Systematics of the genus Trachyphloeomimus Champion (Coleoptera: ) with a revision of the T. championi species group

Pamela J. Horsley

Department of Natural Resource Sciences McGill University, Montreal

August 2009

A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Master of Science

© Pamela J. Horsley 2009

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TABLE OF CONTENTS

List of Tables……………………………………………………………………..iv List of Figures……………………………………………………………………..v List of Appendices……………………………………………………………….vii Acknowledgements……………………………………………………………..viii Abstract…………………………………………………………………………....x Résumé……………………………………………………………………………xi

1. GENERAL INTRODUCTION AND LITERATURE REVIEW………………………..1 1.1 Subfamily ………………………………………………………….2 1.2 Tribe ………………………………………………………….3 1.2.1 Taxonomic History………………………………………………….3 1.3 Genus Trachyphloeomimus Champion………………………………………..4 1.3.1 Taxonomic History………………………………………………….4 1.3.2 Natural History…………………………………….………………...5 1.3.3 Distribution………………………………………………………….5 1.4 Objectives……………………………………………………………………..6

2. MATERIALS AND METHODS……………………………………………………7 2.1 Material Examined…………………………………………………………….7 2.2 Specimen Preparation and Examination………………………………………7 2.3 Descriptive Format…………………………………………………………….8 2.4 Structural Terminology………………………………………………………..8 2.5 Phylogenetic Methods…………..……………………………………………..9 2.5.1 Analysis………………………………………………….…………..9 2.5.2 Outgroups……………………………………………….………….10

3.RESULTS…………………………………….………………………………….11 3.1 Taxonomic Treatment…………..…………….……………………………...11 3.1.1 Tribe Trachyphloeini Lacordaire…………………………………..11 3.1.2 Genus Trachyphloeomimus Champion...…………………..………11

ii 3.1.3 Trachyphloeomimus championi species group…………………….14 3.1.4 Key to species of the T. championi species group…..……………..15 3.1.5 Species descriptions of the T. championi group……….…………..18

T. apionus Horsley, NEW SPECIES……………………………...…18

T. arcuatibius Horsley, NEW SPECIES ..…………………………..20

T. batmobile Horsley, NEW SPECIES ………………………….….22

T. bravoensis Horsley, NEW SPECIES ………..…………………...24

T. calcaritibius Horsley, NEW SPECIES ……..….………………...26

T. carinatus Horsley, NEW SPECIES …………..………………….28 T. championi O’Brien………………….….…………….……….31

T. discus Horsley, NEW SPECIES …………….………………..….33

T. festivipenis Horsley, NEW SPECIES …………….…………..….35

T. heliconodulus Horsley, NEW SPECIES ………….….……….….38

T. jonesi Horsley, NEW SPECIES ……………………..….……..…40

T. junctus Horsley, NEW SPECIES ……………………..……….....42

T. obrieni Horsley, NEW SPECIES ……….……………..……...….44

T. popeyeus Horsley, NEW SPECIES …………………..………..…46 T. spurcus Champion……………………...……………………..49 T. undulatoides O’Brien………………………………………….52 3.2 Phylogenetic Analysis………………………………………………………..54

4. DISCUSSION AND CONCLUSION……………………………………………….56

REFERENCES……………………………………………….…………………….58

TABLES………………………………………………………………………...…61

FIGURES………………………………………………………………………….63

APPENDICES...………………………………………………………….………...80

iii LIST OF TABLES

Table 1. Data matrix used in the phylogenetic analysis of Trachyphloeomimus species and related taxa. Box indicates the species included in the T. championi species group.

iv LIST OF FIGURES

FIGURES 1-6. Dorsal (left) and lateral (right) habitus of the Trachyphloeomimus championi species group: (1-2) T. apionus, female; (3-4) T. arcuatibius, male; (5- 6) T. batmobile, female.

FIGURES 7-12. Dorsal (left) and lateral (right) habitus of the Trachyphloeomimus championi species group: (7-8) T. bravoensis, male; (9-10) T. calcaritibius, male; (11-12) T. carinatus, male.

FIGURES 13-18. Dorsal (left) and lateral (right) habitus of the Trachyphloeomimus championi species group: (13-14) T. championi, male; (15-16) T. discus, female; (17-18) T. festivipenis, male.

FIGURES 19-24. Dorsal (left) and lateral (right) habitus of the Trachyphloeomimus championi species group: (19-20) T. heliconodulus, female; (21-22) T. jonesi, female; (23-24) T. junctus, female.

FIGURES 25-32. Dorsal (left) and lateral (right) habitus of the Trachyphloeomimus championi species group: (25-26) T. obrieni, male; (27-28) T. popeyeus, male; (29-30) T. spurcus, female; (31-32) T. undulatoides, female.

FIGURES 33-36. Ventral (left) and lateral (right) view of the male genitalia of the Trachyphloeomimus championi species group (Scale bar measurements in parentheses): (33) T. apionus (0.2 mm); (34) T. arcuatibius (0.5 mm); (35) T. batmobile (0.54 mm); (36) T. bravoensis (0.54 mm).

FIGURES 37-40 Ventral (left) and lateral (right) view of the male genitalia of the Trachyphloeomimus championi species group (Scale bar measurements in parentheses): (37) T. calcaritibius (0.38 mm); (38) T. carinatus (0.42 mm); (39) T. championi (0.52 mm); (40) T. festivipenis (0.32 mm).

v FIGURES 41-43. Ventral (left) and lateral (right) view of the male genitalia of the Trachyphloeomimus championi species group (Scale bar measurements in parentheses): (41) T. jonesi (0.38 mm); (42) T. obrieni (0.38 mm); (43) T. popeyeus (0.4 mm).

FIGURES 44-59. Female spermatheca of the Trachyphloeomimus championi species group: (44) T. apionus; (45) T. arcuatibius; (46) T. batmobile; (47) T. bravoensis; (48) T. calcaritibius; (49) T. carinatus; (50) T. championi; (51) T. discus; (52) T. festivipenis; (53) T. heliconodulus; (54) T. jonesi; (55) T. junctus; (56) T. obrieni; (57) T. popeyeus; (58) T. spurcus; (59) T. undulatoides.

FIGURES 60-64. Pronotum (dorsal view) of females in the Trachyphloeomimus championi species group: (60) T. arcuatibius; (61) T. carinatus; (62) T. discus; (63) T. heliconodulus; (64) T. spurcus.

FIGURES 65-67. Hind tibia (lateral view) of males in the Trachyphloeomimus championi species group: (65) T. calcaritibius; (66) T. festivipenis; (67) T. obrieni.

FIGURE 68. Phylogenetic relationships of species of Trachyphloeomimus: Strict consensus tree of 1000 most parsimonious trees of length 368 for 34 unweighted characters. Arrows show placement of species within the T. championi species group.

FIGURE 69. Phylogenetic relationships of species of Trachyphloeomimus: Strict consensus tree of 1000 most parsimonious trees of length 385 for 34 weighted characters. Arrows show placement of species within the T. championi species group. Note shift in placement of T. spurcus and T. jonesi species.

vi LIST OF APPENDICES

Appendix I. Checklist of the species included in the T. championi species group.

Appendix II. Characters and character states used in the phylogenetic analysis of Trachyphloeomimus species and related taxa.

vii ACKNOWLEDGEMENTS

First and foremost, I would like to gratefully acknowledge the following people and institutions for the loan of Trachyphloeomimus specimens: Francois Génier (CMNC), Charles O'Brien (CWOB), Jorge León (ECOS), Richard Hoebeke (CUIC), Robert Jones (UAQM), Roy Danielsson (MZLU), Chris Lyal (BMNH). Without their cooperation this research could not have been completed. I would like to thank Nadine Duperre for graciously and patiently providing the male genitalia sketches. Thank you to Robert Jones, Moises Perez and Jesus Luna Cozar (Chuy) for showing me a great time in Mexico. I would also like to thank Francois Génier for teaching me the wonderful ways of Mantis and how to use it to its’ fullest potential. Your determination, enthusiasm and uncanny attention to detail have taught me how to properly take care of natural history collections. I always know that if there is a problem or a more efficient way to do something, you will find the solution to it, even if it means you have to construct it yourself. I owe Andrew Smith more than the few words I will say here. He has probably the biggest heart, as well as, the most enthusiasm for all things entomology, of anyone I know. I am extremely grateful to have been able to work with and for him these past few years. Thank you for always taking the time to answer my questions and teach me useful things, especially the importance of parentheses. I would like to thank my family for always supporting me and always offering to help in any way they could. My M.Sc. could not have started at a worse time in my life, but it made me realize that nothing should be taken for granted and to live your life the way you want to. I’m happy and relieved you will see me get through this Mom. My life improved drastically when I met my boyfriend Ralph, who has been through it all with me and who I thank the most for his support and love. Last, but not least, I would like to thank my supervisors Terry A. Wheeler and Robert S. Anderson for their guidance, support and patience during the course

viii of this study. Thank you Terry for your reassurance at the start and keeping me in check at the end. It has been an adventure and a privilege working with you Bob. Your knowledge and love of weevils, dream of collecting every leaf litter weevil there is and belief in others has inspired me and helped me immensely. I could not have asked for a better thesis supervisor and mentor. Financial aid for this research was provided by the Natural Sciences and Engineering Research Council of Canada (NSERC Discovery Grants) held by supervisors Terry A. Wheeler and Robert S. Anderson.

ix ABSTRACT

The genus Trachyphloeomimus is redescribed and the T. championi species group is revised. The T. championi group includes three previously described species: T. championi O’Brien, T. spurcus Champion, T. undulatoides O’Brien; and thirteen new species: T. apionus, T. arcuatibius, T. batmobile, T. bravoensis, T. calcaritibius, T. carinatus, T. discus, T. festivipenis, T. heliconodulus, T. jonesi, T. junctus, T. obrieni, T. popeyeus. Relationships among these species are assessed and discussed and the monophyly of the species group is demonstrated. A key to species in the T. championi group and illustrations of species-group level defining characters are provided. Species in the T. championi group are restricted to Mexico, largely in the state of Oaxaca, but the distribution of the genus now includes Mexico, Guatemala, Costa Rica, Nicaragua, Honduras and El Salvador. Six of the sixteen species in the T. championi group are known only from females.

x RÉSUMÉ

Le genre Trachyphloeomimus est redécrit et le groupe d'espèces de T. championi est mis à jour. Le groupe de T. championi inclut trois espèces précédemment décrites: T. championi O'Brien, T. spurcus Champion, T. undulatoides O'Brien; et treize nouvelles espèces: T. apionus, T. arcuatibius, T. batmobile, T. bravoensis, T. calcaritibius, T. carinatus, T. discus, T. festivipenis, T. heliconodulus, T. jonesi, T. junctus, T. obrieni, T. popeyeus. Les liens entre ces espèces sont évalués et discuté et la monophylie du groupe d'espèces est démontrée. Une clé pour les espèces dans le groupe de T. championi et des illustrations des caractères qui définissent le niveau d'espèce-groupe sont fournies. Des espèces dans le groupe de T. championi sont limitées au Mexique, en grande partie dans l'état d'Oaxaca, mais la distribution du genre inclut maintenant le Mexique, le Guatemala, le Costa Rica, le Nicaragua, le Honduras et le Salvador. Six des seize espèces dans le groupe de T. championi sont connus seulement au niveau des femelles.

xi 1. GENERAL INTRODUCTION AND LITERATURE REVIEW

With over 50,000 species in approximately 4500 genera currently described world wide, the Curculionidae is the largest family of organisms (Anderson 2002). Recognized as one of the most difficult groups systematically, their megadiversity and the critical lack of taxonomic specialists make the Curculionidae a challenging group to study. Most weevils are easily recognizable by the presence of an elongate rostrum (snout) with mouthparts situated at the apex and geniculate antennae with a compact club (Anderson 2002). Some subfamilies, namely the Entiminae (broad-nosed weevils), possess a poorly developed rostrum, while the Platypodinae and Scolytinae possess a head that is scarcely prolonged (Anderson 2002). Weevils are generally phytophagous as adults and larvae and those that feed on living plants can loosely be divided into two categories according to feeding habits: the subfamily Entiminae have polyphagous adults and larvae and the remaining subfamilies have oligophagous adults and larvae restricted to a much narrower host range of plants (Anderson 2002). Many weevils are also saprophagous and play an important role in the breakdown of dead plant material in tropical forests. Due to the variety of feeding habits seen in weevils, many are considered economically important pests of ornamental, agricultural and forestry plants and stored grains (Anderson 2002). In contrast, weevils have recently been tested and used successfully in the biological control of pest and invasive species of plants, such as Eurasian watermilfoil (Myriophyllum spicatum L.) and spotted knapweed (Centaurea maculosa Lam.) (Anderson 2002). The Curculionidae is one of the 7 to 18 families, depending on the classification scheme used, within the superfamily Curculionidoidea. According to the most current phylogenetic concepts, as reviewed by Oberprieler et al (2007), the superfamily is divided into seven families: Nemonychidae, Anthribidae, Belidae, Attelabidae, Caridae, Brentidae and Curculionidae. The Curculionidae or true weevils is further divided into 10 to 16 subfamilies. The classification of the Curculionidae into natural subfamilies and tribes, as stated by Oberprieler et al. (2007) is: “the largest outstanding problem in the higher classification

1 of Coleoptera due to the diversity of the group, that is challenged perhaps only by the Staphylinidae (rove )”.

1.1 Subfamily Entiminae The Entiminae, or broad-nosed weevils, is the largest subfamily of weevils with over 12,000 species in approximately 1000 genera (Oberprieler et al. 2007; Anderson and Oberprieler, in press). According to Alonso-Zarazaga and Lyal (1999), the subfamily is currently divided into 55 tribes worldwide. The closest subfamily in terms of species diversity would appear to be the Cryptorhynchinae with slightly less than 7000 world species in almost 700 genera (O’Brien and Wibmer 1978). Entiminae are characterized by a short, stout rostrum that is reduced in length and broader than the rostrum of most other weevils and possesses adelognathous mouthparts (the prementum closing the buccal cavity from beneath) (Oberprieler et al. 2007). They generally do not display sexual dimorphism of the rostrum length used for endophytic oviposition, as is seen in other Curculionidae, but the sexes can usually be distinguished from one another by the overall greater robustness of the female and the concavity of abdominal ventrites 1 and 2 in the male. An important character that most entimines display is the presence of a usually well-defined scar on the apex of the mandible. This scar is left after a specialized deciduous process of the mandible, used by the adult as it splits open the pupal casing, breaks off (Anderson 2002). In a phylogenetic study using mainly larval characters coded for selected subfamilies of the Curculionidae and tribes within the Entiminae, Marvaldi (1997) aimed to test the relationships among the weevils, in particular the broad-nosed weevils. The hypothesis of monophyly of broad-nosed weevils was not accepted by Marvaldi because they constitute at least four monophyletic groups: Ithycerinae, Microcerinae, Brachycerinae sensu stricto and the “Entiminae and allied subfamilies” group, therefore, the change to ectophytic oviposition without the aid of the rostrum has occurred multiple times in the evolutionary history of the Curculionidae. The Entiminae was distinguished from “allied subfamilies” by two larval characters: the shape of the antennal sensorium wider than long, dorsoventrally compressed and cushion-like in form; and the maxillary

2 mala with four ventral setae. The cushion-like antennal sensorium appears to be a good synapomorphy for the entimines (Oberprieler et al. 2007). The Entiminae constitute the majority of the weevil fauna in deserts, mountains and cold climates (Kuschel 1995). Most species are apterous with fused elytra that may prevent dessication in these harsh climates and due to the loss of dispersal powers, could account for the prominence of allopatric speciation at high elevations. Larvae generally feed on plant roots in the soil, while adults feed externally on plant foliage or reproductive structures (Anderson, 2002). Entimines are quite conspicuous in terms of their pest status, generally targeting ornamental plants, as well as crops, particularly citrus and other fruits (Anderson, 2002). Some of the most common entimine pests include Otiorhynchus ovatus Linnaeus (strawberry root weevil) and Cyrtepistomus castaneus (Roelofs) (Asiatic oak weevil) (Anderson, 2002). The classification of Entiminae into natural tribes is chaotic, but currently 55 tribes are recognized worldwide according to Alonso-Zarazaga and Lyal (1999). Many of the tribes and subtribes included in this checklist are based on Palearctic genera and relationships to the southern-hemisphere forms are insufficiently reviewed and are therefore unclear (Oberprieler et al. 2007). In recent years, there has been some attempt to clarify the relationships among the tribes. Thompson (1992) and Marvaldi (1997, 1998) recognize a smaller number of tribes including: Alophini, Pachyrhynchini, Ectemnorhinini, Sitonini and Entimini. Although this is a step in the right direction, there is need to expand the study to include the entire diversity of the subfamily.

1.2 Tribe Trachyphloeini 1.2.1 Taxonomic History The name Trachyphloeini was first proposed by Lacordaire (1863) (as the Trachyphléides) for a group of five genera: (as Cerpoceus, lapsus), , Cathormiocerus, Scoliocerus and Anemophilus, all previously described. Sleeper (1955) published a review of the tribe in North America, which included descriptions of all North American genera and species known at the time. Characters defining the tribe, as described by Sleeper included: the absence of ocular lobes, eyes free and rounded; scrobe variable, never linear and directed beneath at the same time; outer

3 segments of funicle moniliform; elytra scarcely wider than prothorax at base; humeri rounded; metasternum short; intercoxal lobe of first abdominal segment large and broad; corbels of posterior tibiae open, tarsal claws free; apterous. Revisions are needed at all taxonomic levels in order to better resolve the placement and evolutionary relationships within the Trachyphloeini. Presently, the tribe is separated into two subtribes, the Trachyphloeina and Trachyphilina (Alonso-Zarazaga and Lyal 1999). Trachyphilina contains only two genera, Trachyphilodes Voss and Trachyphilus Faust, the latter considered non- Trachyphloeini by Zherikhin and Egorov (1991), but no new placement has been proposed. The second subtribe, Trachyphloeina, is considerably larger with 21 genera recognized. Currently, nine genera are recognized as present in the New World, while the remaining thirteen genera are exclusively Old World. Of the New World genera, five of these are native and four have been introduced. The genera Trachyphloeus Germar and Roelofs have long been known in North America, whereas Trachyphloeosoma Wollaston and Cathormiocerus Schoenherr are each known from one recently recorded introduced species (Anderson 2002). Among the five native genera, Cercopeus Schoenherr, with 11 recognized species in the southeastern United States is the most diverse with the Mexican and Central American Trachyphloeomimus next with eight described species. The remaining genera Cercopedius Sleeper, Chaetichidius Sleeper and Pseudocercopeus Sleeper are monotypic. Adults of all native genera with the exception of Cercopedius, which is associated with Artemisia (sagebrush), are collected in leaf litter, ground debris and under rocks. The relationships and taxonomic status of these last three monotypic genera require reassessment.

