina Xyleborus sp. Onkone Gare l
Xyleborina Xylosandrus morigerus Onkone Gare/Tiputini I2
Premnobina Premnobius clavigennis Onkone Gare I

 

 

Table 3.3 (cont): Morphospecies of ambrosia beetles (Xyleborina + Premnobina) found
in the canopy fogging samples.

277

 

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