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W h eeler , Q u e n t in D u a n e
COMPARATIVE MORPHOLOGY, CLADISTICS, AND A REVISED CLASSIFICATION OF THE GENERA OF LYMEXYLIDAE (COLEOPTERA), INCLUDING DESCRIPTIONS OF TWO NEW GENERA
The Ohio State University PH.D. 1980
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200 \ Z = = = =0. A\'N A330S Vli -S '06 '312: 761-1700 COMPARATIVE MORPHOLOGY, CLADISTICS, AND A REVISED
CLASSIFICATION OF THE GENERA OF LYMEXYLIDAE (COLEOPTERA),
INCLUDING DESCRIPTIONS OF TWO NEW GENERA
DISSERTATION
Presented in Partial Fulfillment of the Requirements for
the Degree Doctor of Philosophy in the Graduate
School of The Ohio State University
By
Quentin D. Wheeler, B .S ., M.S.
*****
The Ohio State University
1980
Reading Committee: Approved By:
Dr. David J. Horn
Dr. Donald E. Johnston
Dr. Gordon R. Stairs A dviser Dr. Charles A. Triplehorn Department of Entomology To my w ife Kim, who understands
i i . ACKNOWLEDGMENTS
A number of people have participated in the successful completion of this project. In all fairness, however, my thanks need be extended to include those who added both materially or philosophically to this study directly and those who gave me encouragement and support during my previous investigations in insect systematics. I will make an effort to thank each person who has substantially contributed to my personal progress and that of my study of Lymexylidae. I apologize to any which are inadvertantly omitted.
Dr. Charles A. Triplehom, my academic adviser for this study and my undergraduate and M.S. studies in entomology, was among the first scientists to inspire my interest in the study of insects. He has proven to be a constant source of sound advice, and an oasis of un wavering support during my years at The Ohio State University.
Dr. Ginter Ekis served as co-adviser during initial work on this project. I am grateful for his support and help, and particularly" for making me aware of many resources - espcially people - around the country and around the world, and for his participation in various discussions on systematics.
A group of colleagues, and friends, which I collectively refer to as the "Ball School" have been of inestimable influence and encourage ment through personal interactions and their outstanding contributions
iiiv. to the literature on beetle taxonomy and theoretical systematics. This
elite group of scientists includes George E. Ball (University of
Alberta, Edmonton), Terry L. Erwin (National Museum of Natural History,
Smithsonian Institution, Washington), David H. Kavanaugh (California
Academy of Sciences, San Francisco), and Donald R. Whitehead (USDA,
Systematic Entomology Laboratory, Smithsonian Institution, Washington).
Lee H. Herman, Jr. (American Museum of Natural History, New York)
gave advice and shared his ideas and methods, providing me with
suggestions which were both constructive and inspiring during visits to the American Museum.
Some particularly helpful colleagues include Stewart B. Peck
(Carleton University, Ottawa), J. Milton Campbell (Biosystematics
Research Institute, Ottawa), Edward C. Becker (B.R.I., Ottawa), Alfred
F. Newton, Jr. and Margaret Thayer (Museum of Comparative Zoology,
Harvard University, Cambridge), John F. Lawrence (CSIRO, Division of
Entomology, Canberra City and formerly of the M.C.Z.), Vicki Funk
(O.S.U., Department of Botany), John LaDuke (O.S.U., Botany), and
Larry Watrous (O.S.U., Department of Entomology).
Ronald B. Madge (British Museum of Natural History, London), very generously shared his insights into lymexylid systematics, including unpublished results of personal research, in an exemplary scientific s p i r i t .
John Nagy prepared most of the habitus drawings of lymexylid beetles in this paper. I commend John on his talent, eye for detail, and careful work; and admire the quality of these drawings, his first
efforts in illustrations of insects. Glen Berkey (Ohio Agriculatural
Research § Development Center, Wooster) prepared excellent reductions
for many plates, and David Stutes is responsible for operation of the
scanning electron microscope and the sem micrographs which are included
in this study.
Support for this study of the Lymexylidae was gathered from
several sources.
I enjoyed full-tim e support for my research during the year 1980
while I was a Presidential Fellow of The Ohio State University.
Larry Watrous (O.S.U., Entomology) and I were awarded a grant from
the Ohio Biological Survey which funded the construction and field
testing of a modification of the flight-intercept trap for beetles.
We were granted permission to test our traps at Cedar Bog and the
Wahkeena Nature Preserve by the Ohio Historical Society.
I was awarded funds for field study of lymexylids in Costa Rica by the Graduate School of The Ohio State University through a Graduate
Student Alumni Award.
The Environmental Sciences Program of the Smithsonian Institu tion supported studies in Panama in 1978 (funds were awarded to Dr.
Terry L. Erwin for studies on carabid beetles of Barro Colorado Island)
The Smithsonian Tropical Research Institute permitted use of the B.C.I. f a c i l i t y .
The Department of Entomology of The Ohio State University has generously supported my research activities for several years. In
v. addition to space for office and laboratory work it has acquired equipment and supplies, accepted page charges for several publications, and paid for travel associated with field work, museum and library studies, and participation in professional society meetings. The latter support came, in part, from the Osborn Fund, the Knull Fund, and the Department's budget: Thanks.
Taxonomic research is heavily dependent upon museum curators opening their hearts and cabinets. I have been able to borrow and study specimens from most parts of the world, and am indeed grateful to each individual and institution which made these loans possible.
These persons and collections are listed in Table 1, along with acronyms which I use in the text in reference to these depositories. VITA
January 3 1954 Born- M ary sv ille, Ohio
1972 High School Diploma, Marysville Senior High School, Marysville, Ohio
1973 Laboratory Assistant, Insect Pathology Laboratory, Department of Entomology, The Ohio State University, Columbus, Ohio
1973 Insect Taxonomist, WHO c/o Insect Pathology Lab., Dept. Entomology, The Ohio State University, Columbus
1974 Field Entomologist, Vector-borne Disease Unit Dept. Health, State of Ohio, Columbus
1975 ' Insect Taxonomist, USDA Forest Service, N.E. Rsch. Lab., Delaware, Ohio
1976 B.S. in Entomology, The Ohio State University, Columbus
1976-1977 Teaching Associate, Dept. Entomology, The Ohio State University, Columbus
1975-1976 Curatorial Assistant, The Ohio State University, Collection of Insects and Spiders, Columbus
1976-1977 Assistant Extension Entomologist, Ohio Cooperative Extension Service, The Ohio State University, Columbus
1976,1978 1979 Research Associate, Dept. Entomology, The Ohio State University, Columbus
1977 M.S. in Entomology, The Ohio State University, Colum bus (Thesis: Revision of the genus Anisotoma of North and Central America)
1978 Research Assistant, Smithsonian Institution, Environ mental Sciences Program funds awarded to Dr. T.L. Erwin, National Museum, Dept. Entomology AT Barro Colorado Island facility of Smithsonian Tropical
v i i . Research I n s titu te , Balboa, Republic o f Panama
Presidential Fellow, The Ohio State University, Columbus
PROFESSIONAL APPOINTMENTS
Assistant Professor of Entomology, Cornell University, Ithaca, New York (appointment begins September 1, 1980)
PROFESSIONAL PAPERS AND SEMINARS
"Classification and phylogeny of the Anisotoma slime- mold beetles", Entomological Society of America, Houston, Texas (November)
"Hennigean phylogenetic systematics and classification of the leiodid beetles, or, what do pigs, beetles, and fungi have in common?", Departm ental Seminar, Department of Entomology, The Ohio State University, Columbus (October)
"Evolutionary trends in the ship-timber beetles, Lymexylidae", Entomological Society of America, North-Central Branch, Indianapolis (March)
"It's a jungle out there!, a travelog of Panama", Graduate Student Seminar Series, Department of Entomology, The Ohio State University, Columbus (A pril)
"Principles of cladistics" (co-speaker with L.E. Watrous), Graduate Student Seminar Ser., Dept. of Entomology, The Ohio State University, Columbus (May)
v i i i . "Slime molds and beetles: perspectives in evolution" Department of Entomology, Cornell University, Ithaca (October)
SPECIAL PROFESSIONAL ACTIVITIES
Co-moderator for discussion group titled "Modern trends in systematics: principles and uses of cladistic analysis", Entomological Society of America, North-Central Branch, Indianapolis (March) (co-moderator, L.E. Watrous)
Invited speaker for symposium "Cladistic techniques: methods and applications", Thomas Henry, moderator. Title of talk: "Are in-groups out?". (Speakers: W.Mathis, T.Schuh, J.Slater, Q.Wheeler), Entomolo gical Society of America, Eastern Branch, B ltimore (September)
GRANTS AND AWARDS
Graduate Student Alumni Award, The Graduate School The Ohio State University, joint award with L.E. Watrous of $1600 for field studies in Costa Rica
Presidential Fellowship of the Ohio State University
Grant, Ohio Biological Survey, $500 awarded jointly with L.E. Watrous for study of efficacy of a new design for a flight-intercept trap
Elected to full membership in Sigma-Xi,The National Research Society, Ohio State University Chapter FIELD STUDIES
1974 1980 Eastern United States and Canada (especially, Ohio Kentucky, Pennsylvania, Connecticut, New York, North Carolina, and Ontario); berlese samples and fungus/ insect collections
1975 Mexico (various localities § habitats); sift leaf litter and fungus/insect collections
1978 Panama (central and western, especially lowland, tropical forest and montane cloud forest; study sites included Volcan de Chiriqui, Cerro Jefe, Cerro Campana, Gamboa area, and Barro Colorado Island) sift leaf litter and fungus/insect collections
1980 Cedar Bog and Wahkeena Nature Preserves (Protected by Ohio Historical Society); test of efficacy of new design of flight-intercept trap funded by Ohio Biological Survey
PROFESSIONAL SOCIETY MEMBERSHIPS
American Entomological Society
Entomological Society of America
Society of Systematic Zoology
Coleopterists* Society
Mycological Society of America
North American Beetle Fauna Project (contrib. author)
x. PUBLICATIONS
Triplehorn, C.A. and Q.D. Wheeler. 1979. Systematic placement and distribution of Uloporus ovalis Casey (Coleoptera: Heteromera: Archeocrypticidae). Coleopterists Bulletin 33: 245-250.
Watrous, L.E. and Q.D. Wheeler. 1980. Out-group comparisons and character analysis. Systematic Zoology: in press.
Wheeler, Q.D. 1977. A new species of Isoplastus from Mexico (Coleo ptera: Leiodidae). Coleopterists Bulletin 31: 77-78.
______. 1977. Placement of Anisotoma fenderi Hatch (Coleoptera: Leiodidae). Entomological News 8 8: 137-138.
______. 1978. Australasian species of genus Leiodes Latreille (Coleoptera: Leiodidae). Entomological News 89: 63-64.
______. 1979. Slime mold beetles of the genus Anisotoma (Leio didae): classification and evolution. Systematic Entomology 4: 251-309.
______. 1979. Revision and cladistics of the Middle American genus Creagrophorus Matthews (Coleoptera: Leiodidae). Quaestiones Entomologicae 15: 447-479.
______. 1980. Studies on Neotropical slime-mold/beetle relation ships, Part I: Natural history and description of a new species of Anisotoma from Panama (Coleoptera: Leiodidae). Proceedings of the Entomological Society of Washington 82: 493-498.
______. 198_. Diagnosis and phylogenetic relationships of the monotypic genus Stetholiodes (Coleoptera: Leiodidae). Ohio Jour nal of Science: submitted.
______. 198_. An hypothesis on the evolution of slime-mold feeding in leiodid beetles, in preparation. TABLE OF CONTENTS
ACKNOWLEDGMENTS...... - ...... —i i i
VITA------...... - ...... v ii
LIST OF TABLES------xvi
LIST OF FIGURES—...... -x v ii
INTRODUCTION------...... - ...... 1
MATERIALS AND METHODS— ------—...... 10
M a te ria ls------10
Procedural Methods ------10
Philosophical Methods------14
HISTORICAL REVIEW — ------...... - ...... 18
Description of Species ------20
Description of Genera— : ------■------21
Description of Larvae ------23
Super fam ilial Status ------28
NATURAL HISTORY------—------29
ECONOMIC IMPORTANCE ...... - ...... - ...... — 38
EXTERNAL MORPHOLOGY OF HYLECOETUS------41
A dult ------42
Larva ------117
Pupa------133
x i i . TAXONOMIC CONS I DERATIONS— ------...... 136
Family Lymexylidae ------*------— 137
D iagnosis ------138
Key to Subfamilies and Genera of Lymexylidae ------139
Genus H ylecoetus ------143
D iagnosis ------146
Description ------147
Key to Species ------148
H. dermestoides ------151
H. lu g u b ris ------157
Genus Lymexylon ------163
D iagnosis ------166
Description (L. navale) ------167
Genus A tra c to c e ru s------210
D iagnosis ------213
D escrip tio n (A.bre v ic o rn is ) ------214
Genus Australymexylon, new genus ------300
D iagnosis ------303
Key to Species ------304
D escrip tio n (A. au s tr a le ) ------306
A. fu sc ip e n n is ------312
x i i i . Melittomma-Complex ------361
Key to Genera of Melittomma-Complex ------365
Genus Melittomma ------366
M. brasiliense ------369
M. sericemn ------376
Genus Melanomelittomma, new genus ------380
M. insulare, new combination ------.380
Genus M elittom m opsis ------386
Genus Hylecoetopsis, Incertae Sedis ------390
CLADISTIC ANALYSIS—...... ------477
Character State Analysis------477
Monophyletic Groups ------503
Classification of Lymexylidae ------:------507
SPECULATIONS ON THE ORIGIN OF LYMEXYLOIDEA ...... - — 508
ZOOGEOGRAPHY— ------...... —...... 531
The Vicariant Model: An Introduction ------531
Distribution Patterns for Lymexylidae: The Data and Some
Comments ------533
Speciation Patterns: A Subjective Analysis— ------534
Historical Zoogeography: A Preliminary Analysis------535
REFERENCES------— 559
APPENDIX A: MISCELLANEOUS OBSERVATIONS ON LYMEXYLIDAE— ...... 542
APPENDIX B: SUPPLEMENTARY ILLUSTRATIVE MATERIAL ...... 551
APPENDIX C: ANOTHER RESPONSE TO E. KING-— 556
x iv . LIST OF TABLES
Table Page
1. Collections from which specimens of Lymexylidae were studied, curators responsible for making loans, and acronyms used in the text to refer to these collections ___
2. Descriptions of larvae of Lymexylidae ...... 24
3. Host information for lymexylid beetles ...... 32
4. Hindwing venation systems of terminology compared with terms used here ...... 56
5. Terminology for female genitalia and associated structures. See Figs. 53-55 ...... 64
6 . Terminology for male genitalia (aedeagus) and related postabdominal structures. Figs. 57-59 ...... 70
7. A checklist of the subfamilies and genera of Lymexylidae. See cladistics section for a discussion of relationships of the taxa...... - ...... 142
8 . List of the species of Hylecoetus, with new synonymies 150
9. List of species transferred from Hylecoetus ...... 158
10. List of described species of Lymexylon ...... 177
11. The described species of the genus Atractocerus. See also discussion in text, and references to Atractocerus in Appendix A ...... 225
12. List of the species of AustralymexyIon, NEW GENUS, including new synonymies ...... 305
13. List of species assigned to Melittomma, sensu novo ...... 367
14. List of species of Melanomelittomma, new genus ...... 381
15. List of species of Melittommopsis ...... 387
xv. Table Page
16. Character state polarities ...... 510
17. My interpretations of plesiomorphic and apomorphic states of characters of the metendosternite of Coleoptera, based on data and discussion of Crowson, 1944: 305 ...... 518
18. Phyletic sequence classification of Lymexylidae ...... 519
19. Characters of ancestral Cucujoidea proposed by R.A. Crowson (1955) ...... 528
20. Series of ancestral distributions indicated by cladogram CFig. 296) ...... 541
x v i. LIST OF FIGURES
Figure Page
1. Hylecoetus dermestoides, habitus, "flabellicornis" morph: head-on view (top), lateral view of forebody (bottom).... 2
2-5. Hylecoetus dermestoides, head: fig. 2, dorsal view; fig.3, ventral view; fig.4, posterior view; fig.5, lateral view. 72
6-9. Hylecoetus dermestoides, maxilla: fig. 6 , female; fig. 7, male; fig. 8 , male, palpomere III and bases of palpomere IV and maxillary palp-organ; fig. 9, apex of secondary (2o) branch of maxillary palp-organ ...... 74
10-12. Hylecoetus dermestoides: fig. 10, antenna; fig. 11, labrum (dorsal); fig. 12, labrum and epipharynx (ventral) 76
1-3-14. Hylecoetus dermestoides, maxillary palp-organ; fig. 13, outer surface; fig. 14, inner surface ...... 78
15-16. Hylecoetus dermestoides: fig. 15, hypopharyngeal setae; fig. 16, epipharyngeal setae ...... 80
17-18. Hylecoetus dermestoides: fig. 17, apex of labium (ven tral); fig. 18, antennomere IV, surface detail ...... 82
19-20. Hylecoetus dermestoides: fig. 19, apex of maxillary palpomere IV; fig. 20, apex of labial palpomere III 84
21-22. Hylecoetus dermestoides: fig. 21, "peg" sensilla at apex of labial palpomere III; fig. 22, "flabellicornis" morph, antenna ...... 86
23-24. Hylecoetus dermestoides, "flabellicornis" morph, flabell- ar process of antenna: fig. 23, outer surface; fig. 24, inner surface ...... 88
25-26. Hylecoetus dermestoides, "flabellicornis" morph: fig.25, apex of maxillary palpus; fig. 26, surface of maxillary p alp -o rg an ...... 90
27-28. Hylecoetus dermestoides: fig. 27, mandible (dorsal); fig. 28, maxillary palpomere IV ...... 92
x v ii. Figure Page
29-30. Hylecoetus dermestoides; fig. 29, antennomere VII surface; fig. 30, epicranial p it ...... 94
31-32. Hylecoetus dermestiodes, labium: fig. 31, lateral view; fig. 32, ventral view ...... 96
33-35. Hylecoetus dermestoides: fig. 33, pronotum, scutellum, and bases of elytra (dorsal1; fig. 34, prothorax (ven tral); fig. 35, prothorax (lateral) ...... 98
36. Hylecoetus dermestoides, mesosternum, metasternum, and associated structures (ventro-lateral view) ...... 99
37. Hylecoetus dermestoides, scutellum (metathoracic), dorsal view ...... 101
38-44. Hylecoetus dermestoides; fig. 38, prothoracic leg; fig. 39, apex protibia, inner margin; fig. 40, apex protibia, outer margin; fig. 41, mesothoracic leg; fig. 42, meta thoracic leg; fig. 43, claw; fig. 44, metatibia, apex and s p u r ...... 103
45 . Hylecoetus dermestoides, left elytron and scutellum ( d o rs a l) ...... 105
46. H ylecoetus derm estoides, hindwing (see Table 4 ) ...... 106
47-49. Hylecoetus dermestoides, metendosternite; fig. 47, lat eral view; fig. 48, dorsal view; fig. 49, ventral view... 107
50-52.- Hylecoetus dermestoides; fig. 50, female abdomen (later a l ) ; f ig . 51, male abdomen (lateral); f ig . 5 2 , fem ale abdomen, base (ventral). Abbreviations: T1-T9 (ter- gites I-IX), S1-S9 Cstemites I-IX), PI (pleural sclerite of abdominal segment I) ...... 109
53-55. Hylecoetus dermestoides, female genitalia (ovipositor); fig. 53, ventral view; fig. 54, lateral view; fig. 55, dorsal view ...... I l l
56. Hylecoetus dermestoides, stylus of female genitalia 113
57-59. Hylecoetus dermestoides, male genitalia (aedeagus) and abdominal segment IX: fig. 57, lateral view; fig. 58, dorsal view; fig. 59, ventral view ...... 115
x v iii. Figure Page
60. Hylecoetus dermestoides, larva (lateral view). Drawing by J . Nagy ...... 121
61-64. Hylecoetus dermestoides, larva: fig. 61, posterior end of abdomen (segments 7-10); fig. 62, tarsungulus of pro leg; fig. 63, proleg; fig. 64, antenna ...... 123
65. Hylecoetus dermestoides, larva, labrum and epipharyngeal region (ventral) ...... 125
6 6 . Hylecoetus dermestoides, larva, maxilla. Apex of stipes, palpus and mala (above) and base of stipes and cardo (below ) ...... 127
67. Hylecoetus dermestoides, larva, apex of mala showing par tial division of lacinia and gelea 128
6 8 . Hylecoetus lugubris, larva, mandible, left-mesal surface. Scanning electron micrograph courtesy of J.F. Lawrence, CSIRO, Division of Entomology, Canberra C ity ...... 129
69-70. Hylecoetus dermestoides, larva: fig. 69, prosthecal spines and molar region of mandible; fig. 70, left mandi ble, dorsal view ...... 130
71. Hylecoetus dermestoides, larva, labium (ventral) ...... 131
72. Hylecoetus dermestoides, pupa (ventral). Drawing by J. Nagy ...... 134
73. Hylecoetus dermestoides, habitus, dorsal view. Drawing by J . Nagy ...... 144
74-79. Hylecoetus spp., habitus drawings showing color patterns of species and morphs (dorsal view). Drawings are semi- diagramatic: fig. 74, H. dermestoides, "dermestoides" morph, male; fig. 75, H. dermestoides, "dermestoides" morph, female; fig. 76, H. dermestoides, "cossis" morph, female; fig. 77, H. dermestoides, "morio" morph, male; fig. 78, H. lugubris, male; fig. 79, H. lugubris, female. 159
80-81. Hylecoetus spp., apex of median lobe: fig. 80, dermes toides; fig. 81, lugubris 161
82. Lymexylon navale, habitus of male, dorsal view. Drawing by J.N agy...... 164
x ix . Figure Page
83-84. Lymexylon navale, head: fig. 83, dorsal view; fig. 84, ventral view ...... 178
85. Lymexylon navale, mandible of adult (dorsal) ...... 180
86-87. Lymexylon navale, male, maxilla: fig. 8 6 , apex of maxi llary palpus (palpomere IV); fig. 87, apex of secondary branch of maxillary palp-organ ...... 182
8 8. Lymexylon navale, male, apex of labial palpus ...... 184
89-90. Lymexylon navale, pronotum, dorsal view: fig. 89, male; fig. 90, female ...... 186
91-93. Lymexylon navale: fig. 91, hindwing, dashed lines indi cate vein present in male only (illustrated wing is of fe male); fig. 9Z, abdomen, basal segments (lateral); fig. 93, abdomen, apical segments (lateral) ...... 188
94-96. Lymexylon navale, metendosternite: fig. 94, dorsal view; fig. 95, base of furcal arm (ventral); fig. 96, laminal region (ventral). Abbreviations: FA- furcal arm; ATS- anterior tendon "stalk"; SK- furcal stalk; AT- anterior tendon; L- lam ina ...... 190
97. Lymexylon navale, male, maxilla ...... 192
98. Lymexylon navale, male, apex of abdomen (lateral) ...... 194
99-102. Lymexylon navale, male postabdomen: fig. 99, lateral lobes (ventral); fig. 100, abdominal sternite VIII (ven tral); fig. 1 0 1 , aedeagus (lateral); fig. 1 0 2 , aedeagus ( d o rs a l) ...... 196
103. Lymexylon navale, apex of median lobe and endophallus, lateral view ...... 197
104-108. Lymexylon navale, female genitalia: fig. 104, dorsal view; fig. 105, coxital fold (dorsal); fig. 106, ventral view; fig. 107, stylus; fig. 108, coxital fold (ventral). 199
109-110. Lymexylon navale, female genitalia: fig. 109, apices of valvifers and bases of coxites (dorsal); fig. 1 1 0 , median sclerite (dorsal). Abbreviations: cx- coxite; ms- median sclerite; pb- proctigeral baculus; v- valvifer ...... 201
xx. Figure Page
111-113. Lymexylon navale, larva, posterior end of abdomen: fig. Ill, segments VI-X; fig. 112, segments VIII-X; fig. 113, apex of segment IX. All views lateral; not drawn to scale ...... 2Q3
114. Lymexylon navale, larva, labrum (ventral) ...... 2Q5
115. Lymexylon navale, mesothoracic leg of larva: fig. 115.1, entire leg; fig. 115.2, apex of tibia and tarsungulus.... 2Q7
116. Lymexylon navale, larva: fig. 116.1, thorax and base of abdomen (lateral); fig. 116.2, maxillary palpus; fig. 116.3, mesal surface of mala (of maxilla); fig. 116.4, antenna; fig. 116.5, labium (ventral). Abbreviations: T1-T3 (thoracic segments I—III); A1-A3 (abdominal segments I-III); EPF (epipleural fold); mt (mentum); prm (pre- mentum); lg (ligula) ...... 209.