1.3 Genus Trachyphloeomimus Champion 1.3.1 Taxonomic History The genus Trachyphloeomimus was described by Champion (1911). He described the type-species for the genus, T. spurcus, and also placed Trachyphloeus solitarius, previously described by Sharp (1911), within Trachyphleomimus. In the same publication, Champion described another new monotypic genus, Thamiras, based on a single specimen from the Los Altos region of Guatemala (Champion 1911). He clearly

4 described this new genus considering a close relationship with Trachyphloeomimus, as he states, “it is related to Trachyphloeomimus” (Champion 1911). Champion differentiated the two genera based on slight differences between segments 1 and 2 of the funicle, the form of the prothorax and the first abdominal suture. Further taxonomic work on Trachyphloeomimus was not carried out until nearly sixty years later, when O’Brien (1972) published a review of the genus. O’Brien concluded that Thamiras and Trachyphloeomimus were congeneric and placed the former as a junior synonym of the latter, based on page precedence. He described five new species (T. alternatus, T. championi, T. mexicanus, T. sharpi and T. undulatoides) in addition to the previously described three species. Trachyphloeomimus has received no taxonomic attention since O’Brien’s study. 1.3.2 Natural History Species of Trachyphloeomimus are most often collected using standard leaf litter sifting methods and berlese extraction (Martin 1977, Anderson and Ashe 2000). Recent collections using Winkler bags have also been successful in producing samples of Trachyphloeomimus specimens. They are generally found at higher elevations (1000- 3700m) in oak/pine and cloud forests, but have been found under rocks and other ground debris in high elevation meadows. It is likely that adults feed on plant foliage and larvae feed on roots, as is typical for most species of Entiminae (Anderson 2002); however, adults have not been recorded on plants and no feeding observations are known. Larval habits of Trachyphloeomimus are mentioned in only one ecological study. Morón-Ríos et al. (1997) found that larvae of a Trachyphloeomimus species they called “T. aff. Spurcus” was second highest in terms of macroarthropod root-feeding density in subalpine grasslands in Mexico, slightly less than species of Melolonthidae (Coleoptera). Other than this record and unpublished specimen label data, not much is known about Trachyphloeomimus natural history. 1.3.4 Distribution Based upon the eight described species, the distribution of Trachyphloeomimus has been historically recorded as Mexico and Guatemala. Since the last revision by O’Brien (1972), many individuals in the genus have been collected during numerous expeditions into the mountains of Central America. Although no details on numbers of

5 species collected or distribution are given, Anderson and Ashe (2000) recorded Trachyphloeomimus from Honduras and El Salvador in their study of diversity patterns of leaf litter weevils and Staphylinidae. This increase in recent collecting efforts has extended the geographic range of Trachyphloeomimus significantly. It now includes northeastern Mexico, south through Guatemala, El Salvador, Honduras, Nicaragua and into northern Costa Rica and the number of undescribed species in the genus has increased considerably.

1.4 Objectives The genus Trachyphloeomimus is a speciose and complex genus of broad-nosed weevils inhabiting Mexico and Central America. At present, 8 species are described but over 70 additional species have been recognized in a preliminary assessment of the diversity in the genus (Horsley, unpublished data). These species can be grouped into a number of distinct species groups based on external morphological characters and characters of the genitalia. The main objective of this thesis is to provide a description, assessment of monophyly, and species-level phylogeny of one of these species groups, the T. championi species group. The T. championi species group is formally described and includes 16 species, 3 of which are redescribed, along with descriptions of 13 new species. Relationships among these species are assessed and the monophyly of the species group is demonstrated. A dichotomous key to species in the T. championi group and illustrations of species group level defining characters are provided. This thesis will provide a solid framework for future revisionary work on the remaining species groups within the genus and further emphasizes our lack of basic knowledge of diversity in many taxa.

6 2. MATERIALS AND METHODS

2.1 Material Examined

Descriptions were based on the examination of more than 3500 adult specimens of the genus Trachyphloeomimus. Specimens have been examined from and are deposited in the following private and institutional collections (curators in parentheses). Collection acronyms follow the Bishop Museum’s website (http://hbs.bishopmuseum.org/codens/ codens-inst.html). BMNH The Natural History Museum, London, England (Chris Lyal) CMNC Canadian Museum of Nature, Ottawa, Canada (François Génier) CUIC Cornell University Insect Collection, Cornell University, Ithaca, NY, U.S.A. (E. Richard Hoebeke) CWOB Charles W. O'Brien personal collection, Florida A&M University, Tallahassee, FL, U.S.A. (Charles W. O'Brien) ECOS (formerly CIES) El Colegio de la Frontera Sur (Ecosur), Museo de Entomologia, San Cristóbal de las Casas, Chiapas, México (Jorge L. León Cortes) MZLU Zoological Institute, Lund University, Lund, Sweden (Roy Danielsson). UAQM Universidad Nacional Autonoma de Mexico (Querétaro), Querétaro, Mexico (Robert Jones)

2.2 Specimen Preparation and Examination

Point- and card- mounted specimens were examined using a Leica MZ6 dissecting microscope. All specimens received in alcohol were cleaned using an ultrasonic cleaner, prior to pointing and labelling. For each species, where possible, multiple males and females were dissected. Dissections accomplished following Martin (1977). All dissected parts were placed in microvials with glycerine. Adult specimens were re-pointed and re-labelled accordingly with the appropriate microvial pinned below.

7 Habitus measurements for descriptions were determined using an ocular micrometer on a WILD M3Z dissecting microscope. All digital photographs were taken using a Nikon digital camera, model DMX1200F. Female spermathecae were first digitally photographed, then using Adobe Illustrator were traced to better show the form of the structure. Touch ups to digital photographs and illustrations and plate construction were accomplished using Adobe Photoshop and Adobe Illustrator. All specimens have been databased using Mantis (v2.0.1) as part of the World Weevil Database (WWD). Labels containing the specimen-specific number have been placed below all other labels for easy identification.

2.3 Descriptive format

Species groups are used rather than subgenera but otherwise the formal splitting of the genus into a number of different named entities has been avoided. The name T. championi for the species groups was chosen arbitrarily. Structural characteristics of the species group are given under the diagnosis of the group and are not repeated for each species within the group. A diagnosis of each species is presented which summarizes the key character states that allow for recognition of that species. Full length descriptions follow thereafter providing more detail for identification purposes. All material examined has been indicated with full citation of label data, notation of sex and depository of the specimens, as outputted from Mantis. Holotypes are designated and label data is copied verbatim to decrease chances of future problems in recognition.

2.4 Structural Terminology

The following definitions and standards were used in the descriptions and diagnoses: Colour and Integument based on dried, pinned specimens. Body length was measured dorsally along the midline, from the apical margin of the pronotum to the elytral apex to avoid inaccurate measurement due to the deflection of the head and rostrum. Body width was measured at the widest point, typically just below the humeri

8 where interval 8 of the elytra is enlarged. Bevel refers to the apex of the tibia following Thompson (1992) and his clarification of terminology to be used in weevil nomenclature. Nasal plate is the area extended from the apical end of the antennal insertion to the apex of the rostrum (Bright and Bouchard 2008). The flagellum is the long slender process at the apex of the internal sac of the male aedeagus (Howden 1982) and the apical sclerite complex is the group of sclerites situated towards the apex of the internal sac (Anderson 1987)

2.5 Phylogenetic Methods 2.5.1 Analysis The phylogenetic analysis was conducted using TNT (version 1.1) with a matrix containing 81 taxa (3 taxa as outgroups, 78 taxa in the ingroup) (Table 1) and 34 multi- state characters (Appendix II). A traditional heuristic search with standard settings of Wagner trees using random seed and ten random addition sequences was executed. The tree bisection reconnection (TBR) swapping algorithm with one thousand trees saved per replication was also employed. A consensus tree was computed to determine the strength of the groupings. Following the heuristic search, character weighting was used for subsequent analyses. A simple two-level weighting system was used to distinguish between the weighted characters. Primary characters were used to establish the basic set of a priori relationships among taxa and the secondary characters were then used to reinforce the previously established relationships or to further resolve the reconstructed phylogeny by providing additional characters. Two characters were deemed primary characters and were given higher relative weight as the determined apomorphic state has not been observed elsewhere in Entiminae. The first of these characters is a modification of the basal margin of the pronotum, resulting in either a sub-glabrous half-circle or raised, distinctly triangular shaped area (Figs. 60-64). This character is exhibited on both males and females, but is generally much more evident on females. This character does not appear to be known elsewhere in Entiminae. The second character is a modification of the hind tibia of males (Figs. 65-67). This area is generally more or less straight from proximal to distal ends, but in some species it has been modified to be weakly to strongly

9 arcuate or bearing a distinct tooth or flange. As with the pronotal character, this character also does not appear to be known elsewhere in Entiminae. The remaining characters are considered secondary characters, subject to homoplasy and are accorded lesser weight. In general, these characters are those that are known to have the apomorphic state occurring elsewhere (often widely) within the Entiminae. Following weighting, the matrix was then re-analyzed with a traditional heuristic search with the same options as previously used. A consensus tree was computed allowing for the comparison of groupings that resulted from the two different searches and the analysis of resulting resolution. 2.5.2 Outgroups Upon recommendation from Charles O’Brien, two species of the southern USA leaf litter-inhabiting genus Cercopeus were used as outgroups. C. komareki is representative of the bisexual species, while C. chrysorrhoeus is representative of the parthenogenic species that occur within the genus. Cercopeus is placed in the same tribe as the genus of study, Trachyphloeini, and is believed to be the sister group to Trachyphloeomimus as they exhibit allopatric distributions but maintain similar life habits. One other taxon was also selected for use as an outgroup in order to represent a distinctly different tribe within the diverse Entiminae. Sciomias latipennis Sharp of the tribe Polydrusini was randomly selected to represent the genus, as a type species has yet to be designated.

10 3. RESULTS

3.1 Taxonomic Treatment

3.1.1 Tribe Trachyphloeini Lacordaire

In his review of the tribe Trachyphloeini of America north of Mexico, Sleeper (1955), following Lacordaire (1863), characterized the tribe as lacking postocular lobes, eyes free and rounded, scrobes variable but not linear and directed beneath the eyes, apical articles of funicle moniliform, scutellum small or absent, elytra slightly wider than width of pronotum at base, intercoxal lobe of first ventrite large and broad, ventrite 2 as long as each of 3 and 4, hind tibial corbels open, tarsal claws free. Most of the taxa are from Europe and when the tribe was established by Lacordaire he considered only European taxa. Sleeper (1955) recognizes six genera as present in North America, of which, five are native, the genus Trachyphloeus being introduced. Trachyphloeomimus is the only Neotropical genus placed in the tribe and its allopatric distribution, similar habits, and structural similarity to Cercopeus, suggests these two taxa likely shared a relatively recent common ancestry. The definition and composition of the tribe are in need of revision, as is the situation for most tribes of broad-nosed weevils. At present, there is no evidence for the monophyly of the tribe and it may not constitute a natural group.

3.1.2 Genus Trachyphloeomimus Champion

Trachyphloeomimus Champion 1911:342. Gender masculine. Type-species Trachyphloeomimus spurcus Champion 1911:342 by original designation. O’Brien 1972 (revision). Lona 1937:345,347 (checklist). Blackwelder 1944:801 (checklist). O'Brien and Wibmer 1982:53 (checklist). Alonso-Zarazaga and Lyal 1999:183 (checklist).

Thamiras Champion 1911:343. Type species Thamiras undulatus Champion 1911:343 by monotypy. O’Brien 1972:165 (synonymy with Trachyphloeomimus).

11

Trachyphloeus Sharp 1911:177 (misidentification).

Diagnosis. Rostrum stout, as long as head or longer, convex in lateral view, straight to wider at apex; epistoma not prominent to strongly emarginated, triangular; scrobes visible in dorsal view, descended and directed at lower anterior margin of eyes; eyes oval, moderately large, coarsely faceted, each with multiple erect setae above eye; antennal scape clavate, most often reaching anterior margin of pronotum, densely clothed with appressed scales and erect hair-like setae; funicle of 7 articles, articles 1 and 2 elongate, 3 and 4 slightly longer than broad, 5 and 6 submoniliform, 7 submoniliform to wider than long; club stout, ovate. Elytra most often tuberculate, especially at declivity on interval 5, tubercles occasionally weakly formed or lacking. Femora disctinctly clavate, inner margins simple. Dorsal vestiture dense, brown to copper-red or shiny metallic, scales rounded and indistinct or striate to stellate in form, dorsal surface often encrusted with dirt. Spermatheca with ramus and nodulus of variable relative lengths. Aedeagus variable in form.

Description. Size moderate (length 2.1–6.9mm; width 1.0–3.0mm).

Colour and integument generally reddish-brown to black. Scale colour generally brown or beige, often dirt covered, occasionally metallic in sheen; scale type variable, most often oval striate, few stellate, occasionally difficult to distinguish. Setae most often sub- erect; colour generally dark brown and beige, often golden on legs and on venter of body; form ranging from straight to distinctly clavate, especially along declivity of elytra.

Rostrum not sexually dimorphic. In lateral view more or less straight to moderately arcuate from base to apex, profile generally straight, sometimes with distinct ridge at point of antennal insertion; epistoma not prominent to strongly emarginate, triangular. In dorsal view, more or less straight to distinctly widened at apex; scrobe deep, glabrous,

12 clearly visible, lateral, descending posteriorly to lower anterior margin of eye; surface foveate or sulcate and squamose, generally ending at point of antennal insertion.

Antennae with scape clavate, slender to robust, generally extended to anterior margin of pronotum; with sparse to dense scales and straight, fine to spatulate setae. Funicle with 7 articles, articles 1 and 2 elongate, article 1 generally 1/4 to 1/3 longer than article 2, articles 3 and 4 slightly longer than broad, articles 5 and 6 submoniliform, article 7 submoniliform to distinctly wider than long. Club oval to elongate-oval, with short fine golden setae.

Head round, squamose. Eyes present, moderately large and convex, distinctly inset, oval, situated dorso-lateral on rostrum at base, coarsely facetted; multiple erect setae above eyes, often on distinct tubercle (eyebrow-like) above to eye.

Pronotum variously and irregularly punctate, occasionally irregularly papillate, each papilla bearing one seta. Scales of various form and size. In dorsal view with shape various; apex often sub-tubulate, constricted laterally and dorsally to various extent, occasionally rounded; basal margin modified medially in some with sub-glabrous triangular or semi-circular area, often raised; widest at mid-length to apical 1/3, width relative to elytra ranges from smaller to wider.

Scutellum small to minute.

Elytra fused, elongate oval to sub-rectangular in dorsal view; humeri variable, rounded to strongly produced anteriorly; interval 8 not evident to distinctly enlarged below humeri; vestiture of various form, size and extent, tuberculate in most species (markedly so in some). Apterous.

Legs with femora distinctly clavate. Tibiae more or less straight to slightly bowed inward in some species, uncinate; corbel beveled. Hind tibiae sexually dimorphic in some species; in male inner margin nearly straight to markedly arcuate or with distinct flange

13 or tooth, with setae short to very long; in female generally straight, occasionally with row of distinct, sharp, tooth-like setae. Apical comb with reddish-brown spines of various lengths, but generally short. Tarsi of 5 articles; article 3 broadly bilobed; article 4 greatly reduced and between lobes of article 3; article 5 elongate and bearing tarsal claw; claws long, simple, free and widely divergent. Ventral pilosity dense.

Abdomen with 5 visible ventrites. Ventrites 1 and 2 fused; ventrites 3, 4 and 5 free. Ventrite 5 variously punctate. Pygidium in female generally unmodified, occasionally sinuate or with medial emargination.

Genitalia. Male with aedeagus with body moderate in size to distinctly robust dorsoventrally; apex of body variable, generally acuminate to some extent; internal sac simple with few sclerotized teeth to very complex with many sclerotized teeth, often arranged in clusters, sclerotized teeth of internal sac variable in size and form; flagellum variable in form, most often with apex curving up. Female with spermatheca variable in terms of relative lengths of ramus and nodulus.

3.1.3 Trachyphloeomimus championi species group

Diagnosis. This species group can be easily distinguished by the presence of a generally distinct sub-glabrous triangular or semi-circular shaped area at the middle of the posterior margin of the pronotum. This structure is generally more pronounced in females but in some specimens is smaller or is obscured by scales or dirt in males. Some species have the area distinctly elevated, while on others, it appears simply that scales are absent. Males in this species group have the inner margin of all legs with long fine golden setae. The inner margin of the hind tibia in males also varies from weakly sinuate to having a distinct flange or tooth. The inner face of the hind tibia in females is weakly to strongly dentate. The male aedeagus has a flagellum ranging from weakly expanded at the apex to distinctly expanded and sclerotized along the outer margins at the apex. The internal sac varies, but generally contains clusters of variably sized teeth throughout.

14 Description. Length 2.5-6.9mm. Width 1.4-3.5mm. Antennae with article 7 of funicle as wide as or wider than long (3 species) or moniliform (13 species). Pronotum with middle of posterior margin modified with sub-glabrous wide, low semi-circular area (6 species) or thin, distinctly raised triangular area (8 species) or not modified (2 species). Elytra with very weak tubercles on interval 5 (2 species) or distinct tubercles on interval 5 (14 species); interval 8 not noticeably expanded behind humeri (2 species) or weakly expanded (11 species) or distinctly, broadly expanded (3 species). Legs of male with setae long (16 species); hind tibia of male weakly sinuate or arcuate (2 species) or with basal large triangular swelling (1 species) or modified with flange with widest point at or near mid-point (6 species) or near apical end (1 species); hind tibia of female with denticles (16 species). Male with aedeagus with flagellum distinctly expanded at apex (7 species) or weakly expanded at apex (3 species) or with extremely thin portion (1 species).

3.1.4 Key to species of the T. championi species group

1 Elytra with tubercles weakly to not developed on interval 5 at declivity ………...... 2 1’ Elytra with tubercles moderately to well developed on interval 5 at declivity ……..4

2 Elytra with intervals with indistinct, short fine acuminate setae

………………………………………………………….……T. jonesi NEW SPECIES 2’ Elytra with intervals with distinct, longer, clavate setae ………………….….….…3

3 Humeri distinctly rounded; setae on most intervals ..…T. spurcus Champion, 1911 3’ Humeri subrectangular; setae primarily on odd intervals

…………………………………………………………T. festivipenis NEW SPECIES

4 Elytra with tubercles at declivity on each of intervals 3 and 5 (in some specimens tubercles may be extended as an anteriorly directed ridge along the interval)

……….…………...……………...………………………T. carinatus NEW SPECIES 4’ Elytra with tubercles at declivity on interval 5 only ……………………………..…5

15

5 Elytra with tubercles at declivity on interval 5 extended anteriorly as a distinct ridge

(giving specimens a ‘winged’ appearance) ………..…...T. batmobile NEW SPECIES 5’ Elytra with tubercles at declivity on interval 5 more conical in form, not extended anteriorly as ridges …………...…………..……………………………………...…6

6 Elytra with tubercles at declivity on interval 5 extended medially, interrupting intervals 4 and 3 (making it appear as if intervals 5 and 3 are convergent at

declivity) ……………………………………………..……T. junctus NEW SPECIES 6’ Elytra with tubercles at declivity on interval 5 not extended medially …………..…7

7 Scales of pronotum and elytra distinctly striate ……………………………….....…8 7’ Scales of pronotum and elytra not striate or individual scales obscured by dirt and debris and hard to see …………………………...…………………………………10

8 Male with tibiae and femora with long dense hairs; body size larger (4.8-6.9mm) …………….…………………………………………...T. championi O’Brien, 1972 8’ Male with tibiae and femora not distinctly hairy; body size smaller (3.5-4.7mm) …9

9 Male with hind tibia with inner margin keeled, uniformly arcuate throughout length; female with basal margin of pronotum at middle with broadly triangular glabrous

area ………………………………………………………T. popeyeus NEW SPECIES 9’ Male with hind tibia with inner margin with distinct blunt tooth just basal to mid- length; female with basal margin of pronotum at middle with narrowly semicircular

glabrous area …………………………………………T. calcaritibius NEW SPECIES

10 Antennae with last article of funicle as wide as long or wider than long; elytra with indistinct arched scales along length of intervals or scales lacking …………...…..11 10’ Antennae with last article of funicle longer than wide; elytra with distinct suberect to erect scales along length of intervals ……………………………………...…....13

16 11 Size small (2.5-2.7mm), body form rounded and robust; elytra with tubercles at declivity very widely spaced, interval 5 appearing divergent apically

……………………..…………………………………...…T. apionus NEW SPECIES 11’ Size larger (3.3-6.5mm), body form slightly more elongate; elytra with tubercles at declivity widely spaced but interval 5 appearing subparallel throughout length ….12

12 Female with pronotum at middle of base with broadly triangular glabrous area

……………………………………………………………....T. discus NEW SPECIES 12’ Female with pronotum at middle of base lacking glabrous area ………………...………………………………….....T. undulatoides O’Brien, 1972

13 Male with inner margin of hind tibia strongly arcuate, keeled and distinctly widest at about midlength; female with hind tibia less strongly arcuate, keeled, but still wider at about midlength …………………………..……………………….…..…14 13’ Male and female with inner margin of hind tibia slightly arcuate, not keeled and not distinctly widest at about midlength ………………………………………..…...... 15

14 Pronotum with lateral margins divergent to basal 1/4, thence sub-parallel to apical

1/4, abruptly constricted to apex ……………………...T. arcuatibius NEW SPECIES 14’ Pronotum with lateral margins divergent from base to widest point at apical 1/3

………………………………………………...……………T. obrieni NEW SPECIES

15 Female with pronotum at middle of base with indistinct narrowly semicircular

glabrous area; pronotum regularly sculptured …………T. bravoensis NEW SPECIES 15’ Female with pronotum at middle of base with distinct acutely triangular glabrous area; pronotum with a number of irregular depressions

………………………………………………….…..T. heliconodulus NEW SPECIES

17 3.1.5 Species desriptions of the T. championi group

Trachyphloeomimus apionus Horsley, NEW SPECIES (Figs. 1-2, 33, 44)

Diagnosis. Small size; body form rounded and robust; elytra with intervals somewhat undulate, interval 5 appearing divergent apically, with widely spaced tubercles at declivity; male with ventral margin of legs with setae just longer than on other faces; aedeagus with flagellum distinctly expanded at apex and lateral margins heavily sclerotized, internal sac with minute sclerotized teeth in center of apical half of body (Fig. 33); female spermatheca plug-like in form (Fig. 44).

Description. Size. Length, male, 2.5mm; female, 2.5-2.7mm. Width, male, 1.4mm; female, 1.4-1.5mm. Colour and Integument. Body and appendages reddish-brown, clothed with indistinct and difficult to see (Figs. 1,2). Rostrum. Apex wider than base; base to point of antennal insertion densely clothed in golden brown indistinct scales, sparse short erect setae along mid-length; base distinctly widest just apical to eyes, distinctly pointed and triangular. Antennae. Scape distinctly clavate, extended nearly to pronotum; last article of funicle as wide as long. Head. Densely clothed with indistinct golden scales; eyes distinctly convex. Pronotum. Sub-cordate; lateral margins divergent from base to apical 1/3, widest points distinctly rounded to abruptly constricted slightly tubular apex (Fig. 1); clothed with indistinct golden scales, with a few copper shimmery scales; setae short and sporadically distributed; middle of posterior margin with very faint sub-glabrous half circle, generally more pronounced in female, often obscured by scales or dirt in male (Fig. 1). Elytra. Basal margin arcuate, interval 3 moderately raised along basal margin; humeri rounded, formed by interval 7; interval 8 weakly expanded as a posthumeral swelling to form widest point; strial punctures generally concealed by indistinct golden scales; all intervals appearing somewhat undulated, moreso on odd intervals; interval 5 appearing divergent apically, with distinct widely spaced tubercle at declivity; setae short and recumbent, sub-recumbent along declivity. Legs. Male with femora clavate, ventral margin with sparse setae just longer than remaining femoral setae,

18 outer face clothed with indistinct golden setae and short beige setae, hind tibiae with ventral margin very weakly sinuate, with sparse golden setae distally longer than other faces of tibiae; mucro relatively large. Female with tibiae with ventral margin dentate with short coarse setae, most coarsely dentate on hind tibia. Abdomen. Female with ventrite 5 distinctly sulcate medially; pygidium sinuate in female. Genitalia. Male with aedeagus in lateral view with body arcuate, at approximate mid-point weakly sinuate ventrally to apex (Fig. 33); in ventral view with body with lateral margins sub-parallel, evenly constricted to rounded weakly truncate apex; internal sac with minute sclerotized teeth in center of apical half of body; flagellum originating at mid-point of body and extending to half way down aedeagal apodemes, apex bell-shaped, markedly expanded with lateral margins heavily sclerotized. Female with spermatheca with ramus and nodulus approximately the same length, both curved outwards, plug-like in shape, nodulus extended just above ramus, then sharply curved outward, slight constriction at junction between ramus and nodulus and cornu (Fig. 44).

Material Examined. 1♂, 2♀♀ (MZLU)

Holotype ♀ (MZLU): a) Mexico: Oaxaca / 70km N Oaxaca / 2400 m 6.IX.1986 / R. Baranowski b) sifting litter, / mixed trop. / mont. forest c) WORLD / WEEVIL / DATABASE / WWD3003107 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Oaxaca City (70km N), 2400 m (17°29'58.63''N, 96°31'33.53''W), 6.ix.1986, R. Baranowski (1♀, 1♂ MZLU).

Etymology. Named from the Greek “apion” meaning pear, in reference to the pear- shaped form of this species.

19 Trachyphloeomimus arcuatibius Horsley, NEW SPECIES (Figs. 3-4, 34, 45, 60)

Diagnosis. Large size; colour dull brown; hind tibiae with ventral margin carinate in male; pronotum with sub-glabrous raised triangular area at mid-posterior margin, generally more pronounced in female, often obscured in male (Fig. 60); pronotum large and round; hind femora with inner ventral margin with sub-erect scales; odd intervals unevenly convex, interval 3 moderately convex at base; interval 5 with moderate tubercle at declivity; interval 1 convex at declivity; aedeagus with internal sac with minute sclerotized teeth throughout central portion of body, large portion of moderately sized sclerotized teeth near base of body filling entire width, smaller clusters above and below with larger sclerotized teeth (Fig. 34).

Description. Size. Length, male, 5.2-6.2mm; female, 5.2-5.7mm. Width, male, 2.2- 2.7mm; female, 2.5-2.8mm. Colour and Integument. Body and appendages brown; body densely clothed with imbricate recumbent oval-striate brown scales (Figs. 3, 4). Rostrum. Straight; base to point of antennal insertion clothed with brown scales; shallowly broadly medially sulcate between inner rows of setae; point of antennal insertion deeply broadly trifoveate, middle fovea largest, clothed with small metallic scales. Antennae. Scape clavate and weakly sinuate, extended to pronotum. Head. Frons weakly medially sulcate. Pronotum. Large, round; lateral margins divergent to basal 1/4, thence with sub-parallel margins to apical 1/4, abruptly constricted to apex; clothed densely with brown scales, medially with metallic scales interspersed with sub-glabrous areas; papillae with sub-erect brown and beige setae; sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male (Fig. 60). Elytra. Basal margin weakly arcuate; humeri very distinct, formed by interval 7; interval 8 moderately broadly expanded behind humeri as a posthumeral swelling; strial punctures densely covered with brown scales; odd intervals convex with sub-recumbent moderately thick brown and beige setae; interval 5 with moderate tubercle at declivity, interval 1 convex at declivity; declivity with setae sub-erect and coarsely spatulate (Fig. 3). Legs. Male with femora clavate with ventral margin strongly sinuate in

20 distal half, ventral margin with long fine golden setae, outer face clothed with golden- brown scales and short sub-erect golden setae, inner face clothed with small beige and metallic scales; hind femora of female with inner ventral margin with sub-erect scales; hind tibiae with ventral margin carinate and sinuate in distal half; tibiae with outer and inner faces clothed contiguously with brown scales, with few metallic scales distally, dorsal and outer margins with sub-erect coarse straight brown setae, ventral margin with long fine golden setae; apical comb with dense coarse golden setae on outer margin; mucro moderate. Female with tibiae with ventral margin dentate with short coarse setae, most coarsely dentate on hind tibia. Abdomen. Ventrite 5 shallowly sulcate, extended from apex to 1/3 from base; pygidium weakly sinuate. Genitalia. Male with aedeagus in lateral view with, body evenly weakly arcuate, apical 1/3 and tip of apex weakly diverted ventrally; in ventral view with, apex of body pointed, flagellum inverted and slightly extended through opening near apex, with thin portion extended between arms, internal sac with minute sclerotized teeth throughout central portion of body, large portion of moderately sized sclerotized teeth near base of body filling entire width, smaller clusters above and below with larger sclerotized teeth (Fig. 34). Female with spermatheca with ramus and nodulus of more or less equal width and length, divergent from one another, nodulus with apex curved outward (Fig. 45).

Material Examined. 8♂♂, 19♀♀ (CMNC, MZLU)

Holotype ♂ (MZLU): a) Mexico: Oaxaca, 21km / N Villa Diaz Ordaz / 3100 m, 10.IX.1990 / leg. R. Baranowski b) sifting litter, / boreal forest c) WORLD / WEEVIL / DATABASE / WWD3000365 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Oaxaca City (79km N), Hwy. 175, 2900 m (17°42'.72''N 96°21'7.92''W), 2.x.1990, R. Baranowski (1♀ MZLU); Oaxaca Ctiy (80km N), 2900 m (17°42'30.6''N 96°21'1.08''W), 8.ix.1994, R. Baranowski (3♀♀ MZLU); Valle Nacional (64.5km SW), Km 117.5, 2600 m (17°14'21.54''N 96°31'36.49''W), 28.vii.1992, R. S. Anderson (92-039) (1♀ CMNC); Valle Nacional (66km S), 2750 m (17°23'17''N,

21 96°30'27.3''W), 4.ix.1990, R. Baranowski (1♀ MZLU); same locality, 2.x.1990, R. Baranowski (3♀♀ MZLU); Valle Nacional (70km S), 2700 m (17°12'3.29''N 96°33'39.27''W), 29.ix.1990, R. Baranowski (1♀ MZLU); Villa Diaz Ordaz (21km N), 3100 m (17°8'51.12''N, 96°22'55.61''W), 7.ix.1990, R. Baranowski (1♂, 2♀♀ MZLU); same locality, 10.ix.1990, R. Baranowski (4♂♂, 7♀♀ MZLU); same locality, 20.ix.1990, R. Baranowski (2♂♂ MZLU).

Etymology. Noun in apposition; named from the Latin for “arc” and “tibia” in reference to the shape of the hind tibia in males of this species.

Trachyphloeomimus batmobile Horsley, NEW SPECIES (Figs. 5-6, 35, 46)

Diagnosis. Largest species; pronotum with small sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male (Fig. 5); hind tibiae with ventral margin carinate at more or less mid-length in male, dentate at apical 1/3 in female; interval 5 with strong tubercle at declivity, extended anteriorly as a distinct ridge giving ‘winged’ appearance; odd intervals weakly convex, with overall randomly uneven appearance; aedeagus with apex extended out and square-shaped in ventral view, flagellum of aedeagus expanded at apex with 2 small portions of medium- sized spicules at mid-length (Fig. 35).

Description. Size. Length, male, 5.3mm; female, 6.6-6.9mm. Width, male, 2.3mm; female, 3.2-3.4mm. Colour and Integument. Body and appendages brown to reddish- brown; clothed densely with very small contiguous recumbent oval-striate golden-brown scales (Figs. 5, 6). Rostrum. Apex straight to scarcely wider than base; base to point of antennal insertion densely clothed with golden-brown scales; point of antennal insertion deeply broadly tri-foveate, sparsely clothed with small metallic scales. Antennae. Scape thin, weakly clavate, extended to pronotum. Head. Basally shallowly rugose; broadly shallowly medially sulcate. Pronotum. Lateral margins divergent from base to widest point at mid-length (Fig. 5); widest point sharply constricted to apex in female, more

22 rounded in male; surface weakly acinose, clothed densely with small golden-brown scales and sparse sub-erect moderately thick straight brown setae, scales slightly darker beyond lateral margins; small sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male. Elytra. Basal margin arcuate; humeri distinct, formed by interval 7; interval 8 moderately broadly expanded behind humeri as a posthumeral swelling; strial punctures minute and often obscured by scales; clothed densely with small golden-brown scales; odd intervals weakly convex, with overall randomly uneven appearance; interval 5 with strong tubercle at declivity (Fig. 5), extended anteriorly forming distinct ridge, tubercle with concentration of setae; mainly odd intervals with sub-recumbent moderately thick spatulate brown and beige setae, occasionally with 2 rows on an interval. Legs. Male with femora clavate with ventral margin strongly sinuate in distal half, ventral margin with sparse sub-erect moderately thick straight brown and beige setae, other faces with recumbent brown and beige setae; outer face clothed with golden-brown scales with a band of beige scales 1/3 from apex, inner face clothed sparsely with metallic scales; tibiae with sub-erect moderately thick straight brown setae, outer face clothed with golden scales, inner face with sparse metallic and golden scales, dorsal margin with coarse setae in male, ventral margin with slightly longer and thinner golden setae; hind tibiae with ventral margin carinate at more or less mid-length in male; apical comb coarsely dentate on outer margin; mucro moderate. Female with hind tibiae with ventral margin dentate in distal 1/3. Abdomen. Suture between ventrites 1 and 2 weakly sinuate, moreso at lateral edges; ventrite 5 with deep broad fovea, broader at apex; pygidium emarginate in female, sinuate in male. Genitalia. Male with aedeagus in lateral view with, body evenly arcuate, apex diverted ventrally; in ventral view with, body nearly parallel-sided, convergent to apex, apex extended out and square-shaped; internal sac with minute sclerotized teeth in central portion of body, extended down to more or less mid-length of aedeagal apodemes; at mid-length of aedeagal apodemes, two large oval-shaped clusters of heavily sclerotized large teeth, flagellum extended between these structures to more or less 3/4 down aedeagal apodemes; flagellum expanded at apex, with 2 small portions of medium-sized spicules at mid-length (Fig. 35). Female with spermatheca with ramus and nodulus

23 bulbous, distinctly wider than cornu; ramus twice as long as nodulus, with 90 degree angle between ramus and nodulus (Fig. 46).

Material Examined. 1♂, 4♀♀ (MZLU)

Holotype ♀ (MZLU): a) Mexico: Oaxaca, 26km / E Teotitlan del Camino / 2250m, 26.IX.1990 / leg. R. Baranowski b) Sifting litter, / mixed oak forest c) WORLD / WEEVIL / DATABASE / WWD 3000217 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Teotitlan del Camino (25km E), 2200 m (18°12'41.53''N 96°51'53.27''W), 25.ix.1990, R. Baranowski (1♂ MZLU); Teotitlan del Camino (26km E), 2250 m (18°12'41.53''N 96°51'53.27''W), 26.ix.1990, R. Baranowski (3♀♀ MZLU).

Etymology. Noun in apposition; named after the batmobile, which this species resembles due to the large ridges along the declivity on interval 5.

Trachyphloeomimus bravoensis Horsley, NEW SPECIES (Figs. 7-8, 36, 47)

Diagnosis. Female with narrow semi-circular glabrous area on middle of posterior margin of pronotum; aedeagus with truncate apex, aedeagal apodemes longer than body, internal sac with minute sclerotized teeth throughout, two parallel elongate-oval moderately coarse areas of sclerotized teeth at apical 1/3 of base of aedeagal apodemes, flagellum of aedeagus distinctly expanded at apex, apical margins sclerotized (Fig. 36).

Description. Size. Length, male, 5.2mm; female, 6.5mm. Width, male, 2.5mm; female, 3.3mm. Colour and Integument. Body and appendages dark reddish-brown, clothed with brown indistinct scales (Figs. 7, 8). Rostrum. Apex distinctly wider than base, lateral margins of apex weakly flanged, central portion of with scarce red metallic small scales; base to point of antennal insertion medially moderately sulcate, clothed with

24 brown-golden indistinct scales and dirt and short erect brown setae. Antennae. Scape clavate, extended to anterior margin of pronotom; last article of funicle longer than wide. Head. Clothed with brown-golden indistinct scales and dirt; distinct raised tubercle dorsal to eyes with a few brown erect setae. Pronotum. Sub-cordate; lateral margins divergent from base to apical 1/3, distinctly rounded, weakly constricted to apex (Fig. 7); clothed with brown and golden indistinct scales and short sub-recumbent brown setae; middle of posterior margin with narrowly semi-circular sub-glabrous area, generally more pronounced in female, often obscured by scales or dirt in male. Elytra. Basal margin arcuate; humeri angular, formed by interval 7; interval 8 distinctly broadly expanded as a posthumeral swelling; strial punctures distinct, dirt encrusted; clothed with indistinct brown and golden scales; interval 5 weakly divergent from base to declivity, tubercle at declivity, extended basally in form of weak ridge (Fig. 7). Legs. Male with femora clavate, ventral margin sinuate, with long golden setae, outer surface with brown indistinct scales and beige setae, inner surface free of setae, with smaller indistinct red and golden metallic shiny scales; hind tibiae weakly arcuate, ventral margin with long golden setae, other surfaces with brown indistinct scales and brown and beige sub-erect setae; mucro moderate; female same as male except hind tibia with setae same length, ventral margin with short dentate golden brown setae apically. Abdomen. Ventrite 5 nearly flat; female with pygidium with apex very weakly sinuate. Genitalia. Male with aedeagus in lateral view with body arcuate, apex weakly diverted ventrally; in ventral view with body parallel-sided, evenly constricted to truncate apex; aedeagal apodemes longer than body; internal sac with minute sclerotized teeth throughout, two parallel elongate-oval moderately coarse areas of sclerotized teeth at apical 1/3 of base of aedeagal apodemes; flagellum extended to approximate mid-length of aedeagal apodemes, flagellum bell-shaped, expanded at apex, apical margins sclerotized (Fig. 36). Female with spermatheca with ramus and nodulus terminating in the same direction, nodulus hardly evident, appearing to originate from base of ramus, ramus at least 3 times length of nodulus (Fig. 47).