117. Atractocerus brevicornis, habitus, female, dorsal view. Drawing by J. Nagy ...... 212
118-119. Atractocerus brevicornis, female, head: fig. 118, dorsal view; fig. 119, ventral view ...... 229.
120. Atracrocerus brevicornis, male head and pronotum (dorsal) 230
121-123. Atractocerus brevicornis, female: fig. 121, apical seg ment of labial palpus; fig. 1 2 2 , recumbant s e n s illa on surface of labial palpomere III; fig. 123, antenna ...... 232
124-128. Atractocerus brevicornis, female maxilla: fig. 124, left maxilla, dorsal view; fig. 125, maxillary palpus and base of maxillary palp-organ (lateral); fig. 126, base of palpomere IV showing recumbant sensilla; fig. 127, apex of palpomere IV; fig. 128, apex of secondary branch of maxillary palp-organ ( 2° ) ...... 235
129. A tracto ceru s b re v ic o rn is , fem ale, m ala...... 237
130-132. Atractocerus brevicornis, male maxilla: fig. 130, palpo- meres III + IV and maxillary palp-organ (note that apices of most secondary branches are omitted from drawing); fig. 131, apex of palpomere IV; fig. 132, secondary branch of maxillary palp-organ ...... 239 Figure Page
133. Atractocerus brevicornis, prothorax (ventral view) ...... 241
134-135. Nota of mesothorax and metathorax: fig. 134, Lymex ylon navale; fig. 135, Atractocerus brevicornis ...... 243
136. Atractocerus brevicornis, right elytron (dorsal) ...... 245
137. Atractocerus brevicornis, mesostemum, metasternum, and associated structures (ventro-lateral view) ...... 247
138-140. Atractocerus brevicornis: fig. 138, male abdominal seg ment VIII (ventral); fig. 139, male abdominal segment IX (ventral); fig. 140, female abdominal segment VIII (ven tral). STS 8 TGS - struts ...... 249
141-142. Melittomma brasiliense: fig. 141, labium (ventral h ) ; f ig . 142, hypopharyngeal region (dorsal h ) ...... 251
143. Atractocerus brevicornis, larva, labrum (ventral) ...... 253
144. Atractocerus brevicornis, female genitalia: fig. 144.1, apex of stylus; fig. 144.2, coxital baculus and asso ciated structures. Abbreviations: cb- coxital baculus; vb- valviferal baculus; sb- spurious baculus ...... 254
145. Atractocerus brevicornis, aedeagus, ventral view. Abbre viations: LL- lateral lobe; TS- tegminal strut; TP- teg- minal plate; ML- median lobe ...... 256
146-148. Atractocerus brevicornis, larva: fig. 146, head, thorax, and base of abdomen (lateral); fig. 147.1, metathoracic leg; fig. 147.2, tarsungulus; fig. 148, apex of abdomen (lateral). Abbreviations: T1-T3 (thoracic segments), Al- A3 § A7-A10 (abdominal segments), an (anus), sp (spira cle) , trs (tarsungulus), tb (tibia), fm (femur), tro (trochanter), cx (coxa), epf (epipleural fold) ...... 258
149-150. Atractocerus brevicornis, larva: fig. 149, antenna; fig. 150, maxilla (mesal view) ...... 260
151. Atractocerus brevicornis, larva, m axilla..- ...... 262 Figure Page
152. Atractocerus brevicornis, larva, labium: fig. 152.1, hypopharyngeal setae; fig. 152.2, median, elevated hypopharyngeal region (dorsal); fig. 152.3, surface of elevated area; fig. 152.4, labium, ventral view ...... 264
153. Atractocerus brevicornis, larval head (dorsal view) 265
154-155. Atractocerus tasmaniensis: fig. 154, antenna (male), antennomeres 2-6; fig. 155, male maxillary palp-organ (and palpomeres II, III, and IV) ...... 267
156. Atractocerus brasiliensis, hindwing. Abbreviations: see ta b le 4 ...... 270
157. Atractocerus procerus, hindwing. Abbreviations: see ta b le 4 ...... 272
158-159. Atractocerus procerus, male: fig. 158, maxillary palp- organ; fig. 159, outer surface of maxillary palp-organ secondary branches ...... 274
160-161. Atractocerus procerus, male: fig. 160, inner surface of maxillary palp-organ secondary branches (2 ); fig. 161, surface of antenna ...... 276
162-163. Atractocerus procerus, elytron: fig. 162 (dorsal); fig. 163, "tubercles" at base ...... - ...... 278
164. Atractocerus procerus, aedeagus, ventral view. Abbre viations: ML- median lobe; LL- lateral lobes; TS- teg- minal strut; TG- tegminal plate ...... 280
165. Atractocerus procerus, aedeagus, dorsal view. Abbre viations: ML -median lobe; LL- lateral lobes; TG- teg minal p l a t e 282
166. Atractocerus procerus, aedeagus, lateral view. Abbre viations: 11- lateral lobe; ml- median lobe; t- tegmin a l p l a t e 284
167. Atractocerus sp. (ex Philippines), aedeagus (ventral).... 286
168-169. Atractocerus brasiliensis, male, apex of abdomen: fig. 168, lateral view; fig. 169, dorsal view ...... 288
x x i i i . Figure Page
170-171. Atractocerus tasmaniensis, metathoracic leg: fig. 170, tibia + tarsus; fig. 171, claw ...... 290
172-173. Atractocerus brevicornis, female, labium: fig. 172, lat- eral view; fig. 173, ventral view. Representative setae show relative sizes and densities, but many are omitted. Abbreviations: mn- mentum; pig- palpiger; pi- palpomere I; p2- palpomere II; lg- ligula; hyp- hypopharynx ...... 292
174. Atractocerus procerus, female, maxillary palpomere IV.... 294
175. Atractocerus tasmaniensis, head of male, dorsal view 295
176-177. Atractocerus brevicornis, metendostemite: fig. 176, ventral view; fig. 177, lateral view. Abbreviations: vf- ventral flange (of furcal stalk), 1 - lamina, fa- furcal arm...... 297
178. Atractocerus procerus, metendostemite ...... 299
179. Australymexylon fuscipennis, habitus of male, dorsal view. Drawing by J. Nagy ...... 302
180-181. Australymexylon australe, antenna: fig. 180, male; fig. 181, female ...... 315
182-183. Australymexylon fuscipennis, antenna: fig. 182, male; f ig . 183, fem ale...... 317
184. Australymexylon australe, hindwing. Abbreviations: see ta b le 4 ...... 3 1 S
185-186. Australymexylon australe, metendostemite: fig. 185, ven tral view; fig. 186, base of furcal arm and place of ant erior tendon insertion, ventral view 320
187-188. Australymexylon fuscipennis: fig. 187, female abdomen, lateral view; fig. 188, male postabdomen, dorsal view. Abbreviations: S2-S7 (sternites II-VII); T2-T7 (tergites II-VII); P2 (pleural sclerite of abdominal segment II)... 322
189-190. Australymexylon australe: fig. 189, male abdominal segment IK (lateral); fig. 190, apex of paratergal stm ts (ventral). Abbreviations: ST9- ninth sternite; Paraterg.9- ninth paratergite ...... 324
x xiv. Figure Page
191. Australymexylon fuscipennis, male, abdominal sternite VIII, ventral view ...... 326
192. Australymexylon australe, male, abdominal sternite VIII, ventral view ...... 323
193. Australymexylon australe, male, abdominal tergite VIII, dorsal view ...... 330
194. Australymexylon australe, female genitalia (ventral) 332
195. Australymexylon australe, coxite and stylus of female genitalia (ventral) ...... 334
196-197. Australymexylon fuscipennis, aedeagus: fig. 196, lateral view; fig. 197, dorsal view. Abbreviations: ML- median lobe, LL- lateral lobe, TS- tegminal strut, TG- tegminal p l a t e ...... 336
198-199. Australymexylon australe, aedeagus: fig. 198, lateral view; fig. 199, ventro-medial view of lateral lobe. Abbre viations: END- endophallus, LL- lateral lobe, TP- tegmin al plate, TS- tegminal strut, ML- median lobe 333
200. Australymexylon australe, aedeagus, ventral view. Abbre viations: ML- median lobe, LL- lateral lobe, TP- tegmin al plate, TS- tegminal strut ...... 340
201-202. Australymexylon australe, larva: fig. 201, apex of abdo men (lateral); fig. 2 0 2 , an ten n a 342
203. Australymexylon australe, larva, tarsungulus ...... 344
204. Australymexylon australe, geographic distribution ...... 345
205. Australymexylon fuscipennis, geographic distribution 346
206. Australymexylon australe, prothorax, ventral view ...... 348
207-208. Australymexylon australe: fig. 207, mesosternum, meta- stemum, and associated structures; fig. 208, anterior area of metasternum with mesocoxae removed. Abbreviations: CX2- mesocoxa, CX3-metacoxa, MTST- metasternum ...... 349
209. Australymexylon australe, apex of procoxa (note slo t).... 350 Figure Page
210. Australymexylon australe, male, maxilla (right, dorsal)... 3 5 2
211. Australymexylon australe, labrum (ventral). Epipharyngeal s e ta e o m itte d ...... 354
212-213. Australymexylon australe, labium: fig. 212, ventral view; fig. 213, dorsal view of hypopharynx ...... 355
214. Australymexylon australe, larva, maxilla, apex ...... 353
215-216. Australymexylon australe, larva: fig. 215, metathoracic leg; fig. 216, labium, left half (ventral) ...... 359
217. Melittomma brasiliense, male, habitus, dorsal view. Draw ing by J. Nagy ...... 353
218. Melittomma brasiliense, male, head (dorsal) ...... 392
219. Melittomma brasiliense, female, head (dorsal) ...... 394
220. Melittomma brasiliense, male, antenna ...... 3 g 6
221-222. Antennae: fig. 221, Melittomma brasiliense, female; fig. 222, Lymexylon navale, male.., t . 398
223-225. Melittomma brasiliense: fig. 223, prothorax (lateral);- fig. 224, pronotum (dorsal); fig. 225, elytron, right, dorsal view ...... 400
226-228. Melittomma brasiliense, male, maxilla: fig. 226, apex of palpomere IV, representative setae, sensilla, and surface detail only (other omitted); fig. 227, maxilla with maxi llary palp-organ; fig. 228, portion of tertiary branch (3°) of maxillary palp-organ (note short tactile setae, and long sensilla) ...... 402
229-230. Melittomma brasiliense, female genitalia: fig. 229, dorsal view; fig. 230, sclerotized process located between basal parts of coxites ...... 404
231-235. Melittomma brasiliense, male: fig. 231, aedeagus (later al); fig. 232, apex of paratergite of abdominal segment nine; fig. 233, aedeagus (ventral); fig. 234, aedeagus (dorsal); fig. 235, abdominal segments VIII and IX inter nested (ventral), note large, appendiculate paratergites of segment IX ...... 406
xxvi. Figure Page
236. Melittomma brasiliense, metendostemite ...... 408
237. Melittomma brasiliense, metepimeron and base of metacoxa (note groove which receives epipleural fold of elytron)... 410
238. Melittomma brasiliense, hindwing. Abbreviations: see table 4.. 7...... 412
239. Melittomma sericeum, hindwing ...... 414
240. Melittomma sp. (Brazil), hindwing ...... 416
241-242. Melittomma sericeum, male, habitus: fig. 241 (dorsal); fig. 242 (lateral view of forebody). Drawing by J. Nagy ...... 418
243. Melittomma sericeum, mesostemum, metastemum, and associ ated structures (ventro-lateral). Abbreviations: st2a- anterior part of mesosternum; st 2b- posterior part of meso- sternum; eps 2 a- elevated, anterior part of mesepisternum; eps 2b- posterior part of mesepisternum; epm 2 a- posterior part of mesepimeron; epm 2b- depressed part of mesepimeron which receives elytral epipleural fold; tn 2 - mesotrochantor; icp- intercoxal process; cx2- mesocoxa; eps3- metepisternum; epm3- metepimeron; cx3- metacoxa; s t3 - m etasternum ; tr 3 - metatrochantor; fm3- metafemur ...... 420
244. Melittomma sericeum, labium (ventral) ...... 422
245-248. Melittomma sericeum, male: fig.’ 245, aedeagus (lateral); fig. 246, spine on surface of paratergite IX; fig. 247, row of spines on mesal margin of paratergite IX; fig. 248, paratergite IX ...... 424
249-250. Melittomma sericeum, male: fig. 249, base of maxilla (NOTE: cardo removed, and all setae omitted); fig. 250, maxilla, including maxillary palp-organ. Abbreviations: ST- s tip e s , PLF- p a lp if e r , LC- la c in ia , GA- g a le a , P l- palpomere I, P2- palpomere II..., AXS- auxiliary sclerite, MPO- maxillary palp-organ, 1°- primary branch (MPO), 20- secondary branch ...... 426
251. Melittomma sericeum, larva, habitus, lateral view ...... 428
252-253. Melittomma sericeum, larva, proleg: fig. 252, mesal sur face; fig. 253, lateral surface 430
x x v ii. Figure Page
254-255. Melittomma sp. (New- Guinea}.; fig. 254, antenna;, fig. 255, apex of antennomere VIII ...... 432
256. Melanomelittomma insulare, new combination, female, habi tus (lateral view of forebody). Drawing by J.Nagy ...... 434
257. Melanomelittomma insulare, female, head (dorsal) ...... 436
258. Melanomelittomma insulare, male, aedeagus, lateral view... 438
259-261. Melanomelittomma insulare, male genitalia: fig. 259, aedeagus (ventral view); fig. 260, apex of lateral lobe (note row of spines); fig. 261, apex of endophallus. Abbre viations: LL- lateral lobe, ML- median lobe, TS- tegminal s t r u t , TP- tegm inal p l a t e ...... 440
262-263. Melanomelittomma insulare, metendostemite: fig. 262, api cal portion; fig. 263, base. Abbreviations: VLF- ventral flange, SK- furcal stalk, L- lamina, FA- furcal arm, AT- anterior tendon, ATS- anterior tendon stalk ...... 442
264. Melanomelittomma insulare, male, antenna ...... 444
265-266. Melanomelittomma insulare: fig. 265, mesal part of mandi ble (dorsal); fig. 266, apex of mandible ...... 446
267-268. Melanomelittomma insulare : fig. 267, maxilla, male; fig. 268, maxillary palpus, female ...... 448
269. Atractocerus brevicornis, female genitalia...... 450
270-271. Melanomelittomma insulare, female genitalia: fig. 270, coxite + stylus; fig. 271, stylus. Abbreviations: VB- valviferal baculus; CXB- coxital baculus; CF- coxital fold452
272. Melanomelittomma insulare, male, abdominal segment IX. Abbreviation: P9- paratergite of abdominal segment IX.... 454
273. Melanomelittomma in s u la r e , procoxa ( a p e x ) ..- ...... 455
274-275. Melanomelittomma insulare, female genitalia, proctigeral baculi (dorsal): fig. 274, basal ends (anterior); fig. 275, distal ends (posterior) ...... 457
x x v iii. Figure Page
276. Melanomelittomma insulare, apex of protibia ...... 459
277. Melanomelittomma insulare, pupa, head and pronotum flat-: e r a l ) ...... 461
278-279. Apex of larval abdomen: fig. 278, Melanomelittomma insu lare; fig. 279, Melittomma brasiliense ...... 463
280-282. Melittommopsis sp., female: fig. 280, maxillary palpus; fig. 281, apex of maxillary palpomere IV (corner); fig. 282, labial palpus ...... 465
283. Melittommopsis sp. (Brazil), mesostemum and associated sclerites. Abbreviations: ST2a- anterior part of meso stemum, ST2b- posterior part of mesostemum, EP2a- ant erior part (elevated) of mesepisternum, EP2b- posterior part of mesepisternum ...... 467
284-285. Melittommopsis sp., female: fig. 284, % apex of female genitalia; fig. 285, stylus ...... - ...... 469
286. Melittomma sericeum, male, apex of abdominal sternite VIII...... 471
287-288. Melittomma sericeum, female genitalia: fig. 287, coxites; fig. 288, distal part of coxite ...... 473
289. Melittomma sericeum, insertion of anterior tendon of metendostemite ...... 475
290. Hylecoetopsis cylindricus, antenna. Redrawn from Germer, 1912 ...... 476
291. Rate of description of Lymexylidae. Black dots- nominal species; open circles- nominal genera; triangles- species listed in most recent lists (including Zoological Record) some of which are known, but unresolved, synonyms. See text for discussion ...... 521
292. My concept (and illustration) of Crowson's scenario of the evolution of Lymexyloidea, Cleroidea, and Cucujoidea (Crowson, 1960). See text for discussion. Abbreviation: MYBP (millions of years before present)..- ...... 523
xxix. Figure Page
293. Cladogram of relationships among the genera of Lymex- ylidae. Numbers refer to characters (apomorphic states) in table 16 ...... 525
294. Cladogram and character matrix for Melittomminae. Numbers refer to apomorphic states in table 16. Abbreviations: AUST- Australymexylon; MELI- Melittomma; MELA- Melano melittomma; MBRA- Melittomma brasiliense (representative of soft-bodied Melittomma); MSER- M. sericeum; MOPS- Melittommopsis ...... 527
295. Distribution matrix for genera of Lymexylidae ...... 538
296. Geocladogram showing relationships among geographic areas indicated by biocladogram of genera of Lymexylidae. Re duced from fig u re 293 540
297. Flow-chart of systematics. Developed by L.E.Watrous and author. See text for discussion ...... 553
298. Flow-chart of cladistic methodology. Developed by L.E. Watrous and author. See text for discussion ...... 555
XXX. Table 1. Collections from which specimens of Lymexylidae were studied, curators responsible for making loans, and acronyms used in the text to refer to these collections.