Material Examined. 1♂, 1♀ (CWOB)

25 Holotype ♂ (CWOB): a) MEXICO, Puebla / San Nicolas Bravo / 14-VI-1992, E. / Barrera, C. Mayorga b) C.W. O’BRIEN COLLECTION c) WORLD / WEEVIL / DATABASE / WWD3000214 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratype: Mexico: Puebla, San Nicolas Bravo, 14.VI.1992, E. Barrera, C. Mayorga. (1♀ CWOB).

Etymology. Named after the type locality.

Trachyphloeomimus calcaritibius Horsley, NEW SPECIES (Figs. 9-10, 37, 48, 65)

Diagnosis. Hind tibiae with ventral margin carinate and with large basal tooth in male (Fig. 65); coarsely striate beige scales; fore femur enlarged; pronotum with small glabrous semi-circular portion at base; pronotum with surface with large prominent papillae; sclerotized ridge surrounding and extended beyond eye; interval 5 with moderate tubercle at declivity (Fig. 9); ventral margin of leg with long fine beige setae in male (Fig. 65); aedeagus with body length small, apex short and rounded in ventral view, internal sac with medial cluster of large teeth at base of aedeagal apodemes, paired cluster of large teeth at more or less 1/3 down aedeagal apodemes, flagellum of aedeagus distinctly expanded at apex, lateral margins more heavily sclerotized at apex (Fig. 37).

Description. Size. Length, male, 3.5-4.6mm; female, 3.6-4.6mm. Width, male, 1.4- 1.9mm; female, 1.7-2.2mm. Colour and Integument. Body and appendages reddish- brown; clothed with large coarsely striate beige scales, imbricate on pronotum and sparse to contiguous on elytra (Fig. 9, 10). Rostrum. Straight; base to point of antennal insertion clothed with imbricate beige scales, bicarinate and medially weakly foveate; point of antennal insertion sub-glabrous with sparse moderately long fine golden setae; scrobe wide and glabrous. Antennae. Scape thin, sinuate and weakly clavate, extended to pronotum. Head. Lateral margins weakly rugose; eyes noticeably convex, with weak

26 ridge surrounding and extended beyond eye. Pronotum. Lateral margins divergent to mid-length, gently constricted to apex; clothed with imbricate recumbent coarsely striate beige scales and sparse sub-erect brown setae; surface with prominent papillae, occasionally glabrous; small glabrous semi-circular portion at mid-posterior margin. Elytra. Basal margin nearly straight; humeri distinct, formed by interval 7; interval 8 weakly broadly expanded behind humeri as a posthumeral swelling; strial punctures small, hardly evident; clothed sparsely with coarsely striate beige scales; odd intervals weakly convex with long sub-erect coarse spatulate brown setae in male, shorter setae in female; interval 5 with moderate tubercle at declivity with concentration of coarse spatulate beige setae. Legs. Male with femora clavate with ventral margin sinuate apically, ventral margin with long fine beige setae (Fig. 65), outer face clothed with slightly imbricate brown and beige scales and moderately long sub-recumbent beige setae, inner face clothed with beige and sparse small metallic scales in males; hind tibia with ventral margin with large basal tooth (Fig. 65), all tibiae with long fine beige setae, outer and inner faces clothed sparsely with recumbent coarsely striate brown scales, dorsal margins with moderately long sub-recumbent brown setae; apical comb with dense coarse golden setae and teeth on outer margin; mucro small to moderate. Female with hind femur with inner ventral margin with sub-erect beige scales in female; hind tibiae with ventral margin finely dentate; tibiae with short moderate sub-recumbent beige setae. Abdomen. Clothed with scales and sparse short setae; suture between ventrites 1 and 2 weakly medially emarginate in male, weakly sinuate in female; ventrite 5 with large broad shallow fovea in male, nearly flat in female; pygidium weakly sinuate in both sexes. Genitalia. Male with aedeagus in lateral view with, body almost c-shaped, body length small, more or less half length of aedeagal apodemes; in ventral view with, body with sides nearly parallel, apex short and rounded; internal sac with medial cluster of large teeth at base of aedeagal apodemes, paired cluster of large teeth at more or less 1/3 down aedeagal apodemes; flagellum just shorter than aedeagal apodemes, distinctly expanded at apex, flagellum more heavily sclerotized on lateral margins at apex; aedeagal apodemes and tegmen heavily sclerotized, dark (Fig. 37). Female with spermatheca hook- like in shape, strongly constricted at junction of ramus and nodulus with cornu, ramus

27 and nodulus pointed inward, nodulus shorter than ramus, appearing to originate from ramus; ramus more or less 2 times wider than nodulus (Fig. 48).

Material Examined. 18♂♂, 13♀♀ (MZLU)

Holotype ♂ (MZLU): a) Mexico: Oaxaca, 26km / E Teotitlan del Camino / 2250 m, 26.IX.1990 / leg. R. Baranowski b) Sifting litter, / mixed oak forest c) WORLD / WEEVIL / DATABASE / WWD3000221 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Teotitlan del Camino (26km E), 2250 m (18°12'41.53''N 96°51'53.27''W), 26.ix.1990, R. Baranowski (17♂♂, 13♀♀ MZLU).

Etymology. Noun in apposition; named from the Latin for “spur” and “tibia” in reference to the prominent spur on the hind tibia of males of this species.

Trachyphloeomimus carinatus Horsley, NEW SPECIES (Figs. 11-12, 38, 49, 61)

Diagnosis. Hind tibiae with ventral margin carinate in male; ventral margin of leg with long fine beige setae in male; pronotum with sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male (Fig. 61); pronotum with median area and apex with rounded copper and red metallic scales and elongate moderate foveae on lateral sides and along midline, deepest near apex; humeri formed by intervals 7 and 8 combined; interval 8 weakly expanded in conjunction with humeri; hind femora with inner ventral margin with erect beige scales; antennal club tapered on both ends; elytra with intervals 3 and 5 with strong tubercles at declivity, tubercles may be extended as an anteriorly directed ridge along the interval; intervals 3 and 9 join near elytral apex and form small tubercle; aedeagus with internal sac with 3 large sclerotized teeth medially at mid-length of body, 2 very large anvil-shaped sclerotized teeth at base of body, 3 large sclerotized teeth at more or less 1/3 from base of

28 aedeagal apodemes, flagellum extended from more or less 1/3 to 2/3 from base of aedeagal apodemes, thinner and more heavily sclerotized at base of flagellum, 2 moderate sclerotized teeth at base, apex more or less twice as thick as base and less sclerotized (Fig. 38).

Description. Size. Length, male, 4.5-4.7mm; female, 4.8mm. Width, male, 2.0mm; female, 2.4mm. Colour and Integument. Body and appendages dark reddish-brown; clothed with imbricate recumbent smooth brown and beige setae (Figs. 11, 12). Rostrum. Straight; base to point of antennal insertion clothed with beige scales, shallowly sulcate between inner rows of setae along length; point of antennal insertion moderately deeply bifoveate, clothed with small metallic scales. Antennae. Scape clavate, extended to pronotum; club tapered on both ends with short golden setae. Head. Shallowly rugose; weak transverse foveae in female. Pronotum. Lateral margins divergent from base to widest point at more or less mid-length, widest point gradually constricted to apex; clothed with smooth brown and beige scales, medially and apically with round copper and red metallic scales; elongate moderate foveae on lateral sides and medially, deepest apically; lateral margins with sub-recumbent coarsely striate beige scales; small sub- glabrous raised triangle at mid-posterior margin, generally more pronounced in female, often obscured in male (Fig. 61). Elytra. Basal margin arcuate; humeri distinct, formed by intervals 7 and 8; interval 8 weakly expanded in conjunction with humeri; strial punctures moderately deep; clothed with contiguous to weakly imbricate recumbent smooth beige scales with a metallic sheen; odd intervals convex; intervals 3 and 5 with moderate tubercles at declivity, stronger in female (Fig. 11); intervals 3 and 9 joined near apex of elytra forming ridge; mainly odd intervals with sparse short sub-recumbent spatulate brown and beige setae. Legs. Male with femora clavate with ventral margin sinuate in distal half, ventral margin with long fine golden setae, outer face clothed with contiguous recumbent oval-striate brown scales with a patch of beige scales at mid-point, outer face with moderately long sub-recumbent beige setae, inner face clothed with beige and metallic scales; hind femora with ventral margin of inner face with erect beige scales; hind tibia with ventral margin sinuate and carinate and weakly dentate, all tibiae with long fine golden setae, outer and inner faces clothed sparsely with recumbent brown,

29 beige and metallic scales, dorsal margin with moderately long sub-erect brown setae,; apical comb with dense coarse golden setae and teeth on outer margin; mucro large. Female with tibiae with ventral margin dentate, clothed with moderately long coarse brown setae. Abdomen. Clothed with scales and fine setae; suture between ventrites 1 and 2 basically straight and rugose; ventrite 5 with large rugose fovea in male, basically covers entire segment, moderately broadly basally convex and rugose apically in female; pygidium weakly sinuate in both sexes. Genitalia. Male with aedeagus in lateral view with, body evenly arcuate; in ventral view with, body with lateral margins sub-parallel, apex abrupt and rounded; internal sac with 3 large sclerotized teeth medially at mid- length of body, 2 very large anvil-shaped sclerotized teeth at base of body, 3 large sclerotized teeth at more or less 1/3 from base of aedeagal apodemes; flagellum extended from more or less 1/3 to 2/3 from base of aedeagal apodemes, thinner and more heavily sclerotized at base of flagellum, 2 moderate sclerotized teeth at base, apex more or less twice as thick as base and less sclerotized (Fig. 38). Female with spermatheca with ramus and nodulus divergent, nodulus approximately twice as long as ramus, nodulus curved inward at apex, ramus directed outward (Fig. 49).

Material Examined. 3♂♂, 1♀ (MZLU)

Holotype ♂ (MZLU): a) Mexico: Oaxaca, 18km N / Oaxaca City, 2400 m / 17.XI.1989 / leg. R. Baranowski b) sifting litter at road- / side, pine forest c) WORLD / WEEVIL / DATABASE / WWD3000466 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Oaxaca City (17km N), 2200 m (17°9'24.02''N, 96°36'52.68''W), 14.ix.1986, R. Baranowski (1♀ MZLU); Oaxaca City (18km N), 2400 m (17°9'54.69''N, 96°36'48.97''W), 17.xi.1989, R. Baranowski (1♂ MZLU); Oaxaca City (20km N), Hwy. 175, 2600 m (17°10'39.85''N, 96°36'43.06''W), 12.ix.1990, R. Baranowski (1♂ MZLU).

30 Etymology. Named from the Latin “carina” meaning keel, in reference to the characteristic keels on the elytra on adults of this species.

Trachyphloeomimus championi O’Brien (Figs. 13-14, 39, 50)

Trachyphloeomimus championi O’Brien, 1972: 170. Trachyphloeomimus championi, O’Brien and Wibmer (1982; checklist).

Diagnosis. Pronotum with middle of posterior margin with sub-glabrous raised small semi-circular portion, generally more pronounced in female, often obscured by scales or dirt in male; pronotum with surface acinose with large deep irregular punctures (Fig. 13); elytra with interval 5 with strong tubercle at declivity; hind tibiae of legs with ventral margin weakly carinate and dentate with long fine golden setae in male, dentate on apical half only in female; aedeagus with internal sac with minute sclerotized teeth extended from between base of apodemes to more or less apical 1/3 length of apodemes, apically with 2 to 3 sclerotized teeth, flagellum of aedeagus evenly widened to apex, apex weakly flared (Fig. 39).

Description. Size. Length, male, 4.9-6.3mm; female, 4.8-6.9mm. Width, male, 2.1- 2.8mm; female, 2.4-3.5mm. Colour and Integument. Body and appendages reddish- brown; clothed with imbricate recumbent oval-striate brown scales (Figs. 13, 14). Rostrum. Apex scarcely wider than base; base to point of antennal insertion clothed with small scales; bicarinate to point of antennal insertion, medially moderately sulcate along length; point of antennal insertion deeply medially foveate, clothed sparsely with small scales. Antennae. Scape weakly clavate, extended beyond eye and almost to anterior margin of pronotum. Head. Shallowly punctuate; moderately sulcate anterior to eyes; frons medially moderately foveate. Pronotum. Transversely oval; lateral margins divergent from base to widest point at more or less mid-length, thence with sub-parallel sides, sharply constricted at apex; surface acinose, with large deep irregular punctures, clothed with scales and sub-erect straight golden-brown setae; shallowly medially

31 sulcate, deepest sub-apically; with sub-glabrous raised small semi-circular portion at middle of posterior margin, generally more pronounced in female, often obscured by scales or dirt in male (Fig. 13). Elytra. Basal margin arcuate; humeri distinct, formed by interval 7; interval 8 weakly expanded behind humeri as a posthumeral swelling; strial punctures shallow, more or less equal to width of intervals; clothed with brown and golden-brown scales; interval 5 with strong tubercle at declivity (Fig. 13), tubercle with concentration of spatulate golden-brown setae; intervals 3 and 9 more convex apically and joined near apex of elytra forming ridge; odd intervals moderately convex, especially interval 3 at base, clothed with sub-recumbent moderately long straight golden-brown setae to declivity, then denser and spatulate on all intervals to elytral apex. Legs. Male with femora clavate, ventral margin strongly sinuate in distal half, with very long fine golden setae, outer face clothed with contiguous golden-brown scales, inner faces with beige scales becoming more imbricate in distal half, sub-recumbent moderate straight golden-brown setae; tibiae with golden-brown scales and sub-recumbent moderate straight golden-brown setae; hind tibiae with ventral margin sinuate, weakly carinate and dentate with long fine golden setae; apical comb with long coarse golden setae; mucro moderate; corbel bevelled. Female with hind tibiae with distal half dentate. Abdomen. Ventrites clothed with scales and fine golden setae, setae generally shorter in female; suture between ventrites 1 and 2 sinuate; ventrite 5 with apex moderately deeply broadly foveate, fovea deeper and broader in female; pygidium with apex sinuate in male, apex deeply broadly emarginate in female. Genitalia. Male with aedeagus in lateral view evenly arcuate; in ventral view with lateral margins sub-parallel, evenly constricted to pointed apex; aedeagal apodemes longer than body of aedeagus; internal sac with minute sclerotized teeth extended from between base of apodemes to more or less apical 1/3 length of apodemes, apically with 2 to 3 sclerotized teeth; flagellum thin, extended to more or less 4/5 length of aedeagal apodemes, evenly widened to apex, apex weakly flared (Fig. 39). Female with spermatheca with cornu strongly arcuate, hook-like in shape, extended to near apex of ramus and nodulus; strong constriction at point of attachment of ramus and nodulus to cornu; ramus and nodulus approximately equal in length, joined basally, nodulus appearing to originate from ramus; ramus more or less 3 times wider than nodulus (Fig. 50).

32

Material Examined. 20♂♂, 18♀♀ (CMNC, CWOB, CUIC)

Holotype ♂ (CWOB): MEXICO: State of Veracruz, Cofre de Perote, n. slope, 10.0 miles south Las Vigas, 9600 ft., 24-VIII-1967, Ball, T. L. Erwin, R. E. Leech. (Not examined).

Distribution. MEXICO: Mexico Distrito Federal, Cofre de Perote, 3500 m (19°30'26.4''N, 97°10'39.62''W), 2.vii.1972, J. Mateu (2♂♂, 6♀♀ CWOB); Veracruz, same locality, 3250 m (19°28'44.2''N, 97°7'58.31''W), 7.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (6♂♂, 3♀♀ CWOB); same locality, 3800 m (19°28'30.96''N, 97°8'40.61''W), 8.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (3♂♂, 2♀♀ CWOB); same locality, 3380 m (19°28'40.39''N, 97°8'9.08''W), 8.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (4♂♂, 1♀ CWOB); Cofre de Perote (15km E Perote), 3200 m (19°29'19.45''N, 97°7'32.09''W), 18.viii.1987, J. K. Liebherr, D. A. Millman (3♂♂, 1♀ CUIC); Las Lajas Micro., 16km S Rte. 140 at Las Vigas, 3000 m (19°32'14.6''N, 97°11'32.11''W), 29.vii.1990, J. K. Liebherr (2♂♂, 4♀♀ CMNC, CUIC); Las Lajas microondas (15km S Las Vigas), 3000 m (19°33'38.06''N, 97°9'18.81''W), 17.viii.1987, J. K. Liebherr, D. A. Millman (1♀ CUIC).

Trachyphloeomimus discus Horsley, NEW SPECIES (Figs. 15-16, 51, 62)

Diagnosis. Pronotum with sub-glabrous raised semi-circular portion at mid-posterior margin (Fig. 62); short recumbent setae; roundish striate scales; funicular segment 7 distinctly wider than long; interval 5 with strong tubercle at declivity (Fig. 15); tibiae with ventral margin sinuate and dentate in distal half; spermatheca with ramus and nodulus dinstinctly bulbous (Fig. 51).