Acronym C urator Collection/Institution
AMNH Dr. L.H.Herman, Jr. American Museum of Natural History, Central Park West at 79th St., New York, New York 10024, U.S.A.
ANIC Dr. J.F.Lawrence Commonwealth S c ie n tific § In d u s tria l Research O rganization, D ivi sion of Entomology, P.O.Box 1700, Canberra City, A.C.T. 2601, A u stra lia
BMNH Ms. C.M.F.von Hayek British Museum CNatural H istory], Department of Entomology, Crom Ms. J.M arshall w ell Road, London SW7 5BD, England
BPBM Dr. G.A.Samuelson Bernice P. Bishop Museum, Department of Entomology, P.O.Box 6037, Honolulu, Hawaii 96818, U.S.A.
CASC Dr. D.H.Kavanaugh California Academy of Sciences, Department of Entomology, Golden Gate Park, San Francisco, California 94118, U.S.A.
CNCI Dr. E.C,Becker Agriculture Canada, Biosystematics Research Institute, Coleoptera Dr. J.M.Campbell Section, Ottawa, Ontario K1A 0C6 Canada
CUIC Dr. L.L.Pechuman Cornell University, Department of Entomology, Ithaca, New York 14853, U.S.A.
DEMB Dr. H.Morge Deutsches Entomologisches Institut der Deutschen Akademie der Dr. R.Gaedike Landwirtschaftswissenschaften zu Berlin, 13 Eberswalde, SchicklerstaBe 5, D.D.R. Table 1. Collections (Cont'd.).
Acronym C urator _____ Collection/Institution
EIHU Dr. S.Takagi Entomological Institute, Faculty of Agriculture, Hokkaido Uni versity, Sapporo 060, Japan
FMNH Mr. H.Dybas Field Museum of Natural History, Divsion of Insects, Roosevelt Dr. R.Wenzel Road at Lake Shore Drive, Chicago, Illinois 60605, U.S.A. Dr. E.H.Smith
FASC Dr. R.E.Woodruff Florida State Collection of Arthropods, Florida Dept. Agriculture, P.O.Box 1269, Gainesville, Florida 32601, U.S.A.
HELS Dr. H.Silfverberg Entomological Department, Zoological Museum of the University, N. Jarnvagsgatan 13, SF-00100, Helsinki, Finland
ICCM Dr. G.Wallace Carnegie Museum of Natural History, Section of Insects § Spiders, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213, U.S.A.
INPA Dr. B.C.Ratcliffe Instituto Nacional de Pesquisas de Amazonia, Caixa Postal 478, 69.000, Manaus, Amazonas, Brazil
IRSN Dr. R.Damoiseau Institut Royal des Sciences Naturelles de Belgique, Rue Vautier, 31, B-1040. Bruxelles, Belgique (Entomologie)
KOBE Dr. O.C.Lomholdt Zoologisk Museum, Universitetsparken 15, DK 2100, Kobenhavn, Denmark
LACO Dr. C.L.Hogue Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, U.S.A. Table 1. Collections (Cont'd.)*
Acronym Curator Collection/Institution
LUND Mr. R.Baranowski Zoological Institute, Department of Systematics, Helgonavagen 3, S-223 62, Lund, Sweden
MCZC Dr. A.F.Newton,Jr. Museum of Comarative Zoology, Department of Entomology, Harvard Ms. M.Thayer University, Cambridge, Massachusetts 02138, U.S.A.
MHNG Dr. C.Besuchet Entomologie, Museum d'H istoire Naturelle, Route de Malagnou 1211, Dr. I.L obl Geneve 6, Switzerland
MNHN Dr. J.J.Menier Entomologie, Museum National d'H istoire Naturelle, 45 Rue de Buffon Dr. A.Villiers 75005 Paris, France
MOSC Dr. S.Keleinikova Zoological Museum, University of Moscow, Herzen str. 6, Moscow K-9, USSR
NEBR Dr. B.C.Ratcliffe Division of Entomology, W-539 Nebraska Hall, University of Nebraska -Lincoln, Univ. Nebr. State Museum, Lincoln, Nebraska 68508, U.S.A.
NMNH Dr. J.M.Kingsolver National Museum of Natural History, Smithsonian Institution, Dr. T.L.Erwin Department of Erttomology, Washington, D.C. 20560, U.S.A.
NMPR Dr. J.Jelinek National Museum in Prague,‘Musuem of Natural History, Department of Entomology, Prague, Kunratice 1, CSSR
NMVC Dr. A.Neboiss National Museum of Victoria, Entomology Department, 17 Victoria Crescent, Abbotsford, Victoria 3067, Australia 05 05 Stockholm, Sweden 104 B erlin , In v a lid e n str. 43, D.D.R. 2417, Kenedobu, Papua, New Guinea Zoologisches Museum, Museum fur Naturkunde der Humbolt-Unviersitet, Forest Insect Collection, Department of Primary Industry, P.O.Box Section for Entomology, Swedish Museum of Natural History, S-104 Postfach 417, Bergring 7, A-1014, Wien, Austria Collection/Institution South Africa Tervuren, Belgium Adelaide, South Australia 5000, Australia Avenue, Columbus, Ohio 43210, U.S.A. Dr. J.DecelleDr. F.JanczykDr. M.Uhlig Musee Royal de l'Afrique Centrale, Naturhistorisches Section Museum Wien,D1Entomologie, 2. B-1980 Zoologische Abteilung Insekten, Dr. C.A.Triplehorn The Ohio State University, Department of Entomology, 1735 Neil Dr. J.N.L.Stibick C urator SWED Dr. P.I.Persson ZMUB PAPU SAFR Dr. SAMU S.Endrody-Younga Dr. E.G.Matthews Transvaal Museum, Paul Kruger Street, P.O.Box 413, South Pretoria,Australian Museum, Entomology Section, North Terrace, OSUC TERV WIEN Acronym Table 1. Collections (Cont'd.)*
XXXIV INTRODUCTION
The lymexylid beetles (Coleoptera: Cucujiformia: Lymexyloidea sensu Crowson, 1955) comprise a relatively small cucujiform family whose members vary in. general appearance from soft-bodied, cleroid-like beetles to heavily sclerotized, melandryid-like beetles, to brachelytr- ous, bizarre forms which are, at a glance, not very beetle-like at all.
The lymexylids are one of a small number of insect lineages which have evolved true fungus-growing habits. Other fungus-growing insects in clude certain Old World termites, Attiine ants, and some other groups of beetles which include certain scolytids and curculionids (Wheeler,
1907; Wilson, 1971; see also "Natural History" below). This remarkable evolutionary achievement includes distantly related insects, which implies at least four independent developments of this ecological strategy, or perhaps more. These fungus-growing activities are be lieved to be more primitive in the Lymexylidae (and correspondingly less complex) than in other fungus-growing beetles.
Historically, the Lymexylidae have long been a focal point for active discussion and fervent controversy since pre-Linnean time.
The large, multi-branched organs arising from the palpi of most male lymexylids have become a benchmark for the family, and have intrigued naturalists for more than two hundred years. These structures seem to function in chemoreception, although this conclusion has followed
1. Fig. 1. Hylecoetus dermestoides, habitus, "flabellicornis" morph: head-on view (top), lateral view of forebody (bottom).
2. 3.
1 detailed morphological research (Slifer et al, 1975) and non-exacting observations (Germer, 1912; Germer § Stech, 1912); no rigorous experi mental ethological research has been undertaken. These maxillary palp- organs have contributed to a "taxonomic isolation" of the family, which is manifested in our century by recognition of a distinct and separate superfamily, the Lymexyloidea.
The aspect of the lymexylids which I find most intrigueing, however, is the apparently endless debate over the natural classification of the family. What are the phylogenetic relationships of Lymexylidae? What is the sister-group: Cucujoidea?, Cleroidea?, Cucujoids + Cleroids?, or some subunit of one of these?
Linnaeus described the species we know today as Hylecoetus dermes- to id e s as a species of C a n th a ris. The soft-bodied H ylecoetu s b e e tle s could indeed suggest an affinity to cantharoid families, and, in fact, this was again advocated Jeannel § Paulien in 1944. For the same reasons, a relationship to Cleroidea has been suggested (see discussion in Crowson 1955, 1960, 1964). Forbes (1922) compared hindwing venation among families of beetles, and inferred a relationship between lymexylid beetles and Heteromera. Sharp § Muir (1912) conducted a study of male genitalia, and recognized the basic sim ilarities of lymexylid genitalia and the trilobe design. Tanner (1927), based on a study of female genitalia of beetles, also found sim ilarities with heteromerous families
Boving § Craighead (1931) noted the strong cucujoid features of the larvae of lymexylid beetles. It was Leng (1920), who placed the Lymexylidae in a separate superfamily (Lymexyloidea, = Leng's re stricted Teredalia). This arrangement has been perpetuated to date, particularly by Crowson as part of an unresolved trichotomy which includes Cleroidea, Cucujoidea (Clavicornia + Heteromera), and Lymexyl oidea.
Most publications dealing with lymexylid beetles do so in a super ficial way. The bulk of the literature is composed of isolated descrip tions of individual species from various, separate geographic locali ties. A lack of understanding and appreciation for the ranges of morphological variation intraspecifically and for geographic d istri butions of populations have fostered a plethora of synonyms in the family. Few authors have compared more than one or two lymexylids in a single study, although Germer's (1912) excellent treatment of the maxillae and antennae of various lymexylids is an outstanding exception.
Modern systematic methods have, in general, not been applied to the lymexylids such as descriptions of genitalia, although there are exceptions (e.g., V illiers, 1969a). In the existing body of literature, some sp ecies a re m isplaced (assigned in c o rre c tly to a genus), and some genera proposed in recent years are clearly synonyms of pre-existing genera. Further, it has become apparent to me in my study that at least two new genera are needed to adequately deal with the known diversity in the family. 6.
All of these factors indicate the need for a modern systematic study of the lymexylid beetles. My goal in the present study is to review the current generic classification of the family Lymexylidae, to hypothesize cladistic relationships of the genera, and to propose a revised and improved classification. To reach this goal, I have examined as many taxa from as many global localities as possible to form an understanding of the diversity of lymexylid beetles among the extant fauna. I have also looked at both adult and larval stages of these beetles, to increase the numbers of characters available for elucidating sim ilarities and dissim ilarities among the genera.
I have not made an attempt to resolve species-level problems for the entire family. This was not my explicit goal, although it is a serious problem which is in need of urgent consideration, and it is not a necessary part of a successful review of the genera themselves. From the material which I have borrowed and studied, however, I have become convinced that the single greatest problem facing species-level studies of the lymexylids is that of unresolved synonymy. My personal estimate, based on the "species" which I have seen from around the world my concepts of the biological species based on morphological gaps), the literature, and identified material, is that only 33% to 50% of the nominal species are actually valid species. I have serious doubts that many species remain undescribed.
A number of types were borrowed from European institutions by mail, but loans of many important types could not be initiated .by this means. The most outstanding example being the Museum d'H istoire Naturelle in
Paris where no less than fifty types of Atractocerus and the Melittomma
complex are housed. Clearly the most efficient means of dealing with all of this material is to visit Paris and make necessary dissections and studies there. This, however, lies within the province of species-
level revisional work and cannot be entirely justified for a generic study. The type-specimens which I have studied are discussed individu ally in Appendix A, and has allowed species-level decision making for
Hylecoetus, Lymexylon, and Australymexylon. Thus, the need for species- level revisions are within Atractocerus and the Melittomma-complex.
The classification which I advocate is based on my cladistic analy sis of the genera. My analysis was done in accordance with the canons discussed by W illi Hennig (1965, 1966) and subsequent Hennigean cladists during the past decade. My classification is the first attempt to reflect the relationships among the genera on a global basis, and is designed as a framework for future research thrusts including detailed revisional studies of the species of each genus, continued refinements of generic concepts (particularly through additional field studies on immature stages and ecological associations of the species), and further probing into the superfamilial relationships of Cleroidea,
Cucujoidea and Lymexyloidea.
In this sense, my study is the beginning of studies at both "lower"
(species) and "higher" (superfamilial) levels. Cladistic relationships suggested here are hypotheses, and as such subject to critical testing and potential refutation. Although it is my belief that the relation ships discussed here are reflections of a genuine pattern of evolution ary relationships, we should continue to actively test these concepts through further character analysis. To paraphrase George Ball (1978),
I am less concerned that my ideas might be proven incorrect than I am that they might not be tested at all.
How can these hypotheses be tested? The first line of research should be directed at resolving species-level problems and simultaneous ly testing the distribution of character states: i.e., do the synapo- morphies which occur in species studied here actually occur in all species of that genus? The other areas of information gathering should include unexploited sources of characters: ethology, pupal structure, host interactions, morphological details not discussed in this study, etc., and a great increase in the out-group comparisons in character analysis. Detailed, comparative morphological studies are desperately needed in the Cucujiformia families of Coleoptera. A few galant efforts have been made during this century (e.g., Sharp § Muirs' 1912 study of male genitalia and Tanner's 1927 examination of female geni talia), but even these are shallow in taxon-representation and details analyzed (neither of the studies cited above, for example, use even compound m icroscopy), but most s tru c tu re s rem ain v ir tu a lly unstudied at higher levels in Coleoptera.
I believe that my study of the lymexylids will serve to stimulate additional research in all of these directions, and can contribute to 9.
each o£ them to a greater or lesser degree. MATERIALS AND METHODS
MATERIALS
Most of the specimens used in this study were borrowed from the institutions and individuals listed in Table 1 (see "Acknowledgments"), and a number of specimens generously given to me by several individuals.
I was able to see a living larva of Melittomma sericeum collected by my wife at Ohio State University's Bameby Center in southeastern Ohio, and to collect and observe Melittomma brasiliense and Atractocerus brasiliensis during my field studies on Barro Colorado Island in
Panama in 1978. In all, I have seen more than three thousand specimens of the family.
PROCEDURAL METHODS
Optical Equipment - Routine sorting and study of mounted speci mens was done with a Wild model M-3 dissecting microscope. Detailed study was conducted w ith a Wild model M-12 compound m icroscope using magnifications up to x450. Scanning electron micrographs were taken with a Hitachi model S-500 scanning electron microscope.
Illustrations - Habitus drawings and illustrations of some large structures were made with a drawing attachment and the dissecting
10. 1 1 . microscope. Most drawings of individual structures, however, were made from disarticulated beetles (see below) using a drawing attachment on th e compound m icroscope. John Nagy prepared most o f th e h ab itu s draw ings; a l l o th er drawings were made by me. David S tu tes operated the scanning electron microscope.
Measurements - Because lymexylid beetles vary a great deal in size, exact measurements of individual sclerites are of little value without making similar measures for many specimens, with some analysis of the range of variation. This type of statistical analysis was decided to be outside of the primary thrust of this study, and its need was supressed by the abundance of "qualitative" differences and sim ilarities among the taxa examined. Where scales are included in figures, they were drawn from an ocular micrometer. Measurements of specimens are crude estimates only, since individual beetles can flex the head and/or contract the abdomen to various degrees, and were made with an ocular micrometer on the dissecting microscope.
"Miscellaneous Observations", Appendix A - I have appended a group of miscellaneous observations on the lymexylids in Appendix A.
In this section, I include various notes on specimens, biological information, types examined, justifications for nomenclatural changes, and other discussion and/or information which did not neatly fit into the individual generic chapters. These observations are of direct value to continued research in the systematics of the lymexylids and are therefore included as an important source of information. 12.
Disarticulations- Most of my previous research has been involved with minute beetles including various leiodids (Wheeler, 1979a, 1979b,
1980) and archeocrypticids (Triplehom § Wheeler, 1979), and I have found it essential to disarticulate specimens so that individual parts could be adequately magnified (using compound microscopy) for comparison and illustration. A thorough discussion of the philosophy and method underlying d is a r tic u la tio n s was made re c e n tly by Watrous (1980); my methods are briefly summarized below.
Step 1: pinned specimens are relaxed in heated detergent solution or in heated Barber's fluid (formula in W h e e l e r , 1979a);
Step 2: specimens heated to boiling in 10% KOH (or NaOH) and left in KOH at room temperature until muscle tissue is mascerated (usu ally about 15 minutes to 1 hour or longer, depending on size and con dition of the specimen);
Step 3: specimens rinsed in distilled water;
Step 4: specimens disarticulated (i.e., head, prothorax, mouthparts, antennae, le g s, abdomen, g e n ita lia , e t c ., separated as needed);
Step 5; parts cleared in lactic acid (generally 1 day to1 week);
Step 6: parts rinsed in distilled water;
Step 7: parts stored in glycerin in small glass cups;
Step 8; parts studied in temporary depression-slide mounts using lactic acid and a cover slip, or a mixture of glycerin/glycerin jelly.
Genitalia are studied in the same manner. In the lymexylids it is easiest to remove the apical (visible) abdominal segment, clear this in 13.
10% KOH, dissect out the genitalia, and study as above.
For this study, entire disarticulated specimens were stored in small glass cups ("microbeakers" of several commercial firm s), of which
7 fit into small, plastic petre dishes. Stacks of petre dishes were stored in wooden racks designed and constructed by Larry Watrous and me after trying several other methods which proved unsatisfactory. PHILOSOPHICAL METHODS
Hammond C1976), Ball (1978) and other authors have emphasized the
importance of clear statements regarding the philosophical bases used
in systematic studies. I concur with this view of the significance of
philosophical statements, and have tried to express my position in past studies of leiodid beetles (Wheeler, 1979a; 1979b). The state ments below should help the reader to understand the rationale behind
individual decisions in this work.
The sources for my ideas are diverse, and include theoretical
systematic discussions with numerous scientists who shared their views on the topic including Lee Herman and Norman Platnick of the American
Museum of Natural History, New York, George Ball of the University of
Alberta, Terry Erwin and Donald Whitehead of the Smithsonian Institution
David Kavanaugh o f th e C a lifo rn ia Academy o f S ciences, and e s p e c ia lly
Larry Watrous of the Ohio State University. A measure of congruence will be noted between my ideas and those expressed by these individuals and others not mentioned here.
I believe that the cladistic approach to formulation of hypotheses about relationships among organisms is the most clearly formulated approach of those presently available. My method is essentially that of Hennig (1965, 1966), but with some recent refinements (e.g., Plat nick, 1979; Watrous 8 Wheeler, in press). In the decade and a half
14. 15, since Hennig's ideas were published in English an incredible body of literature has emerged discussing cladistic theory and applying it to diverse groups of animals and plants. One illustration of the degree of proliferation can be cited from personal experience. Larry Watrous and I began compiling a set of references on cladistics (including pertinent pre-Hennigean titles, such as Maslin, 1952), with applied publications being confined to Coleopteran titles, but indefinately delayed completion after reaching about 450 titles.
What has been the result of all of this activity? One result has been the emergence of several refined approaches to the cladistic . analysis (e.g., Estabrook, 1972, 1974; recent articles by Farris - see
Farris, 1979-; and articles by Platnick and Nelson of the American
Museum in recent issues of Systematic Zoology): some involving computer programs, others concentrating on basic conceptualization of cladistic principles. One net effect of all of this is the persistant need to state your philosophy, even within cladistics.
Larry Watrous and I have co-authored a paper on cladistic theory which is now in press in Systematic Zoology. Many of the fine points of my philosophy are included in our paper, and need not be expressed again he_re.
Appendix B includes two illustrations which were developed by
Watrous and me for a discussion group on cladistic methods at the North
Central Branch meetings of the Entomological Society of America in 16.
Indianapolis in 1979. These illustrations, and the following comments, will nest our paper in the broader topic of cladistic analysis , general ly , and systematics.
What is the place occupied by cladistics in modern systematic research? A species concept is developed from various sources of information - study of types, specimens collected, borrowed or purchased
Based on these ideas, "species" are used to construct matrices which summarize the distribution of character states among the species. These characters are then analyzed cladistically, and an hypothesis about relationships made. From this basis,further hypotheses can be set forth in the areas of biogeography, ecological relationships and evolutionary pathways.
How is a cladistic analysis done? I believe that in its simplest form, the following steps are included: (1) search for characters
(this includes using refined techniques- e.g., scanning electron microscopy-, immature stages, ecological data and other sources);
(2) polarity of character states are hypothesized (see Appendix B:
Watrous § Wheeler, in press); (3) monophyletic groups are delimited based on synapomorphies (this includes considerations of parsimony and character weights, as discussed by Hecht § Edwards, 1976); (4) conflicting characters (indicating a mistake in analysis and a parallelism, convergence or reversal) are critically re-exmained.
Once all data can be explained in the framework of a single cladogram, 17. we have reached our hypothesis of relationships and can, if we choose, proceed to make causal explanations or hypothesize biogeographic histories. Although I do not propose biogeographic hypotheses for the lymexylids, it should be pointed out that the vicariance model for historical analysis is particularly harmonious with cladistics and is, at least to some degree, testable (see Platnick § Nelson, 1979).
Once homologous character states have been recognized (an unavoid ably subjective procedure), and polarities determined (Watrous § Wheel er, in press) the analysis is straitforward until conflicting character states arise. Watrous $ Wheeler conclude that out-group comparisons are the most useful among various criteria in the literature for polarization of states (e.g., Crisci § Stuessy, in press; de Jong, 1980). HISTORICAL REVIEW
"So, what follow s i s not the h isto ry o f carabid classification, but a_ history as seen by one who appreciates and respects the workers of the past, who believes that science advances by evolution rather than by revolu tion , and who, as an evolu tionist, believes that phylogeny and classification must be linked more or le s s closely"
- George Ball, 1979: 63
The lymexylid beetles have attracted the interest of a large
number of coleopterists,and naturalists generally,for more than two
hundred years. The bizarre structure of these beetles and their
relative rarity have spurred on those lucky enough to encounter them
in the field. The result has been, overall, somewhat less than bene
ficial for the taxonomy of the lymexylids. Most species were described
individually, without comparative references to previously named forms
with resultant synonyms. In fact, it is very likely that all (or most) of the extant species of lymexylids are already known to science , and
one of the greatest chores yet to be done for the family is to iron
out these synonymies and provide a stable system of species-level names.
My generic classification for the lymexylids seeks to express what I have been able to learn about the cladistic, or "phylogenetic",
re la tio n s h ip s and a t th e same tim e to make taxonomic changes as
conservative as I can. I hope that the result is a classificatory
18. la. system that is more natural than those which, came before it and one with minimum disruptive effect on the existing body of literature,
Omnispective hindsight is an advantage to one evaluating history,
but we should not fail to credit earlier workers for sound decisions
based on less information than we have. today, and perhaps weighed on
different scales influenced by prevailing social and scientific
thought. Historians and scientists gain respect for past ideas and methods because our own have evolved from these beginnings, usually at the expense of misconcepts left behind. In fact, the scientific method itself proceeds much the same according to philosopher Karl
Popper 0-968) and his followers, Preliminary hypotheses are made, and critical tests are designed in an effort to refute these. If success** fully falsified, an improved or new hypothesis is made.
The lymexylids are a fascinating assemblage of beetles, and their taxonomic history is rich in proportion, A great deal of attention has been showered upon a small group of animals, and the unusual features of these beetles make them easily distinguishable to all students of Coleoptera, I present a brief historical review of the study of lymexylid beetles below. I attempt to expose the highlights and trends, without going too deep into taxonomic details which can be extracted from later chapters, 20.
Description of Species
Uddman (1753) described the first species of Lymexylidae five years prior to the publication of Systema naturae (10th edition) by
Linnaeus in 1758. His species, known today as Hylecoetus flabelli- corn is, was validated by Schneider in 1791. Linnaeus described the second species in 1758, navale, l a t e r moved to a new genus, Lymexylon, by Fabricius (1787); and, in 1761 described the species we now know as Hylecoetus dermestoides.
Subsequent descriptions came slowly at first, but at a relatively steady and increasing rate. Following 1820, there would be a virtual plethora of isolated species descriptions. The latter trend has con tinued into the twentieth century (Fig.291 ), due in large part to
Maurice Pic's publications, and those of a few other authors.
Based on my examination of more than 3,000 specimens from museums around the world, I believe that I have been able to see most of the b io lo g ica l species (considering external morphological features and male genitalic structure). It is my belief that the future will witness a drastic reversal of this trend, initiated by a large number of synonymies reducing the number of valid species by 50% or more.
Descriptions of valid new species will be rare, if there are species yet to be described.
One point to be made in support of this prediction is that most of the species of Lymexylidae are very widespread geographically. 2 1 .
Single species range over broad areas of Central and South America, continental Africa, Southeast Asia and Indonesia, and Australia. The typical pattern includes relatively few species, but with wide-ranging distributions. The propagation of synonymies has also been fired by inordinate degrees of sexual dimorphism. Melittomma brasiliense, for example, has been named several times due to differences in color between males and females, and differences in the structure of the maxillae and the antennae (males have large maxillary palp-organs, and females have bilobed, almost flabellate antennae) as pointed out by Lane (1955).
Description of Genera
Genera have been described at a slow rate (Fig. 291). Lymexylon was proposed by Fabricius in 1775, and Hylecoetus by Latreille in 1806.
Palisot de Beauvois named Atractocerus (see Item 18 in Appendix A) in
1801, and Murray named Melittomma in 1867. Aside from the later de scriptions of genera discussed below, there has been widespread con fusion in the constituency and application of these generic units.
Species actually belonging to Melittomma-complex genera and Australymex- ylon, new genus, were erroneously assigned to Hylecoetus, with similar mix-ups among other genera.
Pic (1939) described a new genus (Heteromeroxylon) included in
Zoological Record under Lymexylidae. An examination of Pic's paper, 22.
however, indicates that it was described in Rhipiphoridae and that
Pic noted a sim ilarity in appearance to Lymexylidae. Lane (1955)
described two new genera from South America. Melittommopsis is only
known from female specimens (neither Lane nor I have been able to see
a single male), and thus it is problematic to effectively evaluate its
actual significance. Hylecoetopsis was erected for the single species
Hylecoetus cylindricus (see Germer, 1912), known only from a single
specimen from Cayenne. Lane did not actually see the type-specimen,
and based his genus on a single drawing of the antenna which had been
published by Germer (1912). I have not been able to learn in what museum the specimen resides. From Germer's illustration, however, it
is questionable that the beetle in question is actually a lymexylid,
and Lane's proposed genus cannot be seriously evaluated at this time.
I might note that some South American Mycteridae do look a little like
certain lymexylids, and that a species of this or some other family may be involved in the cylindricus problem. Kurosawa (1956) named a
genus, Neohylecoetus, based on material from Java and the Philippines.
From his excellent drawings there is no doubt that his species are good Melittomma. I am certain that the genus must be synonymized, and it is very likely that his two new species are invalid as well (i.e., his drawings and reported geographic distributions concur with the structure and distribution of Melittomma javanicum, a common and widespread species). Description of Larvae
Considering the relative obscurity of most lymexylids, an amazing amount has been published regarding the immature stages, particularly larvae. Two species of Hylecoetusf one each of Lymexylonf Melittommaf and Melanomelittomma NEW GENUS f and seven species of Atractocerus have been described. Some authors have done very thorough reports, includ ing neonatal (newly-emerged first instars] larvae, late-instar larvae, eggs, pupae, adults, and damage to wood (e.g., Lyngnes, 1956 for
Hylecoetus dexmestoidesf Simmonds, 1956 fo r Atractocerus brasiliensis; and Fulmek, 1931 fo r A. emarginatus] , The amount of biological infor mation varies, but is sometimes sizable (e.g., Vessey-Fitzgerald, 1941].
The described larvae of Lymexylidae are summarized in Table 2, and the appropriate references cited.
Crowson (1955, 1960, 1970] has consistently advocated the increased use of larval characters in systematic investigations of beetles, but the lack of use persists as a problem of major proportion. Boving §
Craighead (1931] remains the standard reference for beetle larvae at the familial level, although this type of study needs to be increased in scope relative to the number of taxa compared and to the number and detail of structures studied. Peterson (1961] is another good source of information on North American beetle larvae. Table 2. Descriptions of larvae of Lymexylidae.
Taxon Reference Comments
F am ilies Boving & Craighead, 1931 key (illus. Melittomma and Hylecoetus)
F am ilies Peterson, 1961 key (illus. Melittomma and Hylecoetus)
F am ilies Britton, 1974 families occuring in Australia
Subfamilies Boving & Craighead, 1931 Hylecoetinae, Lymexylinae
Subfamilies Crowson, 1955 Hylecoetinae, Lymexylinae
H ylecoetus lu g u j b r is Boving & Craighead, 1931 i l l u s .
H. lugubris Boving S Craighead, 1931 i l l u s .
H. dermestoides Grandi, 1961 description & illu s. neonatal la r v a ; detailed morphology
H. derm estoides Germer, 1912 neonatal larva + mature larva, i l l u s .
H. dermestoides Lyngnes, 1956 photos: egg, neonatal larva, mature larva, pupa, adults, damage
H. dermestoides Egger, 1974 photos
| S j Lymexylon navale Dominik, 1957 habitus drawing Table 2. Larvae of Lymexylidae (Cont'd.).
Taxon ______R eference______Comments
Atractocerus spp. Britton, 1974 key to families
Atractocerus spp. Gardner, 1935 key to some species
Atractocerus brasiliensis Simmonds,1956 detailed description including neonatal larva, mature larva, egg, pupa, and damage to wood; well illus.
Atractocerus brevicornis Gardner, 1934 brief diagnosis, illus.
Atractocerus crassicornis Clark, 1931 general desc., and illu s.
Atractocerus emarginatus Fulmek, 1931 neonatal larva, mature larva, pupa; good illu s.
Atractocerus kreuslerae Clark, 1926 biological observations and damage to wood
A. kreuslerae Britton, 1970 illu s. mature larva
Atractocerus reversus Gardner, 1929 illu s. (mature larva)
A . rev e rsu s Gardner, 1944 neonatal larva + eggs
A. reversus Roonwal, 1972 photo of mature larva + damage to wood tsj (SI Table 2. Larvae of Lymexylidae (Cont’d i).
Taxon ______R eference______Comments______
Atractocerus quercus Gardner, 1935 description, illus.
Melittomma sericeum Boving & Craighe.ad, 1931 key +illus.
M. sericeum Peterson, 1961 key + illus.
Melanomelittomma insulare, N.Comb. Gahan, 1908 desc., illus.
M. insulare Vessey-Fitzgerald, 1941 desc., biology, illus.
M. insulare Simmonds, 1956 notes
M. insulare Lever, 1969 habitus drawing
**NOTE- "mature larva" is used under "Comments" to refer to late instar larvae. In most cases, it is not clear how many larval instars there are and the only possible differentiation of the exist -k> ing literature is into "neonatal" (newly emerged 1st instar larvae) and "mature" (later instars). 5* 27.
Studies referring to larvae appear sporadically in the literature, some recent examples including Crowson (1964), Dybas (1976), Hayashi
(various years: see References), Lawrence (1977), Hlavac § Lawrence
(1979), Kasule (1969), Watt (1974) and Wheeler (1979b). Superfamilial Status
The concept of Lymexyloidea as a distinct and separate superfamily
of Coleoptera seems to have originated with Leng (1920: p.31). The
acceptance of this view by Boving § Craighead (1931) and Crowson (1955)
has served to strengthen its acceptance, and it is yet to be shown
where the natural affinities of Lymexylidae lie.
Early authors stressed the similarity of the lymexylids to soft
bodied beetles, including the Cantharoidea and the Cleroidea. Those
lymexylids which are soft-bodied (e.g., Hylecoetus, Lymexylon,Austra-
lymexylon, Melittomma spp.) are convergent in both the lightness of
sclerotization and the dense pubescence covering the body. Crowson
(1964) has pointed out that these characters are indicative of beetles which are either short-lived or have secretory functions related to the integument. In a recent survey of entomology in Canada (Danks, 1979), Bright
(1979) placed Lymexylidae in Cleroidea. Although a sim ilarity to cleroidea has been suggested on several occassions, this is the only recent attempt (that I am aware of) to actually place the family within Cleroidea. Following my cladistic analysis of the genera of
Lymexylidae, I offer some speculations on superfamilial relationships, and disagree with affiliation of lymexylids and cleroid families (see
Crowson, 1964; Barron, 1967).
2 8. NATURAL HISTORY
"Among the multitudinous activities of insects, none are more marvellous than the fungus-growing and fungus-eating habits of the Attiine ants" -William Morton Wheeler, 1907
"...sym biotic fungi were subsequently discovered to be associated with certain bark-inhabiting scolytids, for example the genus Tps, as well as other kinds of beetles , including occassional species of Anobiidae, Brenthidae, Curculionidae (in Scolytoproctus and Gulamentus) , and all of the Lymexylidae so far examined" -Edward 0. W ilson, 1971
Many species of beetles, flies, and other insects are fungus- feeding, and, in fact, it is probable that mycophagy has been of immense importance in the evolution of many of the present day families of beetles (Crowson, 1955, 1960). A symbiotic relationship , however, between insect and fungus are far less commonplace^ where the insect actually plays an active role in the dissemination and growth of the host. Among the first Coleoptera to reach the evolutionary plateau of fungus-growing habits were the Lymexylidae.
Schedl (1958) recognized a number of categories for tree-infesting insects, ranging from phyllophagy (leaf-feeding) to xylo-mycetophagy
(forms which do not feed on wood itself, but rather on fungi which are cultivated on the tunnel walls), and based on larval feeding habits.
Intermediate behaviors included feeding on living tissue under bark 29. 30.
(phloeophagy) , on outer, dead layers of hark Ccortiphagyl , and living in the wood itself and gaining nutrients with the aid of entosymbionts which are either intracellular or free-floating in the insect gut
Ixylophagy) . Schedl considered xylo-mycetophagy to be the most re fined specialization in his set of feeding strategies, and summarized its occurance as follows: Scolytidae CIpinae: Corthylini, Xyleborini;
Scolytoplatypodinae); Platypodidae; Cce*“tain} Lymexylidae; Curculioni- dae (Scolytoproctus and Gulamentus); and Brenthidae (Calodromini,
Taphroderini). Schedl suggested that the Lymexylidae are perhaps the most primitive of all xylo-mycetophagous b e e tle s .
Schedl's term xylo-mycetophagy was coined for use in his system of larval feeding patterns, but a more general term was needed for those beetles which was less cumbersome. Ambrosia beetles has been used by
Batra Cl964) and others to refer to these forms excluding Lymexylidae) but as Wilson Q971) pointed out, this is appropriately applied only to certain scolytids. Wilson C1971) suggested, as an alternative, sim ply "Fungus-growing" which is at once easy to apply and self- explanatory, and which includes all of the fungal cultivators.
Wheeler C1907) included Attiine ants, certain Old World termites, and certain beetle groups in his considerations of fungus-growing insects in his monographic study of the A ttii ants Csee Wheeler, 1907:
775-785). Wilson Q.971) also summarizes known incidences of fungus- growing in Chapter 7 of The I n s e c t S o c ie tie s . LYMEXYLID BEETLE BIOLOGY
How much do we presently know about the biology of the lymexylid beetles? The available information varies immensely, from the rather well-known Hylecoetus dermestoides to forms known only by taxonomic descriptions of single specimens.
I have pulled together as much data on the hosts of Lymexylidae, including arboreal hosts and microbial hosts, as possible. This is presented in Table 2. Because of the diverse nature of the data on various taxa, I discuss them individually below.
On The B iology o f th e Genus Hyleocetus. Of all the lymexylid beetles, the greatest amount of biological information has been amass ed fo r Hylecoetus. Most of the discussion below applies sensu stricto to the European species H. d erm esto id es. The North American species,
H. lugubris, however is likely to prove very similar in most details.
As of now, both species share the same fungus host species: A scoidea h y le c o e ti.
Lyngnes (1958) made extensive observations on H. d erm esto id es which were infesting logs of birch-wood in Norway, and reached the following conclusions:(1) egg-laying appears to occur at temperatures above 8°C; (2) old wood (1 h years), slightly fermented, was preferred to freshly cut trees; (3) young larvae bore head-first through the bark and into the wood and use their tail-end to push debris out of
31. Table %. Host information for lymexylid beetles.
Lymexylid beetle Host or biological data Arboreal host ______Reference/Country
Hylecoetus Ascoidea hylecoeti Pinus spp. Batra § Francke-Grosmann, dermestoides 1961; B atra, 1967/ Sweden
Hylecoetus Asoidea hylecoeti Pinus, Liriodendron Batra § Francke-Grosmann, lu g u b ris (Poplar) 1961; Batra, 1967; Simmonds, 1956/ USA
Unidentified Chlorophora excelsa Batra § Francke-Grosmann, 1964; B atra, 1967/ West A frica
Melittomma Oak, Chestnut, Elm Blatchley, 1910; Sanderson sericeum § Glenn, 1963; Hagen, 1886/ USA
Melanomelittomma Gram negative rods in fermenting Coconut Palms § Wild Brown, 1954; Lever, 1969/ in su la re wood, many contam inants (y ie s ts , Palms: Sterensonia & Seychelles and Madagascar etc.) Host: ? Nephrosperma; RARELY: Deckenia & Roscheria
Atractocerus Inferred predation on Lampyrid Gorham, 1886/ C entral § spp. beetles (historic interest only; South America unsubstantiated)
Atractocerus Mould § sap Boswellia serrata Beeson, 1961; Roonwal,1972 reversu s ("Sallai"); Lannea Gardner, 1936, 1944/ India coromandelica Ceylon (''Jhingan11); Lannea grandis w N) Table 3. Host information for lymexylid beetles (Cont'd,},
Lymexylid beetle Host or biological data Arboreal host Reference/Country
Atractocerus Burrows sometimes infested with Eucalyptus patens Clark, 1926/Australia k reu slera e fungus which Clark suggests ("Blackbutt"); E. kills the larvae ?? gomphocephala ("Tuart"); E. redunca v. elata ("Wandoo" ); E. ru d is ("Flooded Gum"); E. calophylla ("Marri")
Atractocerus Bombax malabaricum Blair, 1936/ India a l b ita r s is
Atractocerus Q u e r c u s d i l i t a t a Gardner, 1935/ India q u e r c u s
Atractocerus "Nkoba” log (Uganda); Simmonds, 1956/ A frica brevicornis Mahogany lo g (Gold C o a st); Limba lo g (B elgian Congo); Ekebergia ruppeliana (heart- wood, Tanganyika); Midbrae- diodendron excelsum;(Teak): Tectona grandis; Ricinodendron africanum; Triplochiton sclero- xyIon; (Cashew Nut): Anacardium occidentale (Kenya); Swietenia mahogani .
Atractocerus Dipterocarpus indicus Simmonds, 1956/ In d ia , n ig er Madras Table 3. Host information for Lymexylidae (Cont'd.).
Lymexylid beetle Host or biological data Arboreal host Reference/Country
Atractocerus No large necrotic area, wood Mangifera indica Swabey, 1935; Simmonds, brasiliensis dryer than in M. insulare ("Mango"I/ Mora 1956/ Trinidad, Costa host woods excelsa; Prioria Rica, Barbados copaifera; Albizzia lebbeck
Atractocerus Terminalia myriocarpa Simmonds, 1956/ Assam b la ir i
Atractocerus Buchanania la tifo lia / Gardner, 1936/ India, emarginatus Hevea brasiliensis Malaya, Ceylon ("Rubber tree"]; Dipterocarpus zeylanica
Atractocerus Near nest of Cornitermes sim ilis Wasmann, 1902/ B razil termiticola
Atractocerus Associated with termites C??) ? Wasmann, 1902/ W, A frica brevicornis
Melittomma Boletus ???? ? C astelnau, 1832; Simmonds, brasiliense 1956/ Central § S. America
Melittomma sp. Ficus exasperata; Simmonds, 1956/ W, A frica Funtumia africanum; Triplochiton sclero- xylon; Terminalia w superba f* Table 3. Host information for lymexylid beetles (Cont'd,).,
Lymexylid beetle Host or biological data Arboreal host Reference/Country
Lymexylon navale ? Oak; Chestnut Donisthorpe, 1939; Simmonds, 1956/England
04 cn 36.
the tunnel (Lyngnes also suggests that the posterior end provides
defense against intruders and carries particles - perhaps even spores -
into the bore-hole); (4) later stages face the entrance hole and can use the saw-like ninth abdominal segment for boring in the wood (this
bizarre behavior was also reported by Egger, 1974); (5) besides the host fungus, other fungi grow in the tunnels which may be suitable as
food including Isaria, Aspergillus, and Verticillium; (6) fungal spores are transferred from the emergence site to that of mating and ovipostion by female fungus-pouches (see Buchner, 1928 and Francke-Grosmann, 1967)
and male maxillary palp-organs; and (7) Lyngnes questions that the male palpi are used to find the female, and suggests that the males
simply home in on suitable wood substrata (Slifer et al, 1975, however, suggest the opposite; the simple fact that these organs are sexually dimorphic in H ylecoetu s would seem to add weight to S lifer1s position).
Francke-Grosmann (1967) provides an e x c e lle n t summary o f H ylecoetus biology, the emphasized points which differ from Lyngnes' discussion being the following. The fungus-pouches of the female exude spores embedded in a slimy matrix, at the time of oviposition. Further, neonatal larvae twist about their companions and eggs, seemingly to enhance contamination by spores.
There is yet another interesting aspect to the Hylecoetus-Ascoidea story. Cooreman (1952) described the deutonymph of a tyroglyphid mite,
Histiogaster hylecoeti , which are carried under the elytra of the beetles, and which apparently leave the beetle during oviposition and also inhabits the larval tunnels. 37.