Description. Size. Length, female, 3.8-5.0mm. Width, female, 2.1-2.7mm. Colour and Integument. Body brown; appendages reddish-brown; clothed with small recumbent

33 contiguous round striate golden scales (Figs. 15, 16). Rostrum. Apex wider than base; base to point of antennal insertion clothed with golden scales and sparse short sub- recumbent golden-beige setae; point of antennal insertion shallowly foveate with small sparse metallic scales; scrobe wide and glabrous. Antennae. Scape clavate and weakly sinuate, extended beyond eye and almost to anterior margin of pronotum; funicle with segment 7 noticeably wider than long; club squat and ovate with short fine beige setae. Head. Moderate fovea anterior to eyes. Pronotum. Lateral margins divergent from base to widest point 1/3 from apex, widest point rounded, constricted to apex (Fig. 15); clothed with small golden scales and short recumbent moderately thick straight brown and beige setae; sub-glabrous raised semi-circular portion at mid-posterior margin (Fig. 62). Elytra. Basal margin weakly arcuate; humeri distinct, formed by interval 7; interval 8 moderately pointed and expanded behind humeri as a posthumeral projection; strial punctures shallow, covered with scales; clothed with small golden scales, medially with sparse metallic scales; odd intervals convex, some with multiple rows of short recumbent coarse straight beige setae, even intervals with sparse setae; interval 5 with strong tubercle at declivity with concentration of beige setae (Fig. 15). Legs. Femora clavate with ventral margin strongly sinuate in distal half, with moderately long and thick sub- erect beige setae, outer face clothed with golden scales with some metallic, inner face sparsely clothed with small metallic scales; tibiae with ventral margin sinuate, coarsely dentate in distal half, clothed with golden and metallic scales and short sub-recumbent brown and beige setae; apical comb with dense coarse golden setae and dentate on outer margin; mucro large. Abdomen. Clothed with scales and short fine golden setae; suture between ventrites 1 and 2 sinuate; ventrite 5 with distinct moderately deep medially thin elongate fovea, lateral margins with small foveae; pygidium broadly sinuate. Genitalia. Spermatheca with ramus and nodulus with tips distinctly bulbous, polyp-like in form, nodulus thin beside ramus and extended past ramus by a length more or less equal to length of ramus, nodulus diverted outward and is bulbous at apex (Fig. 51).

Material Examined. 33♀♀ (MZLU)

34 Holotype ♀ (MZLU): a) Mexico: Oaxaca, 66km / S Valle Nacional / 2750 m, 4.IX.1990 / leg. R. Baranowski b) Sifting litter, / Salix and pine- / oak forest c) WORLD / WEEVIL / DATABASE / WWD3000429 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Oaxaca City (82km N), 2900 m (17°43'27.84''N 96°20'43.8''W), 8.ix.1994, R. Baranowski (4♀♀ MZLU); Valle Nacional (61km S), 2900 m (17°16'11.2''N 96°31'6.28''W), 29.ix.1990, R. Baranowski (14♀♀ MZLU); Valle Nacional (66km S), 2750 m (17°23'17''N, 96°30'27.3''W), 4.ix.1990, R. Baranowski (11♀♀ MZLU); same locality, 2.x.1990, R. Baranowski (1♀ MZLU); Valle Nacional (70km S), 2700 m (17°12'3.29''N 96°33'39.27''W), 29.ix.1990, R. Baranowski (1♀ MZLU); Valle Nacional (84km S), 2850 m (17°5'30.95''N 96°38'2.98''W), 4.ix.1990, R. Baranowski (1♀ MZLU).

Etymology. Named from the Latin “discus” meaning flat, in reference to the 7th funicular segment being flattened.

Trachyphloeomimus festivipenis Horsley, NEW SPECIES (Figs. 17-18, 40, 52, 66)

Diagnosis. Hind tibiae with ventral margin carinate distally in male; ventral margin of legs with long fine setae (Fig. 66); pronotum with small sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male; interval 5 with weak tubercle at declivity; hind femora with inner ventral margin with erect beige scales in female; aedeagus with internal sac with 5 clusters of elongate moderately sized dark sclerotized teeth forming a happy face with a hat pattern (Fig. 40).

Description. Size. Length, male, 3.0-3.3mm; female, 3.3-4.2mm. Width, male, 1.5- 1.7mm; female, 1.7-2.1mm. Colour and Integument. Body dark brown; appendages reddish-brown; body clothed with imbricate recumbent oval-striate brown scales (Figs. 17, 18). Rostrum. Apex straight to scarcely wider than base; base to point of antennal

35 insertion clothed with brown scales; point of antennal insertion foveate medially, clothed moderately with small metallic scales. Antennae. Scape clavate and sinuate, extended to pronotum, segment 1 noticeably bulbous apically. Head. Eyes distinctly convex; frons sulcate in lateral view. Pronotum. Lateral margins divergent from base to widest point at 1/3 from base, lateral margins sub-parallel from 1/3 from base to 1/3 from apex, sharply constricted to apex; clothed with brown and golden-brown scales, erect moderately coarse and long straight brown and beige setae; small sub-glabrous raised triangular portion at mid-posterior margin generally more pronounced in female, often obscured in male (Fig. 17). Elytra. Basal margin weakly arcuate; humeri subrectangular, formed by interval 7; interval 8 weakly broadly expanded behind humeri as a posthumeral swelling; strial punctures rarely evident, generally obscured by brown and golden-brown scales; odd intervals weakly convex; setae coarse and spatulate, brown and beige in colour, primarily on odd intervals, sub-erect in male, sub-recumbent in female; interval 5 with weak tubercle at declivity (Fig. 17). Legs. Male with femora clavate with ventral margin strongly sinuate apically, ventral margin with long fine straight golden setae, outer face clothed with slightly imbricate brown scales and short sub-recumbent brown and beige setae, inner face clothed with small metallic scales; hind tibiae with ventral margin sinuate and carinate from mid-length to apex (Fig. 66), all tibiae with outer and inner faces clothed contiguously with brown and sparse metallic scales, dorsal margin with short sub-erect coarse straight brown setae, ventral margin with long fine golden setae; apical comb with dense coarse golden setae on outer margin; mucro moderate. Female with hind femora with inner ventral margin with erect beige scales; hind tibiae with ventral margin dentate on distal half in female; tibiae with short sub-erect coarse straight brown setae. Abdomen. Clothed with beige scales with some metallic basally medially and moderately long sub-erect beige setae in male, shorter sub-recumbent setae in female; suture between ventrites 1 and 2 straight; ventrite 5 weakly medially sulcate in female, smaller in male; pygidium weakly sinuate in male, weakly emarginate in female. Genitalia. Male with aedeagus in lateral view with, body arcuate basally and apically with central portion more or less straight, apex straight to weakly diverted dorsally; in ventral view with, body with lateral sides nearly parallel, apex constricted to rounded point, body shorter than aedeagal apodemes; internal sac with 5 clusters of elongate

36 moderately sized dark sclerotized teeth forming a happy face with a hat pattern; cluster 1: at approximate mid-length of body, clusters 2 and 3: paired elongate oval clusters at junction of base of body and base of aedeagal apodemes, cluster 4: just below and central to clusters 2 and 3, cluster 5: U-shaped, at approximately 1/3 length of aedeagal apodemes; flagellum extended out of U-shape cluster to 1/5 from aedeagal apodeme apex, flagellum thin basally, distinctly expanded at apex, lateral margins of apex sclerotized (Fig. 40). Female with spermatheca with nodulus approximately twice length of ramus, nodulus straight then curved outward at mid-length, ramus and nodulus directed outward, "plug"-like in form, junction between ramus and nodulus and cornu constricted (Fig. 52).

Material Examined. 6♂♂, 30♀♀ (CMNC, CWOB, MZLU)

Holotype ♂ (MZLU): a) Mexico: Oaxaca, 21km / N Villa Diaz Ordaz / 3100 m, 7.IX.1990 / leg. R. Baranowski b) Sifting litter, / boreal forest c) WORLD WEEVIL DATABASE WWD3000389 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Oaxaca City (17km N), 2200 m (17°9'24.02''N, 96°36'52.68''W), 14.ix.1986, R. Baranowski (1♀ MZLU); Oaxaca City (20km N), Hwy. 175, 2600 m (17°10'39.85''N, 96°36'43.06''W), 12.ix.1990, R. Baranowski (3♀♀ MZLU); Oaxaca City (75km N), Hwy. 175, 2700 m (17°33'.63''N, 96°31'16.4''W), 4.ix.1990, R. Baranowski (1♀ MZLU); Suchixtepec (4.6km S), 2150 m (16°4'60''N 96°28'0''W), 23.vii.1992, R. S. Anderson (92-023) (3♀♀ CMNC); Valle Nacional (66km S), 2750 m (17°23'17''N, 96°30'27.3''W), 4.ix.1990, R. Baranowski (2♀♀ MZLU); Villa Diaz Ordaz (17km N), 2750 m (17°7'24.06''N, 96°23'3.09''W), 7.ix.1990, R. Baranowski (6♀♀ MZLU); same locality, 10.ix.1990, R. Baranowski (2♀♀ MZLU); same locality, 20.ix.1990, R. Baranowski (2♀♀ MZLU); same locality, 5.ix.1994, R. Baranowski (1♀ MZLU); Villa Diaz Ordaz (21km N), 3100 m (17°8'51.12''N, 96°22'55.61''W), 7.ix.1990, R. Baranowski (1♀, 2♂♂ MZLU); same locality, 10.ix.1990, R. Baranowski (1♀, 3♂♂ MZLU); Querétaro, Caderyta, Chavarrias (3km E), 2850 m (20°49.459'N,

37 99°35.191'W), 27.vii.2006, R. S. Anderson (2006-0019) (2♀♀ CMNC); Veracruz, Cofre de Perote, 3250 m (19°28'44.2''N, 97°7'58.31''W), 7.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (3♀♀ CWOB); same locality, 3800 m (19°28'30.96''N, 97°8'40.61''W), 8.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (1♀ CWOB); same locality, 3380 m (19°28'40.39''N, 97°8'9.08''W), 8.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (1♀ CWOB).

Not included in type series: Chiapas, San Cristobal (3mi W), 2042 m (16°44'58.29''N, 92°40'32.53''W), 28.vii.1983, J. Peck (2♀♀ CWOB).

Etymology. Noun in apposition; named from the latin for “happy” and “penis” in reference to the spicules of the internal sac being arranged in the form of a happy face.

Trachyphloeomimus heliconodulus Horsley, NEW SPECIES (Figs. 19-20, 53, 63)

Diagnosis. Pronotum with metallic scales medially; pronotum with sub-glabrous raised equilateral triangle at mid-posterior margin (Fig. 63); pronotum medially and apically acinose and with round red metallic scales; antennal club tapered on both ends; weak lateral protrusions over eyes; weakly broadly medially foveate between eyes; interval 5 with strong tubercle at declivity (Fig. 19); spermatheca with nodulus helical and at least 4 times as long as ramus (Fig. 53).

Description. Size. Length, female, 5.1-5.6mm. Width, female, 2.4-2.7mm. Colour and Integument. Body brown; appendages reddish-brown; clothed densely with imbricate recumbent oval-striate brown and golden scales; pronotum with medially raised patch of round red metallic scales (Figs. 19, 20). Rostrum. Straight; base to point of antennal insertion clothed with golden-brown scales; shallowly medially foveate; point of antennal insertion broadly moderately foveate, clothed with metallic scales. Antennae. Scape weakly clavate, extended beyond eye and almost to anterior margin of pronotum; club tapered on both ends with short golden-beige setae. Head. Frons weakly broadly

38 medially foveate; eyes large. Pronotum. Lateral margins sub-parallel from base to widest point almost at apex, widest point rounded and constricted to apex; clothed with golden- brown scales, medially and apically acinose with contiguous round red metallic scales; glabrous raised equilateral triangular portion at mid-posterior margin (Fig. 63). Elytra. Basal margin sinuate; humeri distinct, formed by interval 7; interval 8 weakly broadly expanded behind humeri as a posthumeral swelling; strial punctures shallow, more or less equal to width of interval; clothed with brown and golden scales; interval 3 convex basally; interval 5 with strong tubercle at declivity (Fig. 19), with concentration of setae; all intervals with sub-recumbent brown and beige setae, less setae on even intervals to declivity, from declivity to apex setae strongly spatulate. Legs. Femora clavate with ventral margin moderately sinuate in distal half, ventral margin with moderately long sub-erect beige setae, outer face clothed brown scales with a band of beige scales and short sub-recumbent beige setae, inner face densely clothed with beige and sparse metallic scales; tibiae with ventral margin dentate, outer and inner faces clothed with golden-brown and metallic scales, dorsal and ventral margins with moderately long sub- erect brown and beige setae; apical comb with dense coarse golden setae and teeth on outer margin; mucro moderate. Abdomen. Clothed with scales and moderately long sub- erect beige setae; suture between ventrites 1 and 2 weakly sinuate; ventrite 5 medially apically moderately foveate, apex weakly keeled; pygidium weakly emarginate. Genitalia. Spermatheca with ramus short and bulbous relative to nodulus, nodulus thinner and at least 4 times as long as ramus, middle portion sinuate and helical, somewhat plug-like in form, nodulus hook-like and diverted outward at apex (Fig. 53).

Material Examined. 5♀♀ (MZLU)

Holotype ♀ (MZLU): a) Mexico: Oaxaca / 73km N Oaxaca City, / 2600 m, 8.IX.1994 / leg. R. Baranowski b) sifting litter, / pine-oak forest c) WORLD / WEEVIL / DATABASE / WWD3000462 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

39 Paratypes: MEXICO: Oaxaca, Oaxaca City (73km N), 2600 m (17°32'29.92''N, 96°31'11.99''W), 8.ix.1994, R. Baranowski (1♀ MZLU); Oaxaca City (82km N), 2900 m (17°43'27.84''N 96°20'43.8''W), 8.ix.1994, R. Baranowski (2♀♀ MZLU); Oaxaca City (82km N), Hwy. 175, 2900 m (17°43'27.84''N 96°20'43.8''W), 29.ix.1990, R. Baranowski (1♀ MZLU).

Etymology. Noun in apposition; named from the Latin “helico” and “nodulus” in reference to the characteristic spiralling of the nodulus as seen in females of this species.

Trachyphloeomimus jonesi Horsley, NEW SPECIES (Figs. 21-22, 41, 54)

Diagnosis. Body form distinctly rounded; elytra with very weak tubercle on interval 5 at declivity (Fig. 21); setae acuminate short and fine; male with hind tibia with setae just longer than other faces; aedeagus with internal sac with minute sclerotized teeth throughout, at base of body sclerotized teeth are moderately coarse until basal 1/3 of aedeagal apodemes, flagellum with small medial cluster of moderate sclerotized teeth, extended to approximate apical 1/3 of aedeagal apodemes, flagellum weakly expanded at apex, apical margins weakly sclerotized (Fig. 41).

Description. Size. Length, male, 3.3mm; female, 3.9-4.8mm. Width, male, 1.5mm; female, 2.1-2.5mm. Colour and Integument. Body and appendages dark reddish-brown, clothed with indistinct golden brown scales (Figs. 21, 22). Rostrum. Apex nearly straight; apex to point of antennal insertion with small shiny metallic scales; base to point of antennal insertion with central portion very weakly sulcate, clothed with indistinct brown scales and dirt, with short erect acuminate setae. Antennae. Scape clavate, extended to anterior margin of pronotum; last article of funicle longer than wide. Head. Broadly very shallowly transversely foveate; weak tubercles dorsal to eyes with sparse erect setae. Pronotum. Lateral margins divergent from base to widest point at approximate apical 1/3, widest point rounded and gradually constricted to apex (Fig. 21); clothed with indistinct brown scales and short fine acuminate setae; middle of posterior

40 margin with small sub-glabrous raised triangular area, generally more pronounced in female, often obscured by scales or dirt in male. Elytra. Basal margin arcuate; humeri rounded, formed by interval 7; interval 8 very weakly expanded as a posthumeral swelling; strial punctures clothed with scales, approximately same width as intervals; clothed with indistinct scales and short fine acuminate setae; interval 5 with very weak tubercle at declivity (Fig. 21). Legs. Male with femora clavate, ventral margin sinuate in distal half, ventral margin with long golden setae, dorsal and outer faces clothed with indistinct scales and shorter golden acuminate setae; male with hind tibia with ventral margin carinate, widest point just apical to mid-length, ventral margin with setae just longer and thinner than dorsal and outer faces. Female with tibiae with ventral margin sinuate and dentate in distal half. Abdomen. Clothed with scales and acuminate golden setae; suture between ventrites 4 and 5 arcuate; ventrite 5 with weak medial apical fovea; pygidium with apex weakly sinuate. Genitalia. Male with aedeagus in lateral view with body evenly arcuate to apex; in ventral view with body nearly parallel-sided, evenly constricted to pointed apex; aedeagal apodemes longer than body; internal sac with minute sclerotized teeth throughout, at base of body sclerotized teeth are moderately coarse until basal 1/3 of aedeagal apodemes; flagellum with small medial cluster of moderate sclerotized teeth, extended to approximate apical 1/3 of aedeagal apodemes, flagellum weakly expanded at apex, apical margins weakly sclerotized (Fig. 41). Female with spermatheca with nodulus approximately 3 times length of ramus and extended beyond ramus, nodulus sharply curved outward apically, ramus and nodulus joined at the base, divergent from one another (Fig. 54).

Material Examined. 1♂, 19♀♀ (CMNC, UNAMQ)

Holotype ♀ (CMNC): a) MEXICO: QUERÉTARO / Mpio: Colon, Cerro Zamorano / 2770m, 26.VII.2006 / 20°55.967' N 100°11.021' W / R.S. Anderson, hardwood forest litter / 2006-0015 b) WORLD / WEEVIL / DATABASE / WWD3000471 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

41 Paratypes: MEXICO: Querétaro, Colon, Cerro Zamorano, 2770 m (20°55.967'N, 100°11.021'W), 26.vii.2006, R. S. Anderson (2006-0015) (1♀ CMNC); same locality, 26.vii.2006, R. S. Anderson (2006-0014) (15♀♀ CMNC); El Zamorano, (20°55'56.17''N, 100°10'45.78''W), 20.ii.1998, J. L. Cozar (1♀, 1♂ CMNC); Trigos (3km N), 2600 m (20°48.84'N, 100°10.73'W), 3.x.2001, R. Jones (1♀ UNAMQ).

Etymology. Patronymic; named after the collector of the majority of this species, Robert Jones of Querétaro, Mexico.

Trachyphloeomimus junctus Horsley, NEW SPECIES (Figs. 23-24, 55)

Diagnosis. Humeri formed by intervals 7 and 8 combined; interval 8 expanded weakly in conjunction with humeri; pronotum with small sub-glabrous raised triangular portion at mid-posterior margin; pronotum medially and apically with round dark copper metallic scales; pronotum with weak foveae on lateral sides and medially; interval 5 with tubercles at declivity extended medially interrupting intervals 4 and 3, appears as though intervals 5 and 3 are convergent (Fig. 23); intervals 3 and 9 join near elytral apex and form small tubercle; tibiae with ventral margin weakly dentate; spermatheca with nodulus approximately 3 times length of ramus (Fig. 55).