Francke-Grosmann (1967) points out that like many ambrosia fungi of the scolytids, that Ascoidea hylecoeti is found only in association with the host beetle, but can grow in many different arboreal hosts, including in Europe many hardwoods and all softwood species. The fungus provides an excellent source of food for the developing larvae, but excessive growth seems to retard larval growth (Francke-Grosmann,
1953; 1967), and, Batra § Francke-Grosmann (1961) suggest that the enzymatic activity of the fungi might make the cell walls of wood more managable for the larvae.
In the classification of Alexopoulos and Mims (1979: pp. 262-263),
A scoidea belongs to the family Ascoideaceae of the subclass Hemi- ascomycetidae (Class: Ascomycetes). A scoidea is associated with plant exudates and with ambrosia beetles (including Lymexylidae), and
Alexopoulos and Mims characterize the family by multispored asci and hat-shaped ascospores, among other characteristics.
On the Biology of Melanomelittomma insulare. Perhaps the most intensive series of biological studies of any lymexylid beetle were undertaken by the British government to investigate the potential control measures against Melanomelittomma in s u la r e , NEW COMBINATION, a very serious pest of coconut palms in the Seychelles. Among the reports on these scientific activities, those by Simmonds (1956),
Vessey-Fitzgerald ( 1941), and Brown (1954) are particularly informa tiv e . ECONOMIC IMPORTANCE
Although most lymexylids are rare in museum collections and are actively sought by amature collectors and professional coleopterists alike, some species have causedserious economic loss to trees and wood products in the past, a few are pests today, and other species have the potential to become pests in the future as our perceptions of the value of various tropical trees change, and our uses of these trees and their cultivation increase in years ahead. This is particularly true for countries in tropical climates around the world.
Particular details of the biology of many species are discussed and summarized in my chapter on "Natural History", and this information need not be reiterrated here. What I would like to do, however, is briefly point out those lymexylid beetles which have been economically significant up to date in various parts of the globe.
Lymexylon navale - This species, the true ship-timber beetle, was a serious pest of the shipping industry in the eighteenth century in
Europe and Scandanavia. Changes in ship-building materials and other unknown factors, however, have rendered the species relatively harmless today. Although the species occurs widely, it is rather rarely collected at present.
Hylecoetus dermestoides - This species is sometimes a serious, localized pest of cut wood in Europe. Losses are generally not great.
38. 39.
In addition to comments on its biology by Egger (1974) and Lyngnes
(1956), Dominik (1975, 1976) discusses its economic importance and some c o n tro l m easures.
Melittomma sericeum - This species was, during the nineteenth century, a serious pest of chestnut trees in North America (Craighead,
1950, said that 50% to 90% of the chestnuts had worm-holes inflicted by M. sericeum; Champlain § Knull, 1923, reported damage to structural wood by this species as well). After the blight of American Chestnut, however, the species was never again a pest of any economic consequence.
Today th e sp ecies i s uncommon.
Me 1 anome 1-11tomma in s u la r e , new com bination - This sp ecies has caused more damage, measured in dollars, than any other single species of Lymexylidae. It appears to be a species native to Madagascar, which was introduced into the Seychelles, afterwhich it became a very serious pest of the coconut palm (see Brown, 1954). This species is also a pest of coconut palm in Madagascar.
As an example of the seriousness of this problem, between 1953 and 1958, 77% of the coconut trees (about 95,500 trees) on Praslin
Island were infested and after careful removal of infested wood and plugging the damaged area of the bole with tar infestation in 1961 had been reduced to about 18% (Nye, 1961). Other examples of the damages of insulare were cited by Brown (1954). Production on one plantation fell from 22,000 nuts per month in 1948 to about 15,000 per month in
1952 due to the beetle infestations. In one estate of about 350 acres, 40. the annual loss of mature trees was at the rate of about 150 trees per year. As implied by these figures, the problem has been drastically reduced, and is presently under control. Further details can be found in papers by Vessey-Fitzgerald (1941), Brown (1954), Simmonds (1956), and Nye (1961).
Atractocerus kreuslerae - Clark (1926) reported on the damage by the "pin-hole borer" in Western Australian forests, observing that
"Probably no insect does greater damage to the commercial timber growing in Western Australian forests". This species seems to attack a large number of Eucalyptus species. Clark determined, from emergence monitoring traps strapped to infested trees, that the larvae require at least two years to complete development.
Atractocerus reversus - Roonwal (1972) reported that this species is "a serious pest of standing trees, felled logs, billets and plancks of the salai, Boswellia serrata Roxb., and the jhingan, Lannea Coro mandel ica (Hout) Merr. (synonym 1^. grandis (Dennst.) Engl.), in India".
Further details reported by Roonwal center on description of larval boring activities, and biological notes. EXTERNAL MORPHOLOGY OF HYLECOETUS
The lymexylid beetles represent fragments of an ancient lineage and because our fauna today contains only a few species from several major lineages the kinds and widths of the morphological gaps between the genera are truely impressive. In fact, some characters differ between genera in distances normally recognized between familial units.
This, and my use of some unconventional sources of characters indicate the need for clear definition of the terms used, I can think of no more concise definition than a clearly labelled illustration and an accompanying discussion.
For this introduction to lymexylid structure, I have chosen
Hylecoetus dermestoides as the representative species. This beetle is common throughout much of Europe and parts of Asia and is occasion ally of economic importance as well as causing damage to cut, stored supplies of wood. More is known of the biology of this species than any other single Lymexylid species. This kind of information helps to round out our perspective on structure since it is possible in some instances to reasonably infer some functional explanations underlying particular structural configurations. And, on the practical side, there simply were many specimens available so that dissections could be freely made to compare and evaluate structures among individuals. 42,
This morphological account of H ylecoetu s will also supplant a formal description of the genus, as is made for all other genera of the family. Appropriate references, however, are made in the section on
H ylecoetus taxonomy to various parts of the present morphological discussion.
Hylecoetus dermestoides
GENERAL DESCRIPTION. H ylecoetu s d erm esto id es has an elongate more or less parallel body form and is moderately flattened dorso- ventrally. Length varies from about 4.8 mm to more than 16.0 mm. The body is lightly sclerotized throughout, thus resembling superficially beetles of the Cleroidea or possibly the Cantharoidea. Most of the surface of the body is clothed in fine setae. The legs are long and thin in form, and the elytra are long, covering the fully-developed hindwings and much of the dorsal surface of the abdomen.
THE HEAD. The head o f H ylecoetu s is prognathous, broad, and somewhat flattened dorso-ventrally in shape. Dorsally, the head is approximately as wide as long, and is rounded in shape (Fig. 2); there is no distinct constricted area behind the eye delimiting a "neck" region, as is common in other genera (e.g., Melittomma, Lymexylon,
Australymexylon, Atractocerus ) . The vertex has an evident "epicranial pit" (Figs. 2, 30) of unknown function. Crowson (1955) suggested that this pit might be involved in the transfer of spores, but this explana tion seems superfluous in light of what we now know about the fungus- 43. pouches of the female genitalia (see Francke-Grosmann, 1964).
The surface of the head is covered by very dense, rough, and nearly reticulate punctation, and with very fine, dense, suberect setae. The antennae are inserted just anterior and mesad to the eyes.
There is no fronto-clypeal suture (the clypeus is not clearly separated from the frons).. The gula is short and narrow (Fig. 3); the gular sutures nearly parallel, very slightly divergent both anteriorly and posteriorly. There is a distinct gular ridge bordering the opening of the foramen (Figs. 4, 5).
THE ANTENNAE (FIGS.1,2,10,18,23,24,29). Because I am synonymizing
Hylecoetus flabellicoznis w ith Hylecoetus dermestoides I have to deal with both forms of the male antennae in this discussion. I will first describe the form of the antennae in "typical" dermestoides and female dermestoides and then describe the "flabellicornis" antennal form.
The antennae are slender, 11-segmented, and not much longer than the length of the head at the midline. The scape (segment 1) is elongate and slightly clavate in form (Fig. 10); the pedicel (segment
II) is short, not much longer than wide; segment III is elongate, clavate in form; segments IV-X are short, transverse, produced asymme trically (constricted near the middle). The scape has a patch of sparse, short, spinose setae near the base; a few similar setae occur near the base of the pedicel; the scape and pedicel each have long, rather stout setae distributed over their surfaces. The apical portion of segment III, and the entire surface of segments IV-XI, is covered with both the stout, long setae present on the first two 44. segments and short, broad, (apparently) thin-walled setae. The scanning electron micrograph in figure 29 shows a portion of the surface of a female dermestoides antenna; the large, stout setae are fluted while the thin-walled setae have collapsed for the mostpart in this preparation. A third type of seta is also seen which is a small cuticular opening surrounded by minute, stubby projections. Near the top of the photo are some microtrichiae (minute, pointed projections of the cuticle); these are more easily seen in a photo of the male antenna in figure 18. Both thin-walled setae and the larger, fluted setae are evident on the laterally produced apices of the antennal segments IV-X and th e apex o f I I I .
I consider Hylecoetus flabellicornis to be a synonym o f H. dermestoides. H. flabellicornis males were long recognized because of the bizarre form of the antennae, which are strongly flabellate as the specific epithet implies (seen in Figs. 22, 23, 24, and the habitus in
Fig. 1). In these beetles which have the flabellate antennae the maxillary palp-organ is greatly degenerated and is minute in size.
The surface of the flabellate antennae have the typical long, fluted setae, broad more or less imbricate "scales" on the integument, and rather elongate, thin-walled setae (see Figs. 23, 24); the fluted
(and perhaps tactile; see Slifer et al, 1975) setae are lacking on the inner surfaces of the branches of the antenna (the protected areas). 45.
THE LABRUM AND EPIPHARYNGEAL REGION (T ig s. 1 1 ,1 2 ,1 6 ,1 7 1 . In
H y l e c o e t u s the labrum is a broad, flat, small-sized lobe which projects anteriorly from the clypeal region of the head, and is separated from the clypeus by a broad clypeo-labral membrane. It is lightly sclerotiz- ed, particularly along the lateral margins. Drosally, the labrum is covered by long, stout setae. Near the apex of the labrum these setae become very dense toward the raidline and coverge with similar setae which project from the ventral surface to form a dense, medial, setal
"brush" (m°st of the setae forming the brush are often broken off, making these setae appear shorter and far more stout than might other wise be apparent]. A ventral, longitudinal, medial patch of setae
(visually similar to the setae on the dorsal surface with light microscopy] which become more dense apically and also contribute to the brush (Figs. 11» 12, 17].
Tormae. The tormae are a pair of posteriorly-directed struts arising from the labrum which border to a greater or lesser degree the epipharyngeal membrane. In H yleco etu s the tormae are short and expanded ventrally and medially as broad, sclerotized plates. There are a few fine setae along the raesal margins of the tormae (Fig. 12].
Epipharyngeal Membrane. The epipharyngeal membrane i s th in and lies posterior to the labral sclerite and between and ventral to the tormae. It has a few fine setae scattered laterally and bears a dense, mesal patch of microtrichiae which converge along the midline to form a trichial-ridge (Fig. 12; also, in figure 16 the microtrichiae are visible in center of photo, and several labral setae to the left). 46.
THE MANDIBLES (F ig. 271, The mandibles in H yleco etu s are simple, short, and robust in form. The prostheca is nearly absent, only a few setae remaining. The left and right mandibles are nearly identical in structure. The outer surface bears several very long, stout setae. The apical, incisor, lobe consists of an apical dens and a poorly defined, second dens basal to the apical one. There is no distinct mola, although there is a small area of fine setae basal to the prosthecal remnant which could he some vestige of the mola.
THE MAXILLAE AND MAXILLARY PALP’ORGAN (F igs. 2 , 6 -2 , 1 3 , 14, 12,
25, 26f 281 • The degree of sexual dimorphism in maxillary structure in
H ylecoetus (and most other genera of Lyraexylidae) is truely extra ordinary and, in the case of Hylecoetus dermestoides, between male
"morphs" or forms as well. The basic structure of the maxilla is discussed, and then the various modifications.
Cardo. The cardo is relatively small, and in shape it is narrowed basally. Doyen (1966) and Evans (1961) discuss the muscle attachment patterns for most Coleoptera. See figures 6, 7.
S t i p e s . The stipes is not divided into a distinct basistipes and dististipes and further, it is not clearly separated from the lacinia or galea by distinct sutures. The stipes articulates basally with the cardo. See figures 6, 7.
L acin ia. The lacinia is the inner lobe of the maxilla and in
H ylecoetus consists of a small, tapered, membranous lobe which is covered by dense, long, fine setae. 47.
Galea. The galea is the "inner lobe" of the maxilla, located beside the palpifer and palpus. In H ylecoetus the galea is a broad, rounded lobe, larger in size than the lacinia, and it is covered by dense, long, fine setae which are very dense at its apex.
P a lp ife r . The palpifer is a broad, ring-shaped sclerite from which the palpus arises. The palpifer has several long setae on its mesal surface.
Maxillary Palpus. The maxillary palpus is 4-segmented, covered by dense, long, fine setae, and is sexually dimorphic (Figs. 6, 7).
Female maxillary palpus. In fem ale H ylecoetu s palpus segment I is short, about as long as wide. The second segment is long and slight ly expanded apically. The third is similar to the second, but is nearly parallel-sided. The fourth (apical) segment is also elongate, parallel sided, and apically bears numerous sensilla (Figs. 6, 19).
Male maxillary palpus. The first or basal segment is short as in the female. The second is relatively short, and transverse in form.
The third is abruptly and greatly enlarged (Figs. 7, 8) and is broadly rounded in shape. It is this segment which supports the large maxillary palp-organ discussed below. The fourth and apical segment is very thin and long, and bears several sensilla at its apex (Fig.
19).
Maxillary Palp-Organ. In H ylecoetu s the maxillary palp-organ only occurs in the male, and it is connected basally with the apex of the third maxillary palp segment. The third segment is abruptly enlarged and forms a cup-like structure which is almost entirely membranous. The palp-organ arises to the side and near the hase (relative to the opposing sid e o f segment th r e e l o f segment th re e . In th e memBrane connecting the palp-organ to the palpus there is a small, elongate sclerite I have named the auxiliary sclerite which probably serves as a point of muscle attachment (Fig, 8}. The palp-organ is complex, and multi-branched forming a basket-like structure with pairs of the branches forming opposing sides of the enclosure. The palp-organ is compact, and relatively small in repose. It is very membranous, and when filled with fluids greatly expands, each branch becoming erect and the entire structure protruding from the head in mustache-like form. The exposed area of the palp-organ often is greater than that of the dorsal surface of the head. The inner surface of each branch of the palp-organ is covered by long, thin-walled setae Csee discussion of the palp-organ setae of Melittomma sericeum by Slifer et al, 1975; the setae appear to be similar on most Lymexylidae). The outer surface of each branch is covered by shorter, stouter, fluted, presumably tactile-functioning setae (Figs. 13, 14, 26). In the case of H y le c o e tu s the palp-organ consists for the mostpart of large, primary (1°) branches which support many smaller, secondary (2°) branches (these form the sides of the basket). The surface of these membranous branch es are characteristically wrinkled in appearance.
The Labium and Hypopharyngeal Region (Figs. 15, 20, 21, 31, 32).
The labium o f h y le co e tu s is a small, lightly sclerotized, ventral flap which bears a pair of 3-segmented palpi (Figs. 31, 32). 49.
Mentvaa. The mentura is a sm all, lig h tly s c le ro tiz e d p la te . I t is subquadrate in shape and bears several long setae.
P a lp ig e rs. The palpigers are ventro-lateral in placement and consist of poorly defined sclerites. The proximal, ventral portion of the palpigers are so lightly sclerotized that it is difficult to decide if they are actually fused into a single plate as suggested by
Williams (1938: 257). The palpigers bear several long setae apically and laterally. A large membranous area separates the palpigers from th e mentum and from th e p a lp i. W illiam s ( 1 9 3 8) discussed the similarity between the mouthparts of Psephenus and Hylecoetus. This sim ilarity, however, is simple convergence and is due entirely to a general reduction in mouthpart development. In the case of Hylecoetus, s h o rt lived adults account for this reduction trend.
Labial Palpi. The labial palpi are 3-segmented, elongate, feebly sclerotized, and covered by moderately dense, long setae. The first segment is elongate and more or less parallel-sided. The second segment is similar to the first in diameter, but shorter in length. The third and apical segment is long, parallel-sided, and about as long as I and
II combined in females, and about 80% of the combined length of I+II in males. The apex of the apical segment bears several short, peglike sensilla (Figs. 20, 21). 50.
THE PROTHORAX (F igs. 1 , 33, 34, 35, 73). The prothorax in
H ylecoetus is rather small, and the pronotum is nearly quadrate from
a dorsal view. The prothorax is attached to the head by a wide
cervical membrane, within which are found a pair of elongate, cervical
sclerites. The prothorax is also attached to the mesothorax by a
membranous area, and articulates freely.
The Pronotum. From a dorsal aspect, the pronotum is nearly
quadrate; the posterior margin is slightly sinuate, and the anterior
margin is broadly rounded. The dorsal surface is irregular, with some
areas a little depressed and others elevated into "bumps". The integu
ment, as throughout the body, is lightly sclerotized. The surface is
covered by dense, small, fine punctures and is covered by dense, long,
fine setae.
The Pleural and Sternal Regions of the Prothorax. In H ylecoetu s
the pronotum has distinct lateral margins which are almost carinate in
form. Ventrad to these pronotal lateral-margins the pronotum contacts the prosternum anterior to the procoxae. The procoxal cavities are not well defined (Fig. 34), and are broadly open behind. There are no evident tergo-sternal sutures. The prosternum is short and transverse in form (Fig. 34), and there is no posteriorly directed process between th e coxae.
THE MESOSTERNUM AND ASSOCIATED SCLERITES (Fig. 3 6 ) . The meso- stemum is a small, more or less quadrangular, sclerite situated anter ior to the middle coxae. In H ylecoetu s the mesosternum is simple; in some other genera of lymexylids the central portion is elevated. The 51. mesepistemum is a small, simple sclerite located to each side of the mesosternum; each mesepisternal plate is larger in area than the meso- sternum. The suture separating the mesosternum and mesepistemum is obscure; thus, at a glance, there appears to be a single, transverse sclerite (mesosternum + mesepistemum). The raesepimeron is narrow and lies posterior to the mesepistemum and is elongate, curving from the edge of the mesocoxal cavity anteriorly toward the elytral base. The mesocoxae are cylindrical, in part, but the base is enlarged trans versely toward the lateral margin of the body.
The Mesothoracic Notum. The only structure of the mesothoracic notum which I have used comparatively is the scutellum. Thus, I will confine the description of the mesothoracic notum here to this sclerite.
The scutellum is elongate and broad, and narrowed posteriorly (Fig* 37).
In the case of H ylecoetu s the lateral margins are sinuate, and the posterior margin is emarginate at the middle. Along the midline there is a distinct, double, elevated "keel" which is autapomorphic for this genus.
THE METASTERNUM AND ASSOCIATED SCLERITES (F ig . 3 6 ) . The m etastern - um is a large, quadrangular sclerite. Anteriorly, there are large, shallow depressions (the "coxal depressions") in which the mesocoxae rest. The metepisternum and metepimeron are elongate, and situated p a r a lle l to and l a te r a l to th e m etastem um as shown in fig u re 36. The metepisternum and metepimeron are separated by a longitudinal groove or depressed line. The metacoxae are transverse, with the medial portion short, but cylindrical in cross-section and projecting ventrally. 52.
THE LEGS (Figs. 38, 39., 40, 41, 42, 43, 441. The legs are all
very long and thin in form, and covered by dense, very shallow punctures
and long, very fine setae. The tarsal formula is 5-5-5 in H ylecoetu s
(and in all other genera of Lymexylidae as well]. Because the spurs
at the apex of the tibiae vary in number, it is useful to refer to
these by a tibial spur formula (i.e., the number of spurs at the apex
of the tibia for the proleg-mesoleg-and metaleg respectively) . Thus,
in H ylecoetu s the tibial spur formula is 0-1-1 in both sexes. Sexual
dimorphism in the tibial spur formula does occur in some lymexylids
( e .g ., Atractocerus spp., Melittomma'-complex spp.). In each instance,
however, where dimorphism occurs between the sexes, the male has the
lesser number of spurs.
The coxa of the proleg is long and cylindrical. The trochanter
is small in size, and of the "normal" type in Crowon's terminology
(see Crowson, 1955, p. 88 and fig. 105], The femur is elongate and
about twice as broad as the tibia, which is long and narrow, and nearly
parallel-sided. The tarsus has segment I very long, segments II-IV
shorter, and segment V longer than IV. The apex of the protibia has
a comb of small spines on the anterior margin of the apex (Fig. 40),
and posteriorly the opening is enlarged by an excised area above the
apex (Fig* 39).
Except as noted, the other legs are similar in form to the first pair. The claws are similar on all legs; the unguitractor is large
and conspicuous, and bears several setae. The coxa of the middle leg
is shorter than the procoxa, but is also cylindrical in form. The coxa 53. of the hind leg is transverse in form; the middle (mesal) part is ventrally projecting and more or less cylindrical and the lateral part is shallowly concave, this depressed area receiving the hind femur at r e s t.
THE ELYTRON (F ig s. 45, 731. The e ly tr a in H ylecoetus are s o ft
Cvery lightly sclerotized) and covered by very small, shallow punctures and dense, fine setae. The humeral angle is perceptibly swollen. The epipleural fold (epipleuron) is feebly developed in Hylecoetus. I t is broad beneath the humeral angles, and gradually narrows posteriorly
(migrating dorsally) until it ends a short distance beyond the division between the mesosternum and metastemum. Beyond this point the elytron lies on the dorsal surface of the abdomen, and becomes narrowed toward its apex. Although there is a distinct tongue-and-groove arrangement along the basal part of the sutural margins of the elytra, this ends well before the apex of the elytron. Thus, the tips of the elytra are not in contact with one another. The surface of each elytron has several elevated, rib-like, carinae, A short, oblique, obscure carina runs from what I term the "first carina" to the suture; I interpret this as the "sutural carina" along with a slightly elevated ribbing along the sutural margin of each elytron (it is conceivable that these are modifications of the surtural stria, but this is not clear). Carinae
1 and 2 become fused at or a little beyond the middle of the elytron.
Carina 3 ends near the middle of the elytron. 1+2 and 4 continue posteriorly and become proximal distally, A raised ribbing along the lateral margin is called the "marginal carina". 54.
THE HINDWING AND HINDWING VENATION (Fig. 46L. The hindwings o f
all Lymexylidae are relatively* large, membranous structures which are
sometimes completely covered by the elytra (e.g., Melittowma spp.I,
sometimes partially* covered by* the elytra 6 2 .sr., Australymexylon spp.I,
and sometimes fully exposed, the elytra greatly shortened (Atractocerus
sp p .I. The venation, as a general statement, is primitive, nearly
complete, and fits the Cantharoid-type pattern discussed by Crowson
Cl 955].
Terminology* for the venation of the hindwing of beetles has been prolific and more than a little confusing, I am not so much concerned here for anatomical accuracy* (e.g., does a particular vein represent a single vein with an accompanying one ahsent, or is it the fusion of these two?, etc.] of the terminology, as I am its ability to clearly point to a particular structure. I essentially follow Forbes (1922] and Doyen (1966) as closely as possible. Table 3 compares the terms used by several authors with that employed here; this will help in comparing lymexylid venation patterns to those described in the literature by various authors. Among the systems included are Doyen
(1966), Forbed (1922), Crowson (1955), Graham (1922), Selander (1959),
Kempers (1901), King (1955), A rn ett (1971), and W allace § Fox (1975).
Selander, Kempers, and King are included because they each discuss
Lymexylidae specifically.
In Hylecoetus the costa (C) forms the anterior border of the hind wing. The subcosta (Sc) is short, and fused with C at about ^ the wing-length. The radius (R) likewise becomes fused with C+Sc near the midpoint of the hindwing. The medial vein CM) arises spuriously and connects w ith th e ra d ia l c e ll Cforraed near th e apex o f th e wing by a radial crossvein and the "recurrent radius" CDoyen, 1966)- Rr) by the r-m crossvein. An m-cu crossvein connects M to Cu below the radial cell.
The first anal vein CIA) arises spuriously near the wing base, and the posterior branch of the 1A (IA 2 ) arises about halfway out the wing and appears to be a crossvein at first glance. The wedge cell lies between the la-2a crossvein and 2A, and is a very useful point of reference when recognizing veins in the posterior part of the wing. 3A and 4A are simple veins near the base of the wing; these veins are particularly thickened.
These veins are depicted in figure 46, and summarized in Table 3.
Other useful references to the nomenclature of hindwing venation include
Comstock C19®8), and B orror, DeLong, and T riplehorn (1978). Tabl^ 4>. Hindwing venation systems of terminology compared with terms used here.
Structure Wheeler:here Doyen,1966 Forbes,1922 Crowson,1955 Graham,1922 Selandc
Costa C C CC ■C C
Subcosta Sc Sc Sc Sc Sc Sc
Radial R R R R RR
Recurrent R Rr Rr r (d ista l) - RR
Radial sector Rs - Rs RS R2 - radial cr-v r r r (proximal) - r l r r-m crossvein r-m r-m r-m r-m R3+4 r-m
Medial MMMM R5 M
Cubital Cu Cu Cu Cu Ml+2 Cul
C (at margin) Cu Cu M4+Cu - R5+M1+2 M4+Cul m-cu crossv. m-cu m-cu (M) - R5 M4 (?)
1 st Anal 1 1A1 1A (part) 1A - M3+4 Cu2
1st Anal 2 1A2 1A (part) 2A2 - Cul 1A2 la-2a (cr-v) la-2a la-2a - - - 1A2 T able 4. Hindwing venation (Cont'd.).
Structure Wheeler:here Doyen,1966 Forbes,1922 Crowson,1955 Graham,1922 Selander,1959
2nd Anal 2A 2A 2A3+3A1 - Cu2+1A 1A3+2A1
3rd Anal 3A 3A 3A2 - 2A 2A2
4th Anal 4A 4A 4A1 - 3A 3A
Radial cell RC -- RC - -
Wedge c e ll W wedge c e ll W W - -
Structure Kempers,1901 King,1955 Arnett,1971 Wallace & Fox ,1975
Costa I CC C
Subcosta II Sc Sc Sc-
Radial III RR R
r - m - - r-m r-m
Medial IV MM Rs cn Cubital V Cul CU MP "J T able 4. Hindwing venation (Cont'd.).
Structure Kempers,1901 King,1955 Arnett,. Wallace & Fox,1975
Cu (at margin) - - M4+Cu M2+3+4+Cua
1A1 VII Cu2 1A PCu
1A2 VIII 1A1 2A2 Ala+Alb
2A IX 1A3+2A1 2A3+3A1 Alc+A2
3A X 2A2 3A2 A3
4A XI 3A — A4 59.,
THE METENDOSTERNITE CFigs. 47, 48, 49-1. The metendosternite is one of the most thoroughly studied of the endoskeletal system of adult
Coleoptera. Crowson (1938, 1944, 1955) has compared the structure of this sclerotized process throughout much of the beetle order. One of his conclusions dealt with a basic design of the metendosternite from which he believed many of the modifications seen in Polyphaga could reasonably be modified from. The model he called the Hylecoetoid-type named, of course, for Hylecoetus lymexylids.
The basal part of the metendosternite (the end of which is located between the posterior coxae) is in the form of a long, parallel-sided furcal stalk. Near the base of the furcal stalk is a pair of lateral processes, and ventrally - along its prelaminal length - is a ventral longitudinal flange. Anteriorly, the furcal stalk forks above the lamina and from a narrow, frontal part of the furcal stalk arises the two long, antero-laterally directed (and elevated, dorsally) furcal arms. The anterior tendons are very thin, and arise close to each other between the bases of the furcal arms and the apex of a median laminal process (located on the ventral surface of the lamina). Three views of the metendosternite are seen in figures 47-49,
THE ABDOMEN. The male and female abdomens in Hylecoetus d if f e r considerably in basic structure; this is somewhat of an anomaly within the context of Lymexylidae. They are similar, however, with regard to overall shape, surface sculpturing, setation, and degree of sclerotiza- tion. Postabdominal and genitalic differences are discussed separately below. The Abdomen of the Male (fig. 51}. The male abdomen is elongate, and more or less tubular in form. Each tergite and sternite is rather long, and transverse in shape. The abdomen "telescopes", each segment sliding for a short distance into the segment behind (posterior to) it.
Segments 2-8 are similar in form, and all are for the most part fully exposed; segment 8 is smaller than 7 (see Fig. 51). Segment 9 is fully developed, and forms a capsule or sleeve around the aedeagus.
The form of this interesting abdominal.segment is discussed along with the male genitalia below. Tergite 1 is present and well developed.
A small, "floating" pleural sclerite is present laterally and is quite evident. A small, narrow, transverse sclerite ( w h i c h i s very lightly sclerotized) lies anterior to sternite 2 , and re p re se n ts sternite 1. This is an unusual and plesiomorphic condition among
Polyphagan insects.
The Abdomen of the Female (Figs. 50, 52}. The female abdomen is similar to that of the male in general form, except that it has one less segment. The apical exposed segment is the seventh. The eighth is much smaller, and is telescoped into the abdomen. Ventrally the eighth segment has a very long, thin spiculum ventrale. Abdominal segment 1 also has distinct pleural sclerites (Fig. 50) and a distinct sternite (Fig. 52), in addition to a well-developed tergite
(Fig. 50). The female genitalia are discussed separately below. 61.
THE FEMALE GENITALIA (Figs. 53, 54, 55, 56). Female genitalia of most beetles are not routinely referred to in revisional studies, and have received relatively little attention from coleopterists. In some groups, however, the female genitalia can be useful at some level, whether it is species-level (e.g., Watrous, 1980: study of Lathrobium), species-group level (e.g., Wheeler, 1979a: study of Anisotoma), or the generic level (e.g., Wheeler, 1979b: study of Creagrophorus and the Aglyptinus association of genera, including Aglyptinus and
Scotocryptus). Tanner (1927) remains the only comparative study of female genitalia for Coleoptera which was undertaken on a very broad scale. As emphasized by Tanner, however, this was a preliminary study, and should be continued. A major consideration which became apparent in my study of female genitalia in both Leiodidae (Wheeler, 1979a,
1979b) and Lymexylidae is the need for comparative study using compound m icroscopy. Many u sefu l d e ta ils can sim ply not be handled effectively with a dissecting microscope.
The entire female genitalic apparatus can be referred to as the ovipositor or simply the female genitalia. Both the ovipositor and l abdominal segment VIII are teles^ped into the abdomen at rest. The ovipositor is connected to the eighth abdominal segment by a large, tubular membrane (the intersegmental membrane), which, in most
Lymexylidae, is covered by very dense, small setae. Excluding the various baculi, which are typically very heavily sclerotized, most of the ovipositor is either lightly sclerotized (more or less soft) or actually membranous; further, in many cases it is difficult to 6.21
determine where membrane ends and sclerite begins. Each major part
of the ovipositor is discussed individually below.
Valvifers (Figs. 53, 54, 55}. The valvifers are paired, elongate
in form, and most anterior Ci«e-> basal) in position of all the
ovipositor sclerites. The valvifers are poorly defined(that is, the
transition from membrane to the valviferal plate is not abrupt), and ventro-lateral in position Cthey occupy much of the space on the ventral surface for their length of the ovipositor, and cover most of the side as well). The anus opens dorsally between and at about the posterior limit of the valvifers. The elongate valvifers approach the midline ventrally, but do not come into contact. Ventrally, they give rise to a pair of valviferal baculi which, according to Doyen
C1966), function as an articulation for the coxites. In Hylecoetus the valvifers are nearly as long as the coxites.
Coxites (Figs. 53, 54, 55}. The coxites are paired sclerites located posterior (distal) to the valvifers (see figures). The coxites consist of a single sclerite dorsally, but are divided ventrally by a coxital fold (pigs. 53, 54) into a distal part and a basal part, which are subequal in length in Hylecoetus. Each distal part bears an unsegmented, elongate, slightly clavate stylus near the apex. The stylus bears one very long seta at its apex, and several shorter ones
(pig. 56) over its surface.There are also several small, peg-like sensilla. The dorsal and ventral surfaces of the distal part of the coxite bear numerous long, fine setae set in large, membranous sockets. 63.
The basal part of each coxite gives rise to a coxital baculus which articulates with the valviferal baculus which is located anterior to the coxital baculus. The surface of the basal part is covered by numerous setae on its distal half.
Fungus-pouch (Fig. 53) . The fungus pouch o f Hylecoetus is located, for the mostpart, between the basal parts of the coxites, although it extends laterally from the midline internally. It is a membranous infolding and includes a medial groove and lateral extensions (the
"pouches") within the basal parts of the coxites and the valvifers
(in part). Francke-Grosmann (1967) used the term fungus-pouch.
The inner margins of the distal parts of each coxite are lined with very dense, fine, small setae. Dorsally, the entire inner margins of the coxites plus the membranous area between them is clothed by similarly dense setae.
These pouches are known to hold spores of the host fungus,
Ascoides hylecoeti, which are apparently extruded at the time of oviposition (see Francke-Grosmann, 1967 and Natural History below ).
Proctigeral Baculus (Figs. 54, 55). The paired, proctigeral baculi arise dorso-medially and perhaps represent the proctiger (see
Doyen, 1966). These baculi are about the same length as the valviferal baculi, and their anterior apices are poorly sclerotized, becoming expanded into several poorly delimited lobes.
Coxital Baculus (Figs. 53, 54). The c o x ita l b a c u li, mentioned above under the discussion of the coxites, are short, oblique, narrow structures which are situated just posterior (distal) to the valviferal Table S . Terminology for female genitalia and associated structures, see Figs. 53-55.
Term Description Source Synonyms
Valviferal baculus (-i) Long,heavily sclerotized processes;paired; Proposed here baculus (Doyen, on ventral side of genitalia and arising 1966);ventral from the valvifer plates baculus (Ekis, 1977)
Coxital baculus (-i) Small, usually oblique, sclerites located Proposed here baculus (Doyen, at base of each valviferal baculus, and 1966);oblique bac arising from base of coxites ulus (E kis,1977)
Proctigeral baculus Long, scle ro tized processes;paired;on Ekis, 1977 proctiger (Doyen, (~i) dorsal side of genitalia and arising"from 1966) proctiger (or proctigeral remnant plate)
Intersegmental mem Long, large, tubular membrane connecting Doyen, 1966 brane genitalia to abdominal segment VIII; typically very hairy in Lymexylidae
Connecting membrane N on-sclerotized or very lig h tly scle ro tized Doyen, 1966 areas of the genitalia between sclerites and baculi; most conspicuous basad of valvifers, to about apex of valviferal baculi
Coxite (-s) Sclerites surrounding the vulva and Tanner,1927 Hemisternites (Lin- supporting the styli; distad to valvifers; droth,1957);Valvi- probably remnants of sternite IX. fer 2 (Doyen,1966)
O' T able 5. Terminology for female genitalia (Cont'd.).
Term Description Source Synonyms
Basal part (-s) Portion of coxite before (basal) the Proposed here - (of coxite) coxital fold (in Lymexylidae)
Distal part (-s) Portion of coxite beyond (distal) the Proposed here (of coxite) coxital fold (in Lymexylidae)
Coxital fold (-s) Transverse fold near the middle of each Proposed here - coxite on the ventral surface, separat ing a basal and distal portion of each coxite;fold is incomplete:absent dorsally
Valvifer (-s) Sclerite (paired) basad to coxites; Tanner,1927;and,Valvifer 1 (Doy- probably represents halves of tergite IX Lindroth,1957 en,1966)
Medial sclerite (-s) Longitudinal, single, generally lightly Proposed here Ventral lamina sclerotized process lying between coxites (Ekis,1977) on ventral side of genitalia
Stylus (~i) Paired, small, sclerotized processes which Tanner,1927; Gonostylus (Doyen, articulate with coxites; usually bearing Lindroth,1957 1966);Coxital sty many setae and apical in position lus (Ekis,1977)
Fungal pouch Specialized, membranous organ of lymexylid Francke-Gros- Vaginal pocket, female g en italia which functions in s to r - mann,1967 (Buchner,1928) age of spores; located between coxites
Ventral spicule of segment VIII of abdo Lindroth,1957; - Spiculum ventrale O' men; single and very long in Lymexylidae Doyen,1966 tn f i f i ,
b a c u li.
Abdominal Segment VIII. The eighth abdominal segment is small,
and intimately associated with the female genitalia. The ventral
strut (spiculum ventrale) is similar in structure throughout the
Lymexylidae, and serves as a point for muscle attachment.
THE MALE GENITALIA AND POSTABDOMEN (Figs. 57, 58, 59). The male
genitalia (referred to en to to as the "aedeagus") of beetles has been
found particularly useful by coleopterists. The excellent comparative
study made by Sharp and Muir (1912) is still a standard reference work.
The male genitalia of Hylecoetus were discussed by Crowson (1955), who
emphasized their unusual structure. The anomaly of these, however, is
as true by contrast to other Lymexylidae as it is to other Coleoptera.
My choice of terminology in reference to male genitalia diverges to
a small degree from that usually accepted. I have tried to follow,
as closely as possible, the preferred terms of Sharp § Muir (1912).
Lindroth (1957) provided an excellent, and long-needed, comparative review of the terms used for male genitalic components. Some special
structural modifications in the Lymexylidae, however, required some
changes in terms. My selections, and their relationships to terms used by other authors are summarized in Table 5. The table should
alleviate some confusion with regard to terms.
An hypothesis on the origin of unusualities in the aedeagus of
Hylecoetus beetles. Before describing the various parts of the male
genitalia, it would be useful to discuss certain unusual aspects vof 67.
H ylecoetu s male genitalia. During my study of these structures I developed an hypothesis which I feel adequately explains these modi fic a tio n s .
Crowson (1955) discussed the fact that the lateral lobes of
H ylecoetu s are asymmetrical. That is, both lateral lobes lie proxi- mally to one side of the tegminal plate. What might underlie this bizarre placement of the lateral lobes which are symmetrical in all other beetles that I have seen in Lymexylidae and in both Cleroid and Cucujoid lines?
The usual position for the aedeagus within the abdomen for the lymexylid beetles is the in v e r te d position described by Crowson (1955).
This means the tegminal plate lies next to the sternites (ventrally), and the median lobe curves upwards (toward the dorsum) inside the abdomen. The inverted orientation is usually associted with beetles which copulate with the male back to the female. The alternative, uninverted position usually is found in forms which position themselves with the male on top of the female when copulating.
In H ylecoetu s the aedeagus is rotated 90° to its former position
(and the position found in other Lymexylidae: the in v e r te d position).
Now the median lobe curves toward the right side of the beetle's abdomen (not the dorsum). The tegminal plate, formerly lying next to the sternites, is now in a lateral position along the left side of the abdomen. Thus, the lateral lobes have compensated by migrating as far possible toward the right. This brings them very close to the original position, lying along the sternites in a ventral position. 6 8 .
From this position, the lateral lobes curve to the left side. This o means that when the aedeagus is extruded, and rotated 90 back to its former position that the lateral lobes curve upwards, just as they do in o th er lym exylid b e e tle s .
Why Hylecoetus store the aedeagus at 90° rotation is not clear.
But, it is very interesting that the lateral lobes have become asymme trical to offset this rotation.
It is also of interest that the capsule-like or sheath-like abdominal segment IX has sim ilarly become asymmetrical. As seen in figure 58, the tergo-sternal suture is oblique, and the tergum is twisted so that part of it is actually lateral in orientiation. This is apparently associated with the rotation of the aedeagus.
Median Lobe (Figs. 57, 58, 59). The median lobe is very long and thin, although it is broader at the base and narrowed gradually toward the apex. The base is recurved, and the tip is gradually curved. In the abdomen, the median lobe curves toward the right side; when exerted its curvature is upwards (toward the dorsum). The median lobe slides outward through a sleeve-like mechanism created by the tegmen.
Tegmen: the Tegminal P late. The tegmen of the male genitalia of beetles consists of the lateral lobes, and a basal part against which they can articulate (although the ability to articulate is lost in some beetles). In the lymexylids the part of the tegmen basal to the lateral lobes can conveniently be discussed as two major components which are a single, plate-like structure and a pair of struts. 69.
The tegminal plate is a broad sclerite which typically lies on the ventral side of the median lobe, although in Hylecoetus i t has rotated in repose to a position alon the right side of the abdomen.
The plate is poorly defined, so that it is difficult to see the limits of the plate, and where the surrounding membranous areas begin. This tegminal plate articulates with the asymmetrical lateral lobes of
Hylecoetus and also gives rise to the tegminal struts.
Tegmen: the Tegminal S tru ts (Figs. 57, 58) . The tegminal struts are a pair of large, appendiculate structures which connect the tegminal plate to the base of the median lobe. This connection is by soft, unsclerotized tissue so that clearing in KOH or other caustic material causes a separation of the tegminal struts and median lobe base. This connection, however, allows the median lobe to move, sliding through the opening between the plate and struts.
Tegmen: the Lateral Lobes (Figs. 57, 58, 59). The lateral lobes of most of the Cucujoid, Cleroid, and Lymexyloid beetles are small, lobe-like structures arising from the tegminal plate; although there are many subsequent modifications (e.g., the tube Heteromeran type of aedeagus described by Crowson, 1955). In tiylecoetus, the lateral lobes are asymmetrical in position. (There are some interesting modifications in other lymexylids, perhaps the most impressive being the huge lateral lobes seen in Atractocerus spp.). Table 6. Terminology for male genitalia (aedeagus) and related postabdominal structures. Figs.57-59.
Term Description Source Synonyms
Median lobe Distal, unpaired portion of aedeagus which Sharp & Muir, Penis (Lindroth, contains the endophallus and through which 1912 1957); aedeagus, the ejaculatory duct passes sensu stricto
Lateral lobe (-s) Distal part of the tegmen; usually paired Sharp & Muir, Parameres (Lin 1912 droth, 19 57)
Basal piece Basal portion o f the tegmen (e .g ., see Sharp & Muir, Wheeler,1979a),ringlike in some primitive 1912 taxa,e.g., Staphylinoidea; NOT recognized as such in Lymexylidae (= MY tegminal plate + tegminal struts ?)