Description. Size. Length, female, 5.1-5.3mm. Width, female, 2.6-2.8mm. Colour and Integument. Body and appendages dark brown to dark reddish-brown; clothed with contiguous recumbent smooth brown with sparse beige scales (Figs. 23, 24). Rostrum. Straight; base to point of antennal insertion clothed with imbricate brown scales; bicarinate with shallow median fovea; point of antennal insertion moderately deeply trifoveate, clothed with sparse small metallic scales. Antennae. Scape clavate, extended beyond eye and almost to anterior margin of pronotum. Head. Weakly rugose, broadly shallowly transversely foveate; eyes appear small. Pronotum. Lateral margins divergent from base to widest point 1/3 from apex, widest point rounded and constricted to apex; clothed with golden-brown scales, medially and apically with round dark copper metallic

42 scales; weakly foveate on lateral sides and medially; small glabrous raised triangular portion at mid-posterior margin (Fig. 23). Elytra. Basal margin arcuate; humeri distinct, formed by intervals 7 and 8; interval 8 expanded weakly in conjunction with humeri; strial punctures moderately deep and large, more or less equal to width of interval; clothed with golden-brown scales; odd intervals weakly convex; interval 5 extended medially interrupting intervals 4 and 3 at declivity, appears as though intervals 3 and 5 are convergent (Fig. 23); intervals 3 and 9 joined near apex of elytra and forming ridge; mainly odd intervals with short sub-recumbent moderately thick brown setae, at declivity become coarser and spatulate. Legs. Femora clavate with ventral margin sinuate in distal half, clothed with moderately long golden-brown setae and brown scales with few beige scales, inner face densely clothed with golden-beige and sparse metallic scales; tibiae with ventral margin weakly dentate, clothed densely with recumbent brown scales and moderately long sub-erect golden-brown setae; apical comb with dense coarse golden- brown setae and teeth on outer margin; mucro moderate. Abdomen. Clothed with scales and short sub-recumbent beige setae; suture between ventrites 1 and 2 basically straight, lateral margins sinuate; ventrite 5 weakly broadly apically foveate; pygidium weakly sinuate. Genitalia. Spermatheca with nodulus approximately 3 times length of ramus and extended beyond ramus, nodulus sharply curved outward apically, curved portion very short, ramus and nodulus joined at the base, divergent from one another, spermathecal duct heavily sclerotized at point of attachment with nodulus (Fig. 55).

Material Examined. 2♀♀ (CMNC, MZLU)

Holotype ♀ (MZLU): a) Mexico: Oaxaca, 75km / N Oaxaca City, hwy 175 / 2700 m, 4.IX.1990 / leg. R. Baranowski b) Sifting litter, small / meadow in pine- / oak forest c) WORLD / WEEVIL / DATABASE / WWD3000470 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratype: MEXICO: Oaxaca, Llano de las Flores, 2713 m (17°7'38.22''N 96°36'3.1''W), 17.vii.1987, R. Anderson (87-25, 29) (1♀ CMNC).

43 Etymology. Named from the Latin “junctus” meaning joined, in reference to the joining of intervals 3 and 5 across interval 4 by a tubercle on the elytra of this species.

Trachyphloeomimus obrieni Horsley, NEW SPECIES (Figs. 25-26, 42, 56, 67)

Diagnosis. Pronotum with sub-glabrous raised triangular portion at base, generally more pronounced in female, often obscured in male; hind tibiae with ventral margin carinate and sinuate in male (Fig. 67); elytra with interval 5 with moderate tubercle at declivity (Fig. 25); pygidium emarginate in female; aedeagus with internal sac with minute sclerotized teeth throughout central portion of body, two parallel elongate-oval coarsely dentate sclerotized portions at base of aedeagal apodemes, flagellum of aedeagus markedly expanded at apex, apical margins distinctly sclerotized, with oval dentate structure slightly basal and ventral to apex (Fig. 42).

Description. Size. Length, male, 4.0-4.9mm; female, 4.0-5.8mm. Width, male, 1.8- 2.2mm; female, 1.9-2.9mm. Colour and Integument. Body brown, appendages reddish- brown; clothed with imbricate recumbent oval-striate grey-brown and golden scales (Figs. 25, 26). Rostrum. Apex straight to scarcely wider than base; base to point of antennal insertion clothed with brown scales; shallowly medially sulcate along length. Antennae. Scape weakly clavate and sinuate, segment 7 more robust in female. Head. Shallowly rugose; frons transversely weakly depressed. Pronotum. Cordate, widest at apical 1/3, lateral margins divergent from base to widest point 1/3 from apex; widest point rounded to constricted apex (Fig. 25); unevenly shallowly rugosely variolate, foveae clothed with recumbent oval-striate brown scales; raised areas between foveae sub-glabrous, occasionally with scales; sparsely clothed with sub-erect moderately thick brown setae; sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male. Elytra. Basal margin weakly arcuate; humeri weak, formed by interval 7; interval 8 broadly very weakly expanded behind humeri as a posthumeral projection; strial punctures shallow, clothed with recumbent oval-striate brown scales; odd intervals very weakly convex, with sub-erect

44 straight brown and beige setae until declivity, spatulate setae from declivity to elytral apex; even intervals with sparse setae before declivity then slightly more dense from declivity to elytral apex; interval 5 with moderate tubercle at declivity (Fig. 25), tubercle with small cluster of beige setae; intervals 3 and 9 joined near apex and forming small tubercle. Legs. Male with femora clavate, ventral margin strongly sinuate in distal half, with long fine golden setae, other faces with sub-recumbent moderately thick beige setae, all faces clothed with brown scales; tibiae with ventral margin sinuate and with long fine golden setae, clothed sparsely with brown scales, dorsal margin with sub-erect moderate straight brown setae; middle tibiae weakly carinate; hind tibiae strongly carinate, widest at mid-length; apical comb coarsely dentate on outer margin; mucro large (Fig. 67). Female with tibiae with ventral margin sinuate and dentate in distal half. Abdomen. Clothed with brown and beige scales, with sparse moderately long fine golden setae; suture between ventrites 1 and 2 sinuate, moreso along lateral margins in female; ventrite 5 apically deeply broadly foveate; pygidium with apex sinuate in male, apex emarginate in female. Genitalia. Male with aedeagus in lateral view with body evenly arcuate, apex weakly diverted ventrally; in ventral view with body parallel-sided, evenly constricted to rounded apex; aedeagal apodemes more or less same length as body; internal sac with minute sclerotized teeth throughout central portion of body, two parallel elongate-oval coarsely dentate sclerotized portions at base of aedeagal apodemes; flagellum extended to approximate mid-point of aedeagal apodemes, bell-shaped, markedly expanded at apex, apical margins distinctly sclerotized, with oval dentate structure slightly basal and ventral to apex (Fig. 42). Female with spermatheca with ramus and nodulus curving outwards, nodulus extended past ramus and curving out, slight constriction at junction between ramus and nodulus and cornu (Fig. 56).

Material Examined. 142♂♂, 34♀♀ (CMNC, CWOB)

Holotype ♂ (CWOB): a) MEXICO: Oax., Hwy. / 175, 53mi. NE. Oaxaca / 9800’ 3 June, 1983 C&L / O’Brien&GB Marshall b) under stones c) C.W. O’BRIEN / COLLECTION d) WORLD / WEEVIL / DATABASE / WWD3000040 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

45

Paratypes. MEXICO: Oaxaca, Llano de las Flores, 2713 m (17°7'38.22''N 96°36'3.1''W), 17.vii.1987, R. Anderson (87-25, 29) (7♂♂, 6♀♀ CMNC); Oaxaca (53mi NE), Hwy. 175, 2987 m (17°33'26.11''N, 96°27'35.25''W), 3.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (132♂♂, 27♀♀ CWOB); Valle Nacional (79.5km SW), Km 132, Llano de las Flores, 2650 m (17°7'38.22''N 96°36'3.1''W), 29.vii.1992, R. S. Anderson (92-042) (1♂, 1♀ CMNC).

Etymology. Patronymic; named after Charles and Lois O’Brien who collected the majority of the specimens.

Trachyphloeomimus popeyeus Horsley, NEW SPECIES (Figs. 27-28, 43, 57)

Diagnosis. Pronotum with sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male; pronotum with 3 patches of brown scales at base (Fig. 27); hind tibiae with ventral margin carinate and sinuate from near base to apex in male, with long erect fine golden-beige setae; strial punctures large and deep; interval 5 with moderate tubercle at declivity; aedeagus with apex rounded, nearly flat; internal sac with 4 pairs of dark moderate in size sclerotized teeth clusters, flagellum of aedeagus distinctly expanded at apex, lateral margins sclerotized (Fig. 43).

Description. Size. Length, male, 3.7-4.2mm; female, 3.7-4.7mm. Width, male, 1.6- 1.9mm; female, 1.8-2.4mm. Colour and Integument. Body and appendages dark reddish-brown to reddish-brown; clothed densely with imbricate recumbent oval-striate brown scales, with random clusters of golden-brown and beige scales (Figs. 27, 28). Rostrum. Straight; base to point of antennal insertion clothed with imbricate brown and golden-beige scales; shallowly medially sulcate between rows of setae; point of antennal insertion with moderate broad fovea; nasal plate glabrous. Antennae. Scape clavate and sinuate, extended beyond eye and almost to anterior margin of pronotum. Head.

46 Transversely weakly broadly sulcate. Pronotum. Lateral margins divergent from base to widest point 1/3 from apex, widest point rounded and constricted to apex; clothed densely with large golden-brown and beige scales, with 3 patches of darker brown scales at base, 1 at mid-posterior margin, 2 on posterior dorsolateral margins; lateral edges with beige scales; sub-glabrous raised triangular portion at mid-posterior margin, generally more pronounced in female, often obscured in male (Fig. 27). Elytra. Basal margin nearly straight; humeri distinct; formed by interval 7; interval 8 moderately expanded behind humeri as a posthumeral swelling; strial punctures large and deep, more or less equal to width of intervals; clothed with brown scales with random patches of golden-beige scales; odd intervals weakly convex, with sub-erect moderately thick straight brown and sparse beige setae in male, setae shorter and sub-recumbent in female; interval 5 with moderate tubercle at declivity, tubercle with golden-beige scales (Fig. 27). Legs. Male with femora clavate with ventral margin strongly sinuate in distal half, ventral margin with long fine golden setae, outer face clothed with weakly imbricate golden brown scales with a band of beige scales at 1/3 from apex, inner face clothed sparsely with small beige and few metallic scales; hind tibiae with ventral margin sinuate and carinate (Fig. 28), all tibiae with outer and inner faces clothed sparsely with golden-brown scales, dorsal margin with moderate sub-erect coarse brown setae, ventral margin with long erect fine golden-beige setae; apical comb coarsely dentate on outer margin; mucro moderate. Female with hind tibiae with ventral margin dentate in distal 1/3; all tibiae with ventral margin with short coarse setae. Abdomen. Suture between ventrites 1 and 2 weakly sinuate medially; ventrite 5 with deep elongate fovea; pygidium weakly sinuate in male, deeply emarginate in female. Genitalia. Male with aedeagus in lateral view with, body evenly weakly arcuate, apex diverted ventrally; in ventral view with, body weakly sinuate, thinnest at mid-point, apex rounded, nearly flat; internal sac with 4 pairs of dark moderate in size sclerotized teeth clusters; pair 1: just basal to opening of internal sac, elongate, extended to more or less mid-length of body; pair 2: just apical to base of body, small, circular; pair 3: at base of aedeagal apodemes, moderate in size, circular; pair 4: just basal to apex of flagellum, moderately elongate; flagellum extended from between cluster pair 3, at base of aedeagal apodemes to 2/3 down aedeagal apodemes, flagellum distinctly expanded at apex, lateral sides heavily sclerotized (Fig. 43). Female with

47 spermatheca with cornu extended to junction of nodulus and ramus with base of cornu; nodulus at least 2.5 times longer than ramus, extends past very small ramus, both ramus and nodulus weakly curved outward (Fig. 57).

Material Examined. 6♂♂, 12♀♀ (CMNC, CWOB, MZLU)

Holotype ♂ (MZLU): a) Mexico: Oaxaca, 61(handwritten) km S / Valle Nacional, 2900 m / 10.XI.1989 / leg. R. Baranowski b) sifting litter in / open boreal forest c) WORLD / WEEVIL / DATABASE / WWD3000344 (all typeset; genitalia extracted, in vial with glycerine, pinned beneath).

Paratypes: MEXICO: Oaxaca, Hwy. 175, Valle Nacional (37mi S), 2926 m (17°33'53.09''N, 96°30'28.71''W), 3.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (1♂ CWOB); Valle Nacional (35.9mi S), 2713 m (17°34'32.55''N, 96°30'50.43''W), 11- 18.viii.1973, A. Newton (1♂ CWOB); Valle Nacional (37mi S), 2591 m (17°16'54.5''N 96°30'51.41''W), 24.v.1971, S. Peck (Ber. 206) (1♀ CMNC); Valle Nacional (38mi S), Hwy. 175, 2987 m (17°27'41.81''N, 96°30'26.04''W), 16.v.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (1♀ CWOB); Valle Nacional (54km SW), Km 107, 2650 m (17°19'18.68''N 96°28'39.54''W), 28.vii.1992, R. S. Anderson (92-036) (2♀♀, 1♂ CMNC); Valle Nacional (55.5km SW), Km 108.5, 2800 m (17°28'24.7''N, 96°31'7.68''W), 28.vii.1992, R. S. Anderson (92-037) (2♀♀ CMNC); Valle Nacional (61km S), 2800 m (17°16'11.2''N 96°31'6.28''W), 3.x.1986, R. Baranowski (1♀ MZLU); same locality, 10.xi.1989, R. Baranowski (3♀♀ MZLU); same locality, 21.xi.1989, R. Baranowski (2♀♀, 2♂♂ MZLU).

Etymology. Named after the cartoon character Popeye, as this species has enlarged profemora resembling that of popeye’s arms.

48 Trachyphloeomimus spurcus Champion (Figs. 29-30, 58, 64)

Trachyphloeomimus spurcus Champion, 1911: 342. Trachyphloeomimus spurcus; O’Brien and Wibmer (1982; checklist).

Diagnosis. Pronotum acinose; pronotum with small sub-glabrous semi-circular portion at mid-posterior margin (Fig. 64); pronotum medially sulcate, deepest sub-apically; elytra with interval 8 not expanded behind humeri; elytra with odd intervals weakly convex; hind tibiae with ventral margin weakly sinuate and dentate from mid-length to apex; sub- erect beige scales near ventral margin of inner hind femora; sternite 5 with deep broad median apical fovea; pygidium strongly emarginate, almost furcate. Spermatheca hook- like in form with ramus and nodulus equal in length, distinct constriction between ramus and nodulus and curved cornu (Fig. 58).

Description. Size. Length, female, 3.2-4.7mm. Width, female, 1.6-2.4mm. Colour and Integument. Body brown; appendages reddish-brown; clothed with contiguous recumbent oval-striate brown scales (Figs. 29, 30). Rostrum. Apex scarcely wider than base; base to point of antennal insertion clothed with scales, weakly medially sulcate along length, extended to point of antennal insertion; point of antennal insertion with sparse small metallic scales. Antennae. Scape clavate, extended to pronotum. Head. Clothed with scales; weak lateral protrusions over eyes with few sub-erect brown setae; frons weakly transversely sulcate. Pronotum. Lateral margins divergent from base to widest point at more or less mid-length, widest point rounded and constricted to apex; surface acinose, densely clothed with scales and sub-erect straight brown setae; medially sulcate, deepest sub-apically; small sub-glabrous semi-circular portion at mid-posterior margin (Fig. 64). Elytra. Basal margin arcuate; humeri distinctly rounded, formed by interval 7; interval 8 not expanded behind humeri as a posthumeral swelling; strial punctures shallow, more or less equal to width of intervals; clothed with brown scales; odd intervals weakly convex, sub-erect coarse spatulate brown and beige setae on most intervals, more dense from declivity to elytral apex; interval 5 at declivity with weak to

49 moderate tubercle (Fig. 29). Legs. Femora clavate, ventral margin strongly sinuate distally, ventral margin with sub-erect moderately thick beige setae, outer face clothed with brown scales, inner face sparsely clothed with metallic and beige scales; hind femora with inner surface near ventral margin with sub-erect beige scales; hind tibiae with ventral margin weakly sinuate and dentate in distal half; tibiae clothed with brown scales, dorsal margin moderately clothed with sub-erect coarse straight brown setae, ventral margin with slightly longer and thinner golden setae; apical comb coarsely dentate on outer margin; mucro moderate. Abdomen. Clothed with beige and metallic scales and sub-erect straight beige setae; suture between ventrites 1 and 2 weakly sinuate; ventrite 5 medially deeply broadly foveate, swollen medially anterior to fovea; pygidium strongly emarginate, nearly furcate. Genitalia. Spermatheca with cornu strongly arcuate, hook-like in form, length extended to point of attachment of ramus and nodulus to basal end of cornu; large constriction at point of attachment of ramus and nodulus to cornu; ramus and nodulus equal in length, joined basally, basally attached portion equal in length to separate ramus and nodulus; ramus slightly thicker than nodulus, wider at apex (Fig. 58).

Taxonomic Notes. Only female specimens are known at this time. External characters and spermathecae show slight variation, but not enough to distinguish species reliably, therefore, all are considered conspecific. If males or more specimens are collected it may be possible to recognize more than the one species. This is one of the more widespread species, occurring in numerous states in Mexico. There are some small differences between populations but these have not been studied in detail. Paratypes as stated in O’Brien (1972) were not examined, although two specimens from a different paratype series were observed.

Material Examined. 94♀♀ (BMNH, CMNC, CUIC, CWOB, UNAMQ)

Holotype ♀ (BMNH): a) “23477” (handwritten on green paper label) b) “Truqui / Mexico” (handwritten) c) “Fry Coll. / 1905. 100.” (typeset) d) “Type” (2 labels, typeset on round orange and white label) e) “sp. figured” (typeset) f) “B.C.A., Col., IV. pt. 3. /

50 Trachyphlœomimus (typeset) / spurcus Ch. (handwritten) g) “SYN- / TYPE” (typeset on round blue and white label) h) WORLD / WEEVIL / DATABASE / WWD3003406 (typeset). (Examined).

Distribution. MEXICO: Bejucos, Temescaltepec, (18°46'33.65''N, 100°25'38.28''W), 3.vii.1933, H. E. Hinton, R. L. Usinger (1♀ CMNC); Popo Park, (19°4'47.44''N, 98°46'52.68''W), 18.viii.1968, R. Muniz V. (1♀ CWOB); Guanajuato, Santa Rosa, Santa Rosa (3km N), 2620 m (21°4.849'N, 101°11.52'W), 30.vii.2006, R. S. Anderson (2006-0029) (7♀♀ CMNC); same locality, 2620 m, 30.vii.2006, J. L. Cozar (15♀♀ CMNC); same locality, 2620 m, 30.vii.2006, P. J. Horsley (2006-018A) (1♀ PJHC); Mexico Distrito Federal, La Puerta, Toluca Est., (19°16'58.17''N, 99°39'20.59''W), 11.vii.1972, J. Mateu (1♀ CWOB); Mexico City (21km S), Hwy. 95, (19°13'29.14''N, 99°8'23.56''W), 5.ix.1982, C. W. O'Brien, L. O'Brien, G. J. Wibmer (4♀♀ CWOB); Parque Nacional Cumbres de Ajusco, Llano de Cantimplora, 3340 m (19°13'24.1''N, 99°15'17.53''W), 4.ix.1982, C. W. O'Brien, L. O'Brien, G. J. Wibmer (2♀♀ CWOB); Parque Nacional Miguel Hidalgo, 2900 m (19°19'32.49''N, 99°19'9.5''W), 17.ii.1972, L. R. O'Brien (4♀♀ (BMNH, CWOB); same locality, 3109 m, 29.v.1974, C. W. O'Brien, L. O'Brien, G. B. Marshall (1♀ CWOB); Paso de Cortes (19km W), 2550 m (19°12'49.74''N, 99°6'48.94''W), 2.vi.1979, A. N. Garcia Aldrete (2♀♀ CWOB); Popocatepetl, 3000 m (19°4'3.6''N, 98°41'42.38''W), 22.viii.1981, J. Palacios (4♀♀ CWOB); Pradera de Salazar, (18°55'18.78''N, 99°14'.07''W), 17.ii.1972, P. Reyes (1♀ CWOB); same locality, 17.ii.1972, C. W. O'Brien (1♀ CWOB); Salazar, (18°55'18.78''N, 99°14'.07''W), 19.v.1972, J. Mateu (2♀♀ CWOB); San Francisco de los Ranchos, 19.x.1983, H. Brailovsky, Barrera (2♀♀ CWOB); Zoquiapan, 3500 m (19°30'N, 98°45'W), 18.iii.1992, A. Moron-Rios (1♀ CMNC); same locality, 18.iii.1992 (1 ♀ CMNC); Xilotepec, Hwy.57, Km 107, 2700 m (20°4.797'N, 99°37.906'W), 25.vii.2006, R. S. Anderson (2006-0013) (1♀ CMNC); Morelos, Tres Cumbres (7mi S), (19°1'16.05''N, 99°9'52.39''W), 7.vii.1975, Q. D. Wheeler (1♀ CMNC); same locality, 7.vii.1975, D. S. Chandler (1♀ CWOB); Puebla, Tlachichuca (7.6km E), 2700 m (19°6'18.48''N, 97°23'26.66''W), 22.viii.1987, J. K. Liebherr, D. A. Millman (1♀ CUIC); Vicente Guerrero (7.6km E), 2420 m (19°1'30.88''N, 98°6'55.22''W), 16.vii.1992, J. S.

51 Ashe (1♀ CMNC); Querétaro, Xilitla (29.1km W), 1524 m (21°22'6.63''N, 99°16'28.07''W), 3.vi.1987, R. Anderson (2♀♀ CMNC); Colon, Cerro Zamorano, 3100 m (20°55.967'N, 100°11.021'W), 26.vii.2006, R. S. Anderson (2006-0014) (2♀♀ CMNC); Trigos (5km NW), 3100 m (20°56.23'N, 100°11.09'W), 6.ii.2006, R. Jones, M. Perez (#2) (1♀ UNAMQ); Huimilpan, La Beata (1km NW), 2500 m (20°18.42'N, 100°14.49'W), 2.xii.2003, R. Jones (sample #2) (1♀ UNAMQ); Pinal de Amoles, Cruz de Palo (ca. Pinal de Amoles), 2450 m (21°8.095'N, 99°38.021'W), 28.vii.2006, R. S. Anderson (2006-0025) (3♀♀ CMNC); Pinal de Amoles (3.4km E), 2120 m (21°9.095'N, 99°36.659'W), 28.vii.2006, R. S. Anderson (2006-0024) (3♀♀ CMNC); San Joaquin, above Camp Alegre, 2500 m (20°54.82'N, 99°34.8'W), 14.viii.2004, R. Jones (#2) (1♀ UNAMQ); Ranas Zona Arqueologica, 2350 m (20°55.514'N, 99°33.315'W), 27.vii.2006, R. S. Anderson (2006-0020) (1♀ CMNC); San Luis Potosi, San Luis Potosi (34km E), 2400 m (22°5'1.97''N, 100°39'20.39''W), 2.vi.1987, R. Anderson (1♀ CMNC); San Luis Potosi (42km E), 2377 m (22°2'55.79''N, 100°36'52.37''), 2.vi.1987, R. Anderson (14♀♀ CMNC); Tlaxcala, Tlaxca (6.8km N), 2820 m (19°39'17.89''N, 98°6'6.42''W), 10.vii.1992, J. S. Ashe (#119) (4♀♀ CMNC); Veracruz, Cofre de Perote, 3250 m (19°28'44.2''N, 97°7'58.31''W), 7.vi.1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (3♀♀ CWOB); La Paisano, 9.8km S Las Vigas, 2800 m (19°34'33.96''N, 97°9'33.98''W), 17.viii.1987, J. K. Liebherr, D. A. Millman (2♀♀ CUIC); Las Lajas microondas (15km S Las Vigas), 3000 m (19°33'38.06''N, 97°9'18.81''W), 17.viii.1987, J. K. Liebherr, D. A. Millman (1♀ CMNC).

Trachyphloeomimus undulatoides O’Brien (Figs. 31-32, 59)

Trachyphloeomimus undulatoides O’Brien, 1972: 176. Trachyphloeomimus undulatoides; O’Brien and Wibmer (1982; checklist).

Diagnosis. Pronotum with middle of posterior margin lacking glabrous area; elytra undulate and distinctly uneven (Fig. 31); spermatheca with nodulus approximately twice length of ramus (Fig. 59).

52

Description. Size. Length, female, 3.3-4.1mm. Width, female, 1.8-2.2mm. Colour and Integument. Body and appendages dark reddish-brown, clothed with few striate scales, mostly indistinct and covered in dirt, resulting in brown colour (Figs. 31, 32). Rostrum. Apex nearly straight; base to point of antennal insertion clothed with scales and sub-erect setae, deeply medially sulcate to mid-length of eyes; point of antennal insertion to apex sub-glabrous, with median carina. Antennae. Scape clavate, extended just beyond posterior margin of eye; funicle with last article as wide as long. Head. Clothed with indistinct scales; lateral protrusions dorsal to eyes lacking, multiple erect setae present. Pronotum. Lateral margins evenly divergent from base to approximate mid-length, sub- parallel and weakly sub-cordate in form; clothed with indistinct brown scales and sub- erect brown and beige clavate setae, few beige striate scales in median apical 1/3 weak sulcus; surface uneven; middle of posterior margin lacking glabrous area. Elytra. Basal margin weakly sinuate; humeri angular, formed by interval 7; interval 8 moderately narrowly expanded as a posthumeral swelling; strial punctures covered in scales, difficult to see; clothed with indistinct brown scales and dirt encrusted; mainly odd intervals with clavate sub-erect beige and brown setae; surface distinctly uneven; interval 5 with strong tubercle at declivity; interval 1 broadly expanded at declivity (Fig. 31). Legs. Femora clavate, ventral margin sinuate in distal half, dorsal and outer faces covered with scales and sub-erect clavate brown and beige setae, inner face with metallic and beige scales; hind tibiae weakly with ventral margin sinuate and dentate in distal half, clothed with brown-golden scales and sub-erect setae; apical comb coarsely dentate on outer margin; mucro moderate. Abdomen. Clothed with beige and scales and sub-erect golden setae; suture between ventrites 1 and 2 nearly straight; ventrite 5 markedly trifoveate near lateral margins and medially just apical to center; pygidium emarginate. Genitalia. Spermatheca with nodulus approximately twice length of ramus, ramus nearly twice as thick as nodulus, nodulus divergent from ramus beginning at approximate quarter-length, nodulus with apex curved outward (Fig. 59).

Material Examined. 13♀♀ (BMNH, CMNC, CWOB)

53 Holotype ♀ (CWOB): MEXICO: State of Mexico, Parque Nacional Miguel Hidalgo, ca. 2900 meters, under stones, C. W. O’Brien. (Not examined).

Paratypes: (as stated in O’Brien, 1972) MEXICO: Mexico Distrito Federal, Parque Nacional Miguel Hidalgo, 2900 m, 19° 19' N, 99° 19' W, 17 February 1972, S. Rosenthal (1♀ BMNH, Examined); Pradera de Salazar, 18° 55' N, 99° 14' W, 17 February 1972, C. W. O'Brien (1♀ CWOB, Examined); State unknown, Llano Grande, 6 May 1962, M. Olguin (1♀ RMVC, Not examined).

Distribution. MEXICO: Mexico Distrito Federal, Parque Nacional Miguel Hidalgo, 3109 m, 19° 18' N, 99° 19' W, 29 May 1974, C. W. O'Brien, L. O'Brien, G. B. Marshall (3♀♀ CWOB); Salazar, 18° 55' N, 99° 14' W, 19 May 1972, J. Mateu (1♀ CWOB); Salazar, Hwy. 15, 3170 m, 19° 1' N, 99° 20' W, 6 June 1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (4♀♀ CWOB); Puebla, La Cumbre (8km SW), 1600 m, 23 July 1987, R. Anderson (87-39) (1♀ CMNC); Zacapoaxtla (9mi S), 2286 m, 19° 48' N, 97° 38' W, 11 June 1983, C. W. O'Brien, L. O'Brien, G. B. Marshall (1♀ CWOB); Veracruz, La Paisano, 9.8km S Las Vigas, 2800 m, 19° 34' N, 97° 9' W, 17 August 1987, J. K. Liebherr, D. A. Millman (1♀ CMNC); Las Vigas (5.6km S), 2750 m, 19° 35' N, 97° 7' W, 8 May 1999, A. Howden, R. Arce (1♀ CMNC).

3.2 Phylogenetic Analysis

As with most groups of Curculionidae in which there are numerous, closely related and structurally very similar species, resolution of phylogenetic relationships is often a difficult and complex procedure and is rarely attempted (Anderson and O’Brien 1996). This study appears not to be an exception and resolution of relationships within the species of Trachyphloeomimus has proven to be a taxing analysis (see Figs. 68-69). Nevertheless, progress has been made and the results presented herein represent the first steps towards developing a phylogenetically-based classification of these weevils at a level below the genus group.

54 The results of the phylogenetic analysis are shown on Figures 68-69. Initially, unweighted analyses failed to demonstrate the monophyly of the T. championi group and failed to resolve much in the way of relationships. This appears to be attributed to a “swamping effect” by characters known to be homoplasious elsewhere in Entiminae over those characters with states not known to occur elsewhere in Entiminae, and thus of greater phylogenetic value. For this reason, subsequent analyses followed the method of Anderson (1987) who employed a weighted character analysis emphasizing character states unique within Entiminae as having the greatest significance and therefore the highest weight. Following weighting of these characters, a more fully resolved and logically acceptable hypothesis of relationships was achieved. This set of relationships recognizes the monophyly of the T. championi group based largely upon the characters of form of the basal margin of the pronotum (Figs. 60-64) and form of the male hind tibia (Figs. 65-67). As noted, apomorphic states of these characters are not known in other broad-nosed weevils and thus there is no a priori evidence of homoplasy. The “problem” taxa in the initial unweighted analysis were T. spurcus and T. jonesi; however, after character re-weighting they were placed within the T. championi species group. Their placement therein is supported by other compatible characters to be discussed following. Once the monophyly of the T. championi species group was established, other characters emerged as compatible with this result. In addition to character 10 (pronotum with basal margin modification) and character 21 (male hind tibia modified), characters 23 (female hind tibia dentate), 24 (female pygidium form), 14 (8th interval form), 19 (interval 3 at base), and 20 (tubercle pattern), are further compatible with the more heavily-weighted characters and aid in the resolution of this group of species. Clearly, much remains to be resolved concerning relationships among Trachyphloeomimus species. As seen in Figure 69, the majority of the species relationships are unresolved, with the exception of only the T. championi group and a second unnamed group of species. An additional problem with this genus (and weevils generally) is the occurrence of parthenogenicity with numerous species propagating without the need of males. For example, within the T. championi group, six species are known only from females, thereby reducing the numbers of characters available for placement of these species.

55 4. DISCUSSION AND CONCLUSION

What started out as a revision of a genus of weevils in which there were eight previously recognized species has “blossomed” (perhaps “erupted” is a better term) into a megadiverse assemblage of approximately 80 species. Most of these species are similar in structure and differ in combinations of character states rather than possession of unique character states not found anywhere else in the genus. Anderson (in press) has found a similar situation in weevils of the genus Theognete Champion, a group in which the one described species has now almost 100 additional congeners. Other examples are the genera Anchonus, Eurhoptus and Tylodinus, and in fact, this situation is very common among many groups of understudied Neotropical weevils (Anderson and O’Brien 1996). As species wthin the T. championi species group are generally found at high montane elevations from approximately 2000 meters to well over 3000 meters, it is not surprising to find such a richness of species diversity restricted to the central Mexican highlands. These species are much more likely to be endemic to the region due to the complexity and isolating effects of the topography of Mexico (Anderson and O’Brien, 1996). Many of the collecting sites for the T. championi species group are only kilometres apart, but populations are highly localized and generally with each new collection site, a new species is discovered. Even with this being said, it’s certain there are still more Trachyphloeomimus species to be found. Trachyphloeomimus is a diverse genus of weevils and with still large gaps in collecting and the number of actual species in Central America known, it is difficult to make an accurate assessment of relationships. As the picture becomes more complete with time, the ability to better predict character evolution should lead to a better understanding of the relationships among species of Trachyphloeomimus. Another interesting aspect of the T. championi species group is the fact that six of the sixteen species are known from females only. The presence of parthenogenetic species that are more widely distributed, as exemplified by T. spurcus, seems to indicate an enhanced ability to disperse and colonize. Although there are slight differences in the external appearances of populations in T. spurcus, there is not enough evidence to warrant splitting of the species into multiple species.

56 While this thesis has not resolved all relationships among Trachyphloeomimus species, it has established a set of relationships and a start at a species group level classification facilitating future study of the genus. The fact that the T. championi species group was found to be monophyletic is encouraging. Future work should continue to emphasize character states unique within the genus as of the highest importance for resolving phylogenetic relationships and molecular methods should be employed to add additional characters and resolve the ‘swamping effect’ of the secondary characters. Certainly, within Trachyphloeomimus and the weevils in general, there is no shortage of work needing to be done. The main objective of this thesis was to provide a better systematic understanding of the genus Trachyphloeomimus. The T. championi species group was found to be monophyletic and displays two unique characters not found in any other Entiminae. The results of this study will provide a useful framework for future revisionary work and has taken a step in the right direction to better our understanding and knowledge of the megadiversity of weevils, especially in Latin America.

57 REFERENCES

Alonso-Zarazaga, M.A. and C.H.C. Lyal. 1999. A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae). Entomopraxis. Barcelona, Spain. Anderson, R.S. 1987. Systematics, phylogeny and biogeography of New World weevils traditionally of the tribe Cleonini (Coleoptera: Curculionidae; Cleoninae). Quaestiones Entomologicae 23(4): 431-709. Anderson, R.S. 2002. American beetles; Polyphaga: Scarabaeoidea through Curculionoidea. ed. R.H. Arnett, M.C. Thomas, P.E. Skelley, J.H. Frank. Boca Raton, Fla.: CRC Press. Anderson, R.S. In press. A taxonomic monograph of the Middle American leaf-litter inhabiting weevil genus Theognete Champion (Coleoptera: Curculionidae; ; Lymantini). Zootaxa, in press. Anderson, R.S. and J.S. Ashe. 2000. Leaf litter inhabiting beetles as surrogates for establishing priorities for conservation of selected tropical montane cloud forests in Honduras, Central America (Coleoptera; Staphylinidae, Curculionidae). Biodiversity and Conservation 9(5): 617-653. Anderson, R.S. and R.G. Oberprieler. In press. Curculionoidea. In: Capinera, J.L. (Ed.), Encyclopedia of Entomology (Edition 2), 4 Volumes. 4000 pp, 1300 illus. Springer-Verlag. Anderson, R.S. and C.W. O'Brien. 1996. Curculionidae (Coleoptera). pp. 329-351, In: Llorente Bousquets, J.E., García Aldrete, A.N. and González Soriano, E. (Editores). Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una Síntesis de su Conocimiento. México. xvi + 660 pp. Blackwelder, R.E. 1947. Checklist of the Coleopterous of Mexico, Central America, the West Indies, and South America. Part 6. Bulletin, United States National Museum. 185: i-iv, 765-925. Bright, D.E. and P. Bouchard. 2008. The Insects and Arachnids of Canada Part 25: Coleoptera, Curculionidae, Entiminae; Weevils of Canada and Alaska Volume 2. NRC Research Press Ottawa, Canada.

58 Champion, G.C. 1911. Biologia Centrali-Americana, Insecta, Coleoptera, Curculionidae. Supplement to the Thecesterninae and Otiorhynchinae. 4(3): 342- 343. Evenhuis, N.L. (21 April 2009). The insect and spider collections of the world website. Available at: http://hbs.bishopmuseum.org/codens/ codens-inst.html (07 January 2010). Howden, A.T. 1982. Revision of the New World genus Hadromeropsis Pierce (Coleoptera, Curculionidae, Tanymecini). Contributions of the American Entomological Institute 19(6): 1-180. Kuschel, W. 1995. A phylogenetic classification of Curculionoidea to families and subfamilies. Memoirs of the Entomological Society of Washington 14: 5-33. Lacordaire, [J], T. 1863. “Histoire Naturelle des Insectes. Genera des Coleopteres.” Vol. 7. p. 191-196. Roret, Paris. Lona, C. 1937. Coleopterorum Catalogus, pars 160, Curculionidae, Otiorhynchinae II, p. 227-412. W. Junk, Berlin. Martin, J.E.H. 1977. The Insects and Arachnids of Canada Part 1: Collecting, Preparing and Preserving Insects, Mites and Spiders. NRC Research Press, Ottawa, Canada: p. 1-182. Marvaldi, A.E. 1997. Higher level phylogeny of Curculionidae (Coleoptera: Curculionoidea) based mainly on larval characters, with special reference to broad- nosed weevils. Cladistics 13: 285-312. Marvaldi, A.E. 1998. Larvae of Entiminae (Coleoptera: Curculionidae): tribal diagnoses and phylogenetic key, with a proposal about natural groups within Entimini. Entomologica scandinavica 29: 89-98. Morón-Ríos, A, V.J. Jaramillo and R. Dirzo. 1997. Species composition of root-feeding macroarthropods in a subalpine grassland and associated with pine forest in Mexico. The Canadian Entomologist 129: 71-80. Oberprieler, R.G., A.E. Marvaldi and R.S. Anderson. 2007. Weevils, weevils, weevils everywhere. Zootaxa 1668: 491-520.