Tegminal p la te Large (usually), basal part of the tegmen, Proposed here Tegmen (Sharp & gives rise to both lateral lobes and the Muir,1912) tegminal struts (Usually ventrally placed)
Tegminal s tru ts Paired, appendiculate stru ctu res arisin g from the tegminal plate; these struts typically connect the plate with the base o f the median lobe
Aedeagus The entire genital apparatus of the male, Sharp & Muir, - consisting o f the median lobe, and tegmen 1912 (in Lymexylidae: tegminal plate + tegminal struts + lateral lobes)
Endophallus Membranous sac which is typically inverted Internal sac (Sharp in to the median lobe and which i s extruded & Muir,1912 and "•i o T able 6. Terminology for male genitalia (aedeagus J (C o n tld%),
T erm Description source Synonyms
Endophallus (cont' d.) during copulation; the opening of the Lindroth,lH57) ejaculatory duct is usually on the endophallus; in Lymexylidae the endo phallus is often partially exposed ventrally, under the tegminal plate
Abdominal segment IX In Hylecoetus IX forms a sleeve around the aedeagus; this segment is usually greatly reduced, often only represented by small tergal and sternal p la tes and some lateral elements
Abdominal segment VIII Segment VIII forms a sleeve around the aedeagus in most forms, other than Hylecoetus; IX being greatly reduced
Median process The middle part of the lateral lobes Proposed here - in Atractocerus spp. (see Fig. 167)
Lateral denticuli One to several pairs of tooth-like Proposed here - processes located between the median process and the lateral process of the lateral lobes of Atractocerus spp.
Lateral process Long and often cup-like or arm-like Proposed here - major processes of the lateral lobes in Atractocerus spp. (see Fig. 167) Figs. 2-5. Hylecoetus dermestoides, head: fig. 2, dorsal view; fig.3, ventral view; fig. 4, posterior view; fig. 5, lateral view.
72. MAXILLARY PALPUS
LAB? UM LABIAL PALPUS • • / POSTERIOR /MANDIBLE TENTORIAL LABRUM
,GENA
ANTENNA
GULAR SUTURE
I EPICRANIAL
LABRUM VERTEX FRONS / MAXILLARY PALP- OCCIPITAL. / ORGAN ✓FORAMEN v ✓ GENA V' \ \ *
_ / ANTENNIFER ANTENNAL SOCKET Figs. 6-9. Hylecoetus dermestoides, maxilla: fig. 6, female; fig. 7, male; fig. 8, male, palpomere III and bases of palpomere IV and maxi llary palp-organ; fig. 9, apex of secondary branch (2°) of maxillary palp-organ. SECONDARY\ BRANCH (2*) MAXILLARY PALP-ORGAN—. __
~ ~~ "SENSILLA — ^
------PALPOMERE IV------
PALPOMERE 11
* ------PALPOMERE II
— -PALPOMERE I GALEA ------"PALPI FER PRMARY BRANCH (W (MAX. PLR ORGAN) ’ CARDO — — PALPIFER AUXILLARY SCLERITE
'LACINIA'' STIPES'
MAXILLARY MPO(2°) PALP O R G A N ------PALPOMERE IV
PALPOMERE 111 AUXILLARY SCLERITE
" 4 Figs. 10-12. Hylecoetus dermestoides: fig. 10, antenna; fig. 11, labrum (dorsal); f ig . 12, labrum and epipharynx (v en tra l).
76. SETAL BRUSH -XI LABRAL SETA
-X LABRUM
-IX
VIII
/ / - ' CLYPEO-LABRAL \ \.A / MEMBRANE > ^ -VII TORMA.
SETAL BRUSH --VI
-V
-IV TRICHIAL LABRUM RIDGE "
Ill
PEDICEL(II)
TORMA
/ \ SCAPE (I ) EPIPHARYNGEAL MICROTRICHIAE MEMBRANE Figs. 13-14. Hy1ecoetus dermestoides, maxillary palp-organ; fig. 13, outer surface; fig. 14, inner surface. 79. Figs. 15-16. Hylecoetus dermestoides: fig. 15, hypopharyngeal setae fig. 16, epipharyngeal setae.
Figs. 17-18. Hylecoetus dermestoides: fig. 17, apex of labrum (ven tral); fig. 18, antennomere VI, surface detail. 83. Figs. 19-20. Hylecoetus dermestoides: fig. 19, apex of maxillary palpomere IVj fig. 20, apex of labial palpomere III.
Figs. 21-22. Hylecoetus dermestoides: fig. 21, "peg" sensilla at apex of labial palpomere III; fig. 22, "flabellicornis" morph, antenna.
86.
Figs. 23-24. Hylecoetus dermestoides, "flabellicornis" morph, flabell ar process of antenna: fig. 23, outer surface; fig. 24, inner surface
Figs. 25-26. Hylecoetus dermestoides, "flabellicornis" morph: fig. 25, apex of maxillary palpus; fig. 26, surface of maxillary palp-organ. 91.
(t L V
M Figs. 27-28. Hylecoetus dermestoides: fig. 27, mandible (dorsal); fig. 28, maxillary palpomere IV.
92.
Figs. 29-30. Hylecoetus dermestoides; fig. 29, antennomere VII sur face; fig. 30, epicranial pit.
94.
Figs. 31-32. Hylecoetus dermestoides, labium: fig. 31, lateral view; fig. 32, ventral view.
96. -SENSILLA
— Ill------
LIGULA
PALPI GER
MENTUM * \ C31 Figs. 33-35. Hylecoetus dermestoides: fig . 33, pronotum, scutellum, and bases of elytra (dorsal); fig. 34, prothorax (ventral); fig. 35, prothorax (lateral). Fig. 36. Hylecoetus dermestoides, mesostemum, metasternum, and asso ciated structures (ventro-lateral view).
99. MESOSTERNUM MESEPISTERNUM j MESEPIMERON TROCHANTINs s ' EPIPLEURON -COXAL m e s o c o x a CAVITY
FEMUR— METEPISTERNUM
TIBIA— — ELYTRON
COXAL DEPRESSIOI
METEPIMERON METASTERNUM 36
METACOXA TROCHANTER Fig. 37. Hylecoetus dermestoides, scutellum (metathoracic), dorsal view.
101.
Figs. 38-44. Hylecoetus dermestoides; fig. 38, prothoracic leg; fig. 39, apex protibia, inner margin; fig . 40, apex protibia, outer margin; fig. 41, mesothoracic leg; fig. 42, metathoracic leg; fig. 43, claw; fig. 44, metatibia, apex and spur. •COXA TIBIA' FEMUR
TROCHANTER . FEMUR- TROCHANTER
TIBIA-
TIBIAL COMB"
TIBIAL SPUR
-TARSUS -TARSUS
TIBIAL SPUR,
CLAW- 104 HUMERAL -SCUTELLARY 105. ANGLE-^ KEEL
. SCUTELLARY CARINA 2 CARINA
-SUTURAL CARINA
CARINA 1
MARGINAL CARINA . r f CARINA 1*2
CARINA 3 CARINA 4
45
Fig. 45. Hylecoetus dermestoides, left elytron and scutellura (dorsal). 106 46 7 RADIUS I CUBITUS / I I , • iVicdiocubital v. cn ! '' ' V V ANAL BRANCH,FIRST ' nANTERIOR ' \ V V / / / \ POSTERIOR BRANCH, FIRST ANAL \ I I \ Fig. 46. Hylecoetus dermestoides, hindwing (see Table 4 ). Figs. 47-49. Hylecoetus dermestoides, metendostemite: fig. lateral view; fig. 48, dorsal view; fig. 49, ventral view. 108. FURCAL ARMS =L~ N) LATERAL PROCESS ANTERIOR TENDONS BASE LAMINA 47 VENTRAL LONGITUDINAL FLANGE RIGHT ARM LEFT ARM RIGHT ARM ANTERIOR ANTERIOR TENDONS TENDONS FURCAL ARMS ■= — MEDIAN LAMINAL PROCESS --LAMINA---- FRONTAL PART OF STALK FURCAL STALK -VENTRAL LONGITUDINAL FLANGE LATERAL PRO CESS------ BASE 48 49 Figs. 50-52. Hylecoetus dermestoides: fig. 50, female abdomen (lateral); fig. 51, male abdomen (lateral); fig. 52, female abdomen, base (ventral). Abbreviations: T1-T9 (tergites I-IX), S1-S9 (stern- ite s I-IX ), PI (pleural s c le r ite o f abdominal segment I ). 110. I Figs. 53-55. Hylecoetus dermestoides. female genitalia (ovipositor) fig. 53, ventral view; fig. 54, lateral view; fig. 55, dorsal view. 54 55 -INTERSEGMENTAL MEMBRANE INTERSEGMENTAL MEMBRANE -CONNECTING CONNECTING MEMBRANE MEMBRANE PROCTIGERAL BACULUS -PROCTIGERAL BACULUS VALVIFERAL BACULUS VALVIFERAL BACULUS VALVIFER VALVIFER COXITAL BACULUS -VALVIFER / COXITAL / FUNGAL POUCH BACULUS i// -COXITE ^COXITE (BASAL PART) ^ (BASAL PART) .COXITAL COXITE ^-COXITAL FOLD FOLD COXITE (DISTAL PART) COXITE (DISTAL PART) X ^MEDIAN GROOVE (FUNGAL POUCH) -STYLUS « STYLUS 112 Fig. 56. Hylecoetus dermestoides, stylus of female genitalia. 113. Fies. 57-59. Hylecoetus dermestoides, male genitalia (aedeagus) and abdominal segment IX: fig. 57, lateral view; fig. 58, dorsal view; fig. 59, ventral view. 116. ENDOPHALUJS- ^.TEGMINAL PLATE LATERAL LOBES MEDIAN 1 LOBE TEGMINAL STRUT ^MEDIAN LOBE ENDOPHALLUS 'TEGMINAL STRUT . . - j r f i i STERNAL i i i SUTURE' '' ' I ‘ < 7 \ \ MEDIAN LOBE I I 1 / / , 7 ^ - t EGMINAL p late TERGITE VIII j2--ENDOPHALLUS LATERAL LOBES' 58 lateral lobe H y le c o e tu s d e r m e s to id e s : LARVAL STRUCTURES GENERAL DESCRIPTION OF THE LARVA (Fig. 60) . The larva is elongate, and thin in form. The head is small, and more heavily sclerotized than other parts of the body (with the exception of the modified ninth abdominal segment). The pronotum is very much enlarged, the head being retractable under this hood-like pronotum. The meso- and meta thoraces are smaller, and similar in form to the abdominal segments following them. The ninth abdominal segment is greatly modified into a very long, thin, sword-like structure which is direct posteriorly. The tenth abdominal segment is quite small, and located ventrally, below the eighth and ninth segments; the anus is located on the tenth segment. The legs are small in comparison to the body as a whole. These remarks, and the structures described below are based on a late-instar larva. The neonatal larva, or 1st instar larva, differs in several respects, and has been described in detail in an excellent paper by Grandi (1961). The Cranium. The cranium is moderately sclerotized, and oval in shape viewed from a dorsal or ventral perspective. The dorsal surface is covered by sparse, long, fine setae of various lengths. A short, medial, longitudinal suture begins near the posterior margin of the head and extends anteriorly for a short distance; no other epicranial 117, 118. sutures are evident. Although ocelli are present in first instar larvae (Grandi, 1961), they are absent in later instar larvae. The fact that first instar larvae have to choose a boring site and make initial penetration of the wood, this is perhaps understandable. Antennae. The antennae are minute in size, and are located above and behind (postero-dorsal) the bases of the mandibles. Each antenna appears to be only 2-segmented. The actual first segment is indistin guishable from the membrane connecting the antenna to the cranium. The second segment (and first visible one) is very short and broad, almost ringlike in form and bears a large digitiform sensillum (Dybas, 1976; = supplemental process of Peterson, 1961), and about four smaller sensilla. The second visible segment (segment 3) is only slightly longer than the digitiform sensillum of the preceeding segment,and bears about four apical sensilla, two of which are longer than the o th e r s . Labium and Epipharyngeal Region (Fig. 65). The labrum i s elo n g ate, simple, and rounded at its apex; the labrum is located dorsad of the mandibles medially. There are two pairs of large and broad setae on the ventral surface (the lateral setae in Fig. 65), a few intermediate sized setae, and very dense, small, fine setae converging along the midline, and extending over the medial surface of the membranous epipharynx. A number of parallel ridges run from the midline to the lateral margins of the epipharynx; these are sinuate, and located posterior to the setae mentioned above (the epipharyngeal comb in F ig, 65). 119. M andibles. The mandibles are broad in form, rather short in length, and very heavily sclerotized. On the mesal surface, near the base of each mandible is a distinct molar region. This mola has several large teeth apically, but at its base consists of parallel, transverse grooves, and to the side of parallel rows of microscopic spines. Dist- ally there is a small area of several spines which may represent a remnant of the mandibular prostheca. M axillae. The maxillae are lightly sclerotized, and covered by numerous setae and spines. The cardo is a small, elongate, basal sclerite which is set off by a distinct suture from the stipes. The stipes is narrow basally, and expands distally. The stipes is not divided into a basistipes and dististipes, and there is no clear suture between the stipes and the mala Qacinia + galea). The mala or maxillary lobe, is partially divided apically into a broad, inner lacinial area and a narrower, outer galeal area. The mesal surface of the lacinial area has a double row of setae along its margin; the area between these two parallel lines of setae Cthe m alar com b) appears to be less sclerotized than the dorsal or ventral surface of the mala. The p a r tia l d iv isio n o f th e mala was mentioned by Crowson C1955, 1964) in the context of Lymexylidae. This division does not occur widely in the family. The malar comb, however, seems to be a fam ilial-level fe a tu re . Labium. The labium is small and very lightly sclerotized. Between the bases of the palpi there is a narrow, heavily sclerotized region. 120. The labial palpus consists of two palpomeres: I long and about as wide as palpifer (the separation of I from the palpifer is sometimes obscure); II is very short, with a patch of short, apical sensilla (palpomere I has several stout setae on its surface). The ligula is broad, membran ous and covered by microtrichiae which are very dense, (see Fig. 71). The Legs (Figs. 62, 63). The legs are small, and broad in form. The coxa is elongate, with several long setae posteriorly; the tro chanter is scarcely longer than broad, with several long setae; the femur is not much longer than the trochanter, and similarly setose (but with more shorter setae); the tibia is elongate and covered by dense, short- to moderate-length, stout setae. The tarsungulus (Fig. 62) is pleurisetose. Apex of Abdomen (Figs. 60, 61). Abdominal segment IX is heavily sclerotized, and very long, narrow, and forked apically bearing stout teeth along dorsal margins. Segment X is small, membranous, ventral in position, and covered by short spines. Segment VIII is simple, and has several stout teeth dorsally (segment VII also has a few teeth). Fig. 60. Hylecoetus dermestoides, larva (lateral view). Drawing by J . Nagy. 121. Pia*. 61-64. Hylecoetus dermestoides, larva: fig.^ 61, posterior^end of abdomen (segments 7“ 0); fig. 62, tarsungulus of proleg; fig. 63, proleg; fig. 64, antenna. COXA K w W J f t W F S•*: ?- j r . * * ' ^-'TROCHANTOR FEMUR 64 -,/ tibia 63 DfGITIFORM ORGAN "TARSUNGULUS 124 Fig. 65. Hylecoetus dermestoides, larva, labrum and epipharyngeal region (ventral). 126. LATERAL SETAE s . i 65 127. partial division / GALEA LACINIA III— .MALAR COMB II— MALA PALPIGER STIPES 66 CA R DO Fig. 66. Hylecoetus dermestoides, larva, maxilla. Apex of stipes, palpus and mala (above) and base of stipes and cardo (below). PARTIAL DIVISION LACINIA GALEA 67 Fig. 67. Hylecoetus dermestoides, larva, apex of mala showing partial division of lacinia and galea. 129. Fig. 68. Hylecoetus lugubris, larva, mandible, left-mesal surface. Scanning electron micrograph courtesy of J.F. Lawrence, CSIRO, Division of Entomology, Canberra City. 130. PROSTHECAL SPINES 69 i i i MjDLA i * I I / - I i ...... if. MICROTR ICHIAl! ROWS Figs. 69-70. Hylecoetus dermestoides, larva: fig. 69, prosthecal spines and molar region of mandible; fig. 70, left mandible, dorsal view. ft Fig. 71. Hylecoetus dermestoides, larva, labium (ventral). 71 Hylecoetus derm estoides: PUPAL STRUCTURES An illustration of the pupa (Fig. 72) is included, but a detailed description is not presented. The pupa is lightly sclerotized and the head and pronotum are covered by very small, dense spines. The pupa is similar in proportions to the adult, and can readily be sexed by the presence or absence of the maxillary palp-organs (clearly visible in Fig. 72). 133. Fig. 72. Hylecoetus dermestoides, pupa (v en tra l). Drawing by J. Nagy. 134. TAXONOMIC CONSIDERATIONS Below, I present a diagnosis for the family Lymexyl idae, and a key to the subfamilies and genera of the world, Following these items are a series of chapters on each genus of the family which consist of generic diagnoses and descriptions of at least the type- species for each genus, based on both larval and adult structures. The genera are arranged in accordance with my classification of the family, which is summarized in Table 7 in the form of a checklist. For discussion of the derivation of this classification and a phyletic sequence of the genera see the chapter on cladistics below. FAMILY LYMEXYLIDAE Lymexylonidae Leach, 1817: p.? Malacodermi (in part) Gyllenhal, 1808:314 Xylotrogi (in part) Latreille, 1825: 354 Malacodermata (in part) Perty, 1830-1834: 25 Oedomeridae (in part) Stephans, 1832: 52 Lymexylonitae Castelnau, 1836: 58 Atractocerites Castelnau, 1840: 290 T e rd ile s Boheman, 1848: 519 Lymexylones Redtenbacher, 1849: 5 § 36 Ptinidae (in part) Melsheimer, 1853: 86 Lymexylonidae, J. Thomson, 1858: 82 Lymexylonides Redtenbacher, 1858: L § CII Lyttidae (in part) Walker, 1858: 285 Lymexylidae LeConte, 1861-1862: XXIV § 198 Diversipalpes Mulsant § Rey, 1863: ? Xylophaga (in part) Gerstaecker, 1873: 160 Lymexylini Heyden, R eitter, 8 Weise, 1891: 228 LymexylonidaejSchenkling, 1915: 1 Lymexylidae; Leng, 1920: 152 Lymexylidae; Barber, 1952: 163 Consult Crowson (1955) for a discussion of the separation of the superfamilies Lymexyloidea, Cleroidea, and Cucujoidea. Below is a diagnosis of the family. DIAGNOSIS. ADULT BEETLES- Procoxae c y lin d ric a l, p ro je c tin g ; t a r s i 5-5-5; antennae filiform to serrate, rarely flabellate; males usually with maxillary palp-organ arising from apex of palpomere III, rarely present in females also (Atractocerus spp.) and absent in both sexes (Australymexylon); head usually narrowed behind eyes forming "neck" region (neck absent in Hylecoetus); aedeagus of tilobe form, sometimes highly modified; female genitalia with coxites divided by coxital fold into basal part and distal part (division is incomplete dorsally); elytra usually with ribbing (i.e., carinae: typically with 4 carinae plus oblique sutural carinae), sometimes brachelytrous (Atractocerus); female sternum VIII with very long, narrow, anterior spiculum gastrale; male sternum VIII usually with paired struts, often fused; LARVAE- head partially retracted under hoodlike pronotum (pronotum much larger than mesonotum or metanotum); legs very short; antenna minute; ocelli absent in later-instar larvae; each abdominal segment with epipleural fold; abdominal segment IX very large, extending above and posterior to X, and either membranous and bulbous, long and swordlike, or cylindrical, and trunctate (and highly sclerotized); maxilla with mesal margin membranous and bordered by long setae, often in a double, parallel row; ECOLOGICAL RELATIONSHIPS- A ll b eliev ed to be sym biotic w ith am brosia fungi, or other microbes (including bacteria complex). 139. KEY TO SUBFAMILIES AND GENERA OF LYMEXYLIDAE 1 Head with epicranial pit (Tigs* 2, 30),rounded behind eyes without "neck" (Fig. 2); elytra long, surpassing metacoxae (Fig. 73); male abdomen with one more visible sternite than female (Figs. 50, 51); lateral lobes asymmetrical (Figs. 57-59); eyes round, small, entire (Fig. 2); LARVA- abdomi nal segment IX very long, narrow, heavily-sclerotized, "swordlike" (Figs. 60, 61); HOLARCTIC ------Subfamily Hylecoetinae ------Genus Hylecoetus 1' Head without epicranial pit; head narrowed behind eyes forming "neck"; elytra long to brachelytrous; male and female- with equal number of sternites visible; lateral lobes symmetrical; eyes moderate to large, emarginate anteriorly; LARVA- abdomi nal segment IX large, bulbous (Fig. 148) or cylindrical- truncate (Fig. 251) ------2 2 Elytra shortened (Fig. 82) or brachelytrous (Fig. 117); without tongue-and-groove locking mechanism along suture; lateral lobes short and truncate (Fig. 99) or very large and broad or elongate (Figs. 165, 167); LARVA- abdominal segment IX large, membranous, "bustlelike", often with asperites on apex (Figs. 112, 148); PALEARCTIC (Lymexylon) o r PANTROPICAL ------Subfamily Lymexylinae—3 2' Elytra covering abdomen or only slightly shortened (Fig. 179), tongue-and-groove mechanism present, sometimes ending before apex of elytra; lateral lobes large and concave medially (Figs. 197-200) or small, lobe-like (Figs. 233,234), rarely elongate with apical row of teeth (Fig. 258); LARVA- afldominal segment IX heavily sclerotized, cylindrical and tru n c a te (F ig. 251); NORTH AMERICA-AUSTRALIA-PANTROPICAL ------Subfamily Melittomminae ------4 3 Elytra long, surpassing metacoxae (Fig. 82); male with maxillary palp-organ (Fig. 97), female maxilla without organ; lateral lobes short, truncate (Fig. 99); female genitalia with spurious baculus joining valviferal baculus (-Fig. 106); female genitalia lightly sclerotized; LARVA- abdominal segment IX with apical "tooth" (Fig. 113); tarsungulus bisetose (Fig. 115); PALEARCTIC ------Genus Lymexylon 14Q. 3' Elytra brachelytrous, never reaching metacoxa (Fig* 117); male and female with maxillary palp-organ ( s m a l l e r f emale , Fig. 124); antenna short, spindle-like (Figs. 119, 123); lateral lobes very large and broad (Fig. 167) or elongate (Fig. 165); hindwing without crossveins, folding only longitudinally (Fig. 157); eyes very large (Fig. 118); female genitalia heavily sclerotized (in part); LARVA- abdominal segment IX with apical aspirites (fig. 148); tarsungulus with several setae (fig. 147); PANTROPICAL------Genus A tractocerus 4 Eyes moderate-sized, widely separated (Fig.179); maxilla simple (Fig. 210), maxillary palp-organ lacking in male and female; male abdominal sternum eight with setose pads (Figs. 191-193); lateral lobes of male genitalia large, concave medially (figs. 196-200); LARVA- abdomi nal segment IX heavily sclerotized apically, with a few minute teeth around margin (Fig. 201); AUSTRALIA— ------Genus Australymexylon, new g. 4' Eyes large, often proximal; maxilla of male with maxillary palp-organ (Figs. 227, 228); male abdominal sternum eight simple; lateral lobes variable, never concave mesally (Figs. 233, 258); paratergites of abdominal segment nine sometimes appendiculate CFigs. 