59 O’Brien, C.W. 1972. A review of the Mexican and Central American genus Trachyphloeomimus, with new species and new synonymy (Coleoptera: Curculionidae, Otiorhynchinae). The Coleopterists Bulletin 26(4): 165-178. O’Brien, C.W. and G.J. Wibmer. 1982. Numbers of genera and species of Curculionidae (Coleoptera). Entomological News 89(2 & 3): 89-92. Sharp, D. 1911. Biologia Centrali Americana, Insecta, Coleoptera, Rhynchophora, Curculionidae: Otiorhynchinae. [part “Apterae”] 4(3): 169-178. Sleeper, E.L. 1955. A review of the Trachyloeini of America north of Mexico (Coleoptera, Curculionidae). The Ohio Journal of Science 55(5): 279-292. Thompson, R.T. 1992. Observations on the morphology and classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. Journal of Natural History 26: 835-891.

60

TABLE 1. Data matrix used in the phylogenetic analysis of Trachyphloeomimus species and related taxa. Box indicates the species included in the T. championi species group.

61 Character 0123456789101112131415161718192021222324252627282930313233

S. latipennis 0220010000000300110000001000105420 C. komareki 00200001000003000100000??0?102056? C. chrysorrhoeus 002000010000030001000??02?0??????7 T. championi 1100000101122111112111112110000235 T. obrieni ?100010101232111112111112110002234 T. bravoensis ?100010101132110102111112000002336 T. batmobile 1100010101232110112111112220002336 T. calcaritibius 11?00101011?0111112111112100020235 T. spurcus 010000010113200011000??13?1??????5 T. popeyeus 1100010101232111112111113210002337 T. arcuatibius ?100010101222021112111112100000233 T. festivipenis 0100010101222111112101113100000334 T. discus 71?001111123011100211??12?2??????9 T. heliconodulus 410001010123001111211??12?2??????1 T. carinatus ?10001010123012111116111224000003? T. junctus 410001010113012111214??12?1??????2 T. jonesi ?100000101120001011101114000000232 T. apionus 7100001111030111102211012110002334 T. undulatoides 710000110103011111221??12?0??????? T. rotundus 3100000101022120011050101012002511 T. scintillatus 31?0000101022231112230114212?02511 T. cyclopeus 3100010101032231112230114202002511 T housemani 3100000101032231112230111202002501 T. unicus 31000001010222311121301??1?200251? T. interformus 3100000101030130110030101001012021 T. pinaultae 3100010101020010110010001001?02?3? T. uyucus 3100000101022010110010?01101012?11 T. nasutus 31001101010300111111100??0?101001? T. echinatus 3100000101030010112110?01001000011 T. mexicanus 0100000101000000101110001010200142 T. hispidus 0100000101000010101010201010200142 T. huitepecensis 610001010100000011100000101020224? T. paramexicanus 010100010100000010111??01?1??????2 T. altus 010000010100000000100??01?40?????2 T. xicotepecensis 01000101010001011031301??0?020?24? T. singulus ?1000001010301211131?00??1?020?14? T. prolixus 6100010101020000011100000010002257 T. cozari 6100010101020000011100001000022250 T. quetzalensis 61000101010321001122100??0?002025? T. tuberosus 7100010101030131103270001110101141 T. perezi 71?1000101030130112270001110020033 T. imbricatus 1111010101030131112170001000120247 T. planus 1100010111030111101170001110001047 T. glochinos 111101010103013?10227??01?1??????8 T. implanus 011100010103013111227??03?1??????7 T.cerasquamatus 0111010101030131112170101100011347 T. lepros 611101010103013?11327??01?0??????3 T. rowei ?111010101030120112170001100120041 T. multituberculatus 111101010103014011227??01?1??????6 T. baranowskii 6110010101030131111170001000120047 T. opacus 1110000101030121111170001110001447 T. elsalvadorensis 611101010103013?112170011240021147 T. denticulatus 61100101010301311121700??1?002114? T. jakei 61100001010301301132700??2?002133? T. oblongus 1110010101010111012120011100101737 T. gracilis 6111010101030111011120014100021446 T. smithi 1110010101030131011120011100021146 T. annulus 1111010101030111010180011140?11048 T. diabolus 1110010101030121011180011000?13842 T. incus 6110010101030111011180111010?14640 T. constrictus 1110010101030111012120011100120200 T. tajumulcoensis 1100010101030001011180?14210021346 T. armentivus 1111000101030111011120000000021146 T. lempirensis 1111000101030121011120010000000246 T. confusus 1111010101030121012120011110001146 T. inflatus 1110000101030121011120011440020220 T. undulatus 110000010103012001212001010002124? T. pseudoundulatus 1110000101030130012120012000020147 T. quadratus ?1?101010103010111221??04?1??????3 T. plumosus 51100101010300011111100??0?000033? T. ashei ?1110101010300301121???02?0??????8 T. alternus 710000010103013111112??01?1??????2 T. colemanae 11110?010103011011110??01?0??????7 T. alternatus ?10001010103000111100??01?0??????4 T. sharpi 91000101010200001111100??1?002023? T. solitarius ?10001010102000011000??11?2??????0 T. biprotuberatus ?1110101010301401121?0002440020138 T. andersoni ?111010101030111011110000000002248 T. howdenorum 710001010103010011221000?100200037 T. genieri ?11001011103010111221000?000000034 T. hondurensis ?10000010103011001110??02?0??????6

63

64

65

66

67

FIGURES 33-36. Ventral (left) and lateral (right) view of the male genitalia of the Trachyphloeomimus championi species group (Scale bar measurements in parentheses): (33) T. apionus (0.2 mm); (34) T. arcuatibius (0.5 mm); (35) T. batmobile (0.54 mm); (36) T. bravoensis (0.54 mm).

68

69

FIGURES 37-40 Ventral (left) and lateral (right) view of the male genitalia of the Trachyphloeomimus championi species group (Scale bar measurements in parentheses): (37) T. calcaritibius (0.38 mm); (38) T. carinatus (0.42 mm); (39) T. championi (0.52 mm); (40) T. festivipenis (0.32 mm).

70

71

FIGURES 41-43. Ventral (left) and lateral (right) view of the male genitalia of the Trachyphloeomimus championi species group (Scale bar measurements in parentheses): (41) T. jonesi (0.38 mm); (42) T. obrieni (0.38 mm); (43) T. popeyeus (0.4 mm).

72

73

FIGURES 44-59. Female spermatheca of the Trachyphloeomimus championi species group: (44) T. apionus; (45) T. arcuatibius; (46) T. batmobile; (47) T. bravoensis; (48) T. calcaritibius; (49) T. carinatus; (50) T. championi; (51) T. discus; (52) T. festivipenis; (53) T. heliconodulus; (54) T. jonesi; (55) T. junctus; (56) T. obrieni; (57) T. popeyeus; (58) T. spurcus; (59) T. undulatoides.

74

75

76

77

Strict consensus of 1000 trees (0 taxa excluded) Slatipennis Cchrysorrhoeus Ckomareki Tsolitarius Tcozari Tprolixus Thuitepecensis Tjonesi Tspurcus Txicotepecensis Taltus Thispidus Tparamexicanus Tmexicanus Tsharpi Talternatus Tplumosus Tnasutus Techinatus Tuyucus Tpinaultae Tinterformus Trotundus Thousemani Tcyclopeus Tunicus Tscintillatus Tquetzalensis Thowdenorum Tundulatoides Tapionus Theliconodulus Tdiscus Tjunctus Tcarinatus Tfestivipenis Tarcuatibius Tcalcaritibius Tchampioni Tpopeyeus Tobrieni Tbatmobilensis Tbravoensis Tgenieri Tquadratus Tjakei Telsalvadorensis Tmultituberculatus Trowei Tlepros Tcerasquamatus Timplanus Tglochinos Tperezi Tbiprotuberatus Tashei Tbaranowskii Timbricatus Tdenticulatus Tplanus Ttuberosus Talternus Topacus Tsingulus Tsmithi Tgracilis Ttajumulcoensis Tincus Tdiabolus Tannulus Thondurensis Tcolemanae Tarmentivus Tandersoni Tlempirensis Tconfusus Toblongus Tconstrictus Tinflatus Tpseudoundulatus Tundulatus Figure 68. Phylogenetic relationships of species of Trachyphloeomimus: Strict consensus tree of 1000 most parsimonious trees of length 368 for 34 unweighted characters. Arrows show placement of species within the T. championi species group. 78

Strict consensus of 1000 trees (0 taxa excluded) Slatipennis Cchrysorrhoeus Ckomareki Tsolitarius Thuitepecensis Txicotepecensis Taltus Thispidus Tparamexicanus Tmexicanus Tcozari Tprolixus Tsharpi Talternatus Tplumosus Techinatus Tnasutus Tuyucus Tpinaultae Tinterformus Trotundus Tunicus Thousemani Tcyclopeus Tscintillatus Tquetzalensis Thowdenorum Tundulatoides Tapionus Tcarinatus Theliconodulus Tdiscus Tfestivipenis Tarcuatibius Tobrieni Tpopeyeus Tbatmobilensis Tbravoensis Tjunctus Tcalcaritibius Tchampioni Tjonesi Tspurcus Tgenieri Tquadratus Tjakei Telsalvadorensis Tmultituberculatus Trowei Tlepros Tcerasquamatus Timplanus Tglochinos Tperezi Tbiprotuberatus Tashei Tbaranowskii Timbricatus Tdenticulatus Tplanus Ttuberosus Talternus Topacus Tsingulus Tsmithi Tgracilis Ttajumulcoensis Tincus Tdiabolus Tannulus Thondurensis Tcolemanae Tarmentivus Tandersoni Tlempirensis Tconfusus Toblongus Tconstrictus Tinflatus Tpseudoundulatus Tundulatus Figure 69. Phylogenetic relationships of species of Trachyphloeomimus: Strict consensus tree of 1000 most parsimonious trees of length 385 for 34 weighted characters. Arrows show placement of species within the T. championi species group. Note shift in placement of T. spurcus and T. jonesi species. 79 APPENDIX I. Trachyphloeomimus championi group: Included species (in alphabetical order).

Trachyphloeomimus apionus Horsley, new species Mexico (Oaxaca) Trachyphloeomimus arcuatibius Horsley, new species Mexico (Oaxaca) Trachyphloeomimus batmobile Horsley, new species Mexico (Oaxaca) Trachyphloeomimus bravoensis Horsley, new species Mexico (Puebla) Trachyphloeomimus calcaritibius Horsley, new species Mexico (Oaxaca) Trachyphloeomimus carinatus Horsley, new species Mexico (Oaxaca) Trachyphloeomimus championi O’Brien Mexico (Mexico D.F., Veracruz) Trachyphloeomimus discus Horsley, new species Mexico (Oaxaca) Trachyphloeomimus festivipenis Horsley, new species Mexico (Oaxaca, Querétaro, Veracruz, (Chiapas)) Trachyphloeomimus heliconodulus Horsley, new species Mexico (Oaxaca) Trachyphloeomimus jonesi Horsley, new species Mexico (Querétaro) Trachyphloeomimus junctus Horsley, new species Mexico (Oaxaca) Trachyphloeomimus obrieni Horsley, new species Mexico (Oaxaca) Trachyphloeomimus popeyeus Horsley, new species Mexico (Oaxaca) Trachyphloeomimus spurcus Champion Mexico (Guanajuato, Mexico D.F., Morelos, Puebla, Querétaro, San Luis Potosi, Tlaxcala, Veracruz) Trachyphloeomimus undulatoides O’Brien Mexico (Mexico D.F., Puebla, Veracruz)

80 APPENDIX II. Characters and character states used in the phylogenetic analysis of Trachyphloeomimus species and related taxa.

Character 0: Vestiture, scale type – Ten states of this character are recognized: 0. striae faint to indistinguishable, sometimes shiny and round to irregular in shape 1. oval striate 2. coarsely striate 3. metallic and stellate 4. with smooth copper scales on middle of pronotum, remaining scales flat and without setae 5. plumose 6. pectinate 7. flat and appressed 8. none present 9. erect, non-metallic, white and stellate

Character 1: Rostrum, scrobe – Three states of this character are recognized: 0. wide and shallow, with scales 1. narrow and moderately deep, without scales 2. narrow and deep, without scales, directed basally

Character 2: Rostrum, nasal plate – Three states of this character are recognized: 0. not protruding, flat 1. distinctly protruding 2. sutural line distinct

Character 3: Rostrum, apex (lateral) – Two states of this character are recognized: 0. nearly flat 1. cuts off abruptly near antennal insertion, declivious

81 Character 4: Rostrum, apex (dorsal) – Two states of this character are recognized: 0. nearly straight to weakly expanded 1. distinctly expanded and forming a flange

Character 5: Antennae, scape – Two states of this character are recognized: 0. evenly expanded from base to apex, clavate 1. very thin at base, expanded at apex, sinuate

Character 6: Antennae, funicle – Two states of this character are recognized: 0. 7th segment longer than wide 1. 7th segment as wide or wider than long

Character 7: Antennae, club – Two states of this character are recognized: 0. cylindrical and segmented 1. ovate

Character 8: Antennae, overall form – Two states of this character are recognized: 0. normal 1. short and stout

Character 9: Head, eye form – Two states of this character are recognized: 0. small 1. large

Character 10: Pronotum, basal margin – Three states of this character are recognized: 0. not modified 1. modified, with low, wide half-circle 2. modified, with distinct triangular area

Character 11: Pronotoum, shape – Four states of this character are recognized: 0. lateral margins nearly parallel with width greater than or equal to length

82 1. lateral margins nearly parallel with length greater than width 2.nearly round 3. sub-cordate

Character 12: Pronotum, width relative to humeri of elytra – Three states of this character are recognized: 0. approximately equal 1. smaller 2. larger

Character 13: Elytra, humeral shape – Four states of this character are recognized: 0. rounded 1. angular 2. pointed 3. not pronounced

Character 14: Elytra, 8th interval projection – Five states of this character are recognized: 0. hardly pronounced 1. weakly expanded 2. strongly expanded, broad 3. strongly expanded, pointed 4. double expansion

Character 15: Elytra, setae distribution – Three states of this character are recognized: 0. more or less on all intervals 1. mainly on odd intervals 2. on all intervals

Character 16: Elytra, setae type – Three states of this character are recognized: 0. straight, sometimes with acuminate apex

83 1. straight or spatulate on dorsal surface, spatulate on declivity 2. tear-drop shape

Character 17: Elytra, setae elevation – Two states of this character are recognized: 0. recumbent 1. sub-erect to erect

Character 18: Elytra, elevation of intervals – Four states of this character are recognized: 0. flat 1. odd raised 2. weakly uneven 3. small tubercles all over surface, uneven appearance

Character 19: Elytra, 3rd interval – Three states of this character are recognized: 0. flat at base 1. weakly raised at base 2. distinctly raised at base

Character 20: Eyltra, declivital tubercles – Nine states of this character are recognized: 0. none present to very weak on interval 5 1. on interval 5 2. on intervals 5 and 6 3. on intervals 1, 3, 5(x2) 4. on interval 5, convergent across interval 4 5. on interval 5(x2) 6. on intervals 3 and 5 7. on interval 6 and at least 2 more different intervals 8. on intervals 1 and 5

Character 21: Legs, hind tibia of male – Two states of this character are recognized: 0. more or less straight, with or without denticles on inner face

84 1. modified, inner face arcuate, with flange or tooth

Character 22: Legs, inner legs of male – Three states of this character are recognized: 0. with setae approximately the same length to only slightly longer 1. with setae approximately 1/2 length to the same length as width of femur 2. with setae bristle-like in form on inner femora only

Character 23: Legs, hind tibia of female – Two states of this character are recognized: 0. without denticles 1. with denticles

Character 24: Abdomen, female pygidium – Five states of this character are recognized: 0. truncate 1. rounded 2. weakly emarginate 3. strongly emarginated or notched 4. rounded acuminate

Character 25: Abdomen, 5th sternite of male – Five states of this character are recognized: 0. nearly flat 1. weakly medially depressed 2. distinctly medially depressed 3. convex 4. apically depressed, basally convex

Character 26: Abdomen, 5th sternite of female – Five states of this character are recognized: 0. nearly flat 1. weakly medially depressed 2. distinctly medially depressed

85 3. convex 4. apically depressed, basally convex

Character 27: Male aedeagus, dorso-ventral body form, lateral – Three states of this character are recognized: 0. not distinctly robust 1. moderately robust 2. distinctly robust

Character 28: Male aedeagus, lateral margins, ventral view – Three states of this character are recognized: 0. more or less parallel-sided from base to sub-apical region 1. convergent from base to sub-apical region 2. divergent from base to sub-apical region

Character 29: Male aedeagus, ratio of body length to aedeagal arms – Three states of this character are recognized: 0. approximately the same length 1. body longer than aedeagal arms 2. body shorter than aedeagal arms

Character 30: Male aedeagus, body apex, lateral view – Five states of this character are recognized: 0. straight 1. curved up 2. curved down 3. twisted 4. curved up with apex swollen

86 Character 31: Male aedeagus, body apex, ventral view – Nine states of this character are recognized: 0. rounded 1. rounded with small acuminate point 2. acuminate 3. acuminate, drawn out to a rounded or truncate point 4. acuminate, drawn out to a point 5. more or less truncate 6. circular 7. spade-like 8. asymmetrical

Character 32: Male aedeagus, internal sac sclerotizations – Seven states of this character are recognized: 0. absent 1. present, moderate to large in size, very globular to somewhat elongate in form, located in mid-body to just apical of mid-length 2. present, small size, located apical to mid-length 3. present, elongate form, apex not recurved, with or without concentrations of small spicules 4. present, elongate form, apex recurved, with or without randomly distributed spicules of small to large size 5. present, elongate form, extremely long and thin 6. present, elongate form, base recurved

Character 33: Female spermatheca, shape – Three states of this character are recognized: 0. nodulus and ramus not extruding, ramus moreso prounounced 1. F-shape, nodulus often much longer than very small ramus and curves out weakly to distinctly

87 2. nodulus and ramus approximately equal in length, short and form ball-shape together, pointed outward, somewhat like the form of “pac-man” 3. triangular-shape, ramus and nodulus equal in length, divergent and pointing up 4. plug-shape, nodulus and ramus of many relative lengths, but nodulus never more than 2x length of ramus, both point outwards 5. hook-shape, nodulus and ramus nearly the same length and touching basally 6. nodulus very small and thin, directly off of base of larger ramus 7. ramus very small, directly off base of larger nodulus 8. both very short and point up

88