235, 248); LARVA- most of abdominal segment IX heavily sclerotized, apex varies from n e a rly smooth, to to o th ed ; PANTROPICAL, NORTH AMERICA------Mel i t tomma-Comp 1 ex ------5 5 Adults usually soft-bodied (Fig* 217), rarely heavily sclerotized (Fig. 241,242); male abdominal paratergite IX appendiculate (Figs. 235, 248); lateral lobes of aedeagus small, lobate, setose (Figs. 234, 245); female maxillary palpomere IV narrow apically; mesostenrum simple anteriorly, or elevated and "A"-shaped; LARVA- apex of abdominal segment IX with deep notches around edge delimiting "teeth" (F ig. 251) ------PANTROPICAL Genus Melittomma 5' Adults heavily sclerotized (Fig. 256); abdominal paratergite IX of male simple (in known males); lateral lobes of aedeagus large with apical row of stout teeth (Fig. 258) (in known males); female maxillary palpomere IV narrow or securiform; LARVA- apex of abdominal segment IX nearly smooth, with a few minute spines (in known larvae) ------141, 6 Female maxillary palpomere IV securiform (Fig. ); meso- stemum with anterior area elevated, elongate and very narrow (pointed apically) (Fig- ); basal half of stylus of female genitalia sclerotized, apical % membranous; distal part of coxite of female genitalia with narrow, basal sclerite but otherwise membranous (Fig. ); male unknown; LARVA- unknown; SOUTH AMERICA ------Genus M elittommopsis 6' Female maxillary palpomere IV narrow apically (Fig. ); mesosternum with anterior area elevated "V"-shaped (in- verted); proctigeral baculus of female genitalia fused for much of length (Figs. 274,275); lateral lobes very large, elongate, with rows of teeth apically (Fig. 258); LARVA- apex of abdominal segment IX nearly smooth around edge, but with a few minute spines (Fig. ) ------— MADAGASCAR, SEYCHELLES------Genus Melanomelittomma, new genus Table 7. A checklist of the subfamilies and genera of Lymexylidae. See cladistics section for a discussion of relationships of the taxa. Subfamily Genus Type-species HYLECOETINAE Boving § Craighead, Hylecoetus Latreille Cantharis dermestoides Linnaeus 1931 LYMEXYLINAE Boving § Craighead, Lymexylon Fabricius Cantharis navale Linnaeus 1931, SENSU NOVO Atractocerus Palisot A. necydaloides Palisot (= Necy- dalis brevicornis Linnaeus) MELITTOMMINAE, New Subfamily Australym exylon, new genus Hylecoetus australis Erichson Melittomma Murray Hylecoetus brasiliensis Casteln. Melittommopsis Lane M. juquiensis Lane Melanomelittomma, new genus Melittomma insulare Fairmaire Genus HYLECOETUS INTRODUCTION Species of the genus H ylecoetu s have been a focal point of interest among European naturalists since before Linneaus published h is Systems Naturae marking th e advent o f modern zoological nomen clature in 1758. H ylecoetu s beetles are a distinct group, marked by numerous autapomorphies, and readily recognized by characters presented in the key and in the generic diagnosis below. Intraspecific variation is great in the common Eurasian species H. dermestoides. Several distinct color and structural morphs have historically been recognized; a practice which is continued in Europe (e.g., Dominik, 1975). In my discussion on H. dermestoides below, I include a treatment of these morphs. I do not do this to promote their nomenclatural usage, which I believe should be abandoned. My intent is to provide a continuum with the historic literature until the geographic distributions and biologies of these forms can be more fully evaluated. Thus, our chances of segregating various observations into their appropriate morph are best. Additional comments on some specimens of H ylecoetus are included in Appendix A, and are referred to below as appropriate. New syno nyms are proposed below. I recognize only two species of H ylecoetus as v a lid : H. d erm esto id es (EURASIA) '& H. lu g u b ris (NORTH AMERICA). 143. Fig. 73. Hylecoetus dermestoides, habitus, dorsal view. Drawing by J . Nagy. 144. 145. : ii m s ' t ’si,1; '< iirtalsfck 73 TAXONOMIC CONSIDERATIONS GENERIC DIAGNOSIS - Males with maxillary palp-organ (Fig. 7); females lacking maxillary palp-organ (Fig. 6); head with epicranial pit (Fig. 2); soft-bodied (lightly sclerotized) and dorso-ventrally flatenned (Figs. 1, 73); elytra slightly shortenned, "ribbed" with costae, exposing apical 1-3 abdominal terga (Figs. 36, 73); scu tellu m with single, medial "keel" (Fig. 37); tibial spur formula 0-1-1 (males + females); eyes small, round, entire; head gradually narrowed behind eyes, without "neck" (Fig. 2); procoxae not slotted at apex; hypo- meron of prothorax not slotted; lateral lobes of aedeagus (male geni talia) asymmetrically placed and freely articulated (Figs. 57, 58, 59); metendosternite with large furcal arms, large laminal plates, and with anterior tendons proximal (Figs. 47, 48, 49); LARVAL CHARACTERS - Tarsungulus pleurisetose (Fig. 62); abdominal segment nine extremely long, heavily sclerotized, "sword-like" in shape, and bifurcate at apex (Fig. 61); segments of legs short, densely setose (Fig. 63); GEOGRAPHIC DISTRIBUTION - Europe, U.S.S.R., China, Japan, eastern North America; FUNGAL HOST - Ascoidea hylecoeti (Ascomycetes); ARBOREAL HOSTS - Many hardwoods, and most softwoods in Europe and North America, especially Pinus spp., Liriodendron sp., and others. 146. DESCRIPTION Hylecoetus dermestoides, the type-species of this genus, was selected for the introduction to lymexylid morphology, and was described in great detail. Both imago and immature stages were referr ed to in that section on external morphology, and no further de scription is required here. It should be noted that H. dermestoides and H. lugubris are very similar to one another, and thus diagnoses for these two species (in addition to the morphological discussion) are sufficient. 147. 148. KEY TO SPECIES OF HYLECOETUS Maxillary palpi with large, flabellate organ (Fig. 7) OR with minute maxillary palp-organ AND antennae flabellate (Fig. 1); color entirely black OR testaceous with head, pronotum, scutellum, and elytral apices black ...... ...... (MALES)...... 2 Maxillary palpi lacking organ (Fig. 6); color entirely testaceous to reddish-yellow; black with reddish head, pronotum, and scutellum OR testaceous with elytral apices black and/or head black ...... (FEMALES)...... 3 Apex of median lobe of male genitalia bulbous (Fig. ); color entirely black; antennae always filiform (Fig. 2, 10) and maxillary palp-organ large and fully-developed (Fig. 2, 7); NORTH AMERICA...... lu g u b ris Apex of median lobe of male genitalia narrow (Fig. ); antennae filiform and maxillary palp-organ large OR antennae flabellate (Fig. 1) and maxillary palp-organ small; color entirely black OR testaceous with head, pronotum, scutellum and elytral apices black; EURASIA ...... dermestoides Elytra black, scutellum, pronotum, and head reddish color; NORTH AMERICA...... lu g u b ris Color entirely testaceous to reddish OR testaceous with head AND/OR apices of elytra black to dark reddish-brown; EURASIA...... dermestoides TYPE-SPECIES AND SYNONYMIES H ylecoetu s L a tr e ille Type-species: Cantharis dermestoides Linneaus H ylecoetus Latreille, 1806: 266; Jaquelin du Val, 1863: 206 (subgenus) Elateroides S ch aeffer, 1766: 139 H ylecerus Jaquelin du Val, 1863: 206 (subgenus) H yloceotus M elsheimer, 1853: 86 (in e rro r) H yloecotus L acordaire, 1857: 500, 502 (in e rro r) H yloecotus Chenu, 1860: 223 (in e rro r) Xylecoethus G yllenhal, 1827: 234 THE SPECIES. - Both the North American and the Eurasian species of this genus are discussed individually below, including a treatment of the intraspecific variation of H. dermestoides. Table 8. List of' the species of Hylecoetus, with new synonymies. Species Author, date, year/ status Geographic distribution H. dermestoides Linnaeus, 1761, p . 201 / VALID SPECIES Eurasia: Europe, USSR, China, Japan H. cribricolle F airm aire, 1889, p .45/ NEW COMBINATION China H. flabellicornis Schneider, 1791, p.109/ NEW SYNONYMY Europe H. cossis Lewis, 1896, p.65/ NEW SYNONYMY Japan H. matsushitai Kono, 1938, p . 199/ NEW SYNONYMY Japan H. orientalis Foursow, 1935, p.92/ INCERTAE SEDIS (Proba East Siberia b le synonym) H. lugubris Say, 1835, p.166/ VALID SPECIES North America: USA, Canada Hylecoetus dermestoides Figs. 1-67, 69-77, 80 Cantharis dermestoides Linneaus, 1761: 201. Meloe marci Linneaus, 1766: 681. Lymexylon proboscideus Fabricius, 1781: 256. Mordella barbatus Schaller, 1783: 322. Lytta francofurthanus Herbst, 1784: 145. Lymexylon morio Fabricius, 1787: 165. Lymexylon flabellicornis Schneider, 1791: 109, NEW SYNONYMY Hylecoetus dermestoides (Linneaus), Latreille, 1806: 266. Hylecoetus aestivalis Gistel, 1857: 14. Hylecoetus nycturgus Gistel, 1857: 25. Hylecoetus cossis Lewis, 1896: 65, NEW SYNONYMY Hylecoetus julvipennis Reitter, 1911: 299. Hylecoetus matsushitai Kono, 1938: 199, NEW SYNONYMY DIAGNOSIS.- H. dermestoides can be separated from H. lugubris by the characters in the key to species. 152. DESCRIPTION- H. dermestoides was described in detail above under "Morphology ". NATURAL HISTORY- See section above "Natural History". LARVAE- See larval description under "Morphology". INTRASPECIFIC VARIATION- As I mentioned above, dermestoides has several forms or morphs.which have been recognized for many years in Europe. Because of the large "gaps" between some of these morphs, and because data on their distributions and habits are both sparse and poorly analyzed, I think it is wise to continue to recognize these morphs until said studies are complete. This will also provide a clear connection with the existing body of literature in which refer ences to these forms is made. A simple way of dealing with these several morphs is to present a dichotomous key. I want to emphasize that these names should not be perpetuated as subspecies or any other nomenclatural unit; I am discussing them to assist in the future collection of additional data on distribution and hosts, so that the relationships among these forms can be understood. A perusal of the list of synonyms above and the morphs below will reveal that most of them have had nomenclatural status in the past, including some which are newly synonymized here. A few remarks on some of the morphs are included in Appendix A. 153. KEY TO MORPHS OF HYLECOETUS DERMESTOIDES Antennae slightly serrate, nearly filiform; maxillary palpus simple, without palp-organ (Fig- 6) ------(Fem ales)------2 Antennae nearly filiform AND maxillary palp-organ present, large, fully developed (Fig. 7); OR, antennae flabellate (Fig. 1) and palp-organ greatly reduced, minute ------(M ales)------3 Color entirely testaceous to reddish-yellow ------ ------dermestoides MORPH Color testaceous to reddish-yellow with head AND/OR apices of elytra balck or very dark reddish-brown ------c o ssis MORPH Antennae flabellate; maxillary palp-organ reduced, very small ------ ------flabellicom is MORPH Antennae nearly filiform ; maxillary palp-organ fully developed— 4 Color entirely black ------morio MORPH Color testaceous; head, pronotum, scutellum, and apices of elytra b la c k ------dermestoides MORPH Female Morphs 1. dermestoides-morph. One of the first clues which indicated to me that the long-standing species dermestoides and flabellicornis might actually be conspecific was that the females were inseparable based on color, morphology, and even genitalia. This typical (and by far the most common) "dermestoides" morph is uniformly testaceous to reddish- brown. 2. cossis-morph. Lewis (1896) described as a new species female individuals which had the head and apices of elytra black. I synonymize this species with dermestoides here (see Appendix A: Item 16). This morph has been described on several other occassions. Kono (1938) described a cossis-morph with typical (pale) head color and black elytral apices, which I also synonymize here. Kono mistakenly took the female to be a male without palp-organs (see Appendix A: Item 8). Foursow's description of orientalis (from Hast Siberia) is very likely another synonym based on the cossis-morph. I have not been able to examine type material for the Foursow species, however, and do not propose synonymy at this time. Also, I transfer Fairmaire's species, cribricolle, to Hylecoetus from Melittomma based on the description (Fairmaire, 1889: 45). I have not seen the type for cribricolle but strongly suspect that it too is synonymous with dermestoides. As I note in the appendix, various combinations of the dark/pale head and 155. elytral apices occur. Further, I have seen "cossis" morphs and typical morphs from th e same lo c a lity . Male Morphs 1. flabellicom is-morph. (Tigs- 1> 22-26; Appendix A, Item 12). This "species", first named by Udmann in 1753, and later validated by Schneider in 1791, has been known to European coleopterists for more than two hundred years and was therefore a source for relunctance on my part to suggest synonymy. This concept, however, is not without precedent. Pfeil (1859) and Germer (1912) each pointed out that "flabellicornis" was perhaps only a variation of the male of dermestoides. I believe that this interpre tation is correct. The two forms occur sympatrically (their respective ranges have not been fully analyzed, but I have seen specimens from the same localities in Europe), at least in part, and morphological they are identical except for the form of the antennae and palp-organ (even the aedeagus is identical). The chemoreceptors of the flabellate antennae are not the same as those on the maxillary palp-organ, as discussed in the section on cladistics below. It is interesting that "flabellicornis" also (but rarely) occurs with complete black coloration This, of course, parallels the "morio" morph, long recognized as a form of dermestoides. This bizarre modification of the antennae is truely remarkable, and 156. I can think of no similarly impressive intraspecific variation in any other lymexylid. In a forthcoming revision of the genus Hylecoetus, I plan detailed analyses of the distribution of each morph, including "flabellicornis" in hopes of developing some explanation for its occurance. In the meanwhile, I am confident (as a morphologist) that this form is conspecific with dermestoides. (See also, Appendix A: 18). 2. morio-morph. This long-recognized variation of dermestoides was described by Fabricius in 1787, and is easily distinguished by its color which is entirely black (Fig. 77); this is in sharp contrast with "typical" forms of the species (Fig. 74). As I noted above, the flabellicornis morph also occurs in an entirely black color on occassion. 3. dermestoides-morph. The "dermestoides" morph is by far the most common and widespread of all, and is marked by nearly filiform antennae and large maxillary palp-organs (Pig* 7); the color is testaceous with the head, scutellum, pronotum, and elytral apices black (Fig. 74). Hylecoetus lugubris Say Hylecoetus lugubris Say, 1835: 166 (tyPe presumed lost) Hylecoetus americanus Harris, 1883: 86; Leng, 1920: 152 Diagnosis. See characters in key and figures 78, 79, and 81. Discussion. Aside from color differences, and a slight difference in the shape of the apex of the median lobe of the male genitalia lugu bris is very similar to dermestoides in both adult and larval structure. But the resemblances are carried even further since both species seem to have the same ambrosia fungus as host (Batra 8 Francke-Grosmann,i961). More detailed ecological studies are badly needed for both species, but especially lugubris. Table 19. L ist o f species transferred from Hylecoetus Species Author/date/page New Combination Hylecoetus australis F richson, 1842, p . 147 AUSTRALYMEXYLON, NEW GENUS (Lymexylon) (as type-species) Hylecoetus fuscipennis Lea, 1894: 603 AUSTRALYMEXYLON, NEW GENUS Hylecoetus pervagus Olliff, 1889, p.87 Melittomma Murray Hylecoetus vigilans Lea, 1911, p. 467 Melittomma Murray Hylecoetus linearis Lea, 1894: 603 AUSTRALYMEXYLON, NEW GENUS (as new synonym of A. australe, New Combination) tn oo Figs. 74-79. Hylecoetus spp., habitus drawings showing color patterns of species and morphs (dorsal view). Drawings are semi-diagramatic: fig. 74, H. dermestoides, ’’dermestoides" morph, male; fig. 75, H. dermestoides, "dermestoides" morph, female; fig. 76, H. dermestoides, "cossis" morph, female; fig. 77, H. dermestoides, "morio" morph, male; fig. 78, H. lugubris, male; fig. 79, H. lugubris, female. 160. 75 76 mmIV.*-.••.;:• 77 79 Figs. 80-81. Hylecoetus spp., apex of median lobe: fig. 80, dermestoi- des; fig. 81, lugubris. 161. end Genus lym exylon INTRODUCTION Lymexylon navale is the true "ship-timber beetle ", a common name which is often applied to the entire family Lymexylidae (e.g., Arnett, 1971). This species was a serious pest during the 16th century, as the common name implies, to the shipping industry. The species has the distinction of having been described by Linneaus in his Systema naturae (1758, p .403) . Westwood (1839, p . 274) r e la te s an in te re s tin g anecdote concerning Linneaus and this species: "Linneaus, at the suggestion of the King of Sweden, having investigated and ascertained the real cause of the damage, suggested that the timber should be sunk under water at the time the insect made its appearance in the perfect state, whereby it was secured against attacks". I retain three species in Lymexylon, two of which are of doubtful status as discussed below. These include navale (Europe), ruflcolle (Japan), and oculare (Japan) . Other species included in this genus by Schenkling (1915), include a d ela id a e (transferred to my new genus Australymexylon as a junior synonym of australe), and sericeum ( tr a n s fe rre d to Melittomma by Leng, 1920). Some notes on the type of L. adelaidae are included in Appendix A (Item 3); as are comments on L. ruficolle (Item 15) . L. australis is proposed as the type species fo r Australymexylon, new genus (see Appendix A, Item 11). 163. Fig. 82. Lymexylon navale, habitus of male, dorsal view. Drawing by J . Nagy. 164. 166. TAXONOMIC CONSIDERATIONS L y m e x y l o n Lymexylon F a b ric iu s, 1775, pp. 3, 204 Calolymus Thunberg, 1791, p. 26 Cantharis (in part) Linneaus, 1758, p.403 Pterophorus H erbst, 1784, p. 47 GENERIC DIAGNOSIS. - Body form elongate, narrow, parallel-sided, dorso-ventrally flattened (Fig. 82); head abruptly narrowed behind eyes forming a distinct "neck" region (Figs. 83, 84); eyes large, emarginate anteriorly (Fig. 83); metendosternite with anterior tendons arising near middle of lamina, but distinctly separated (Figs. 94, 95, 96); male maxillary palp-organ large, with primary-tertiary branching pattern and with very long, dense, "thin-walled" sensilla on inner surface of each branch (Figs. 97, 87); median lobe of aedeagus elongate, curved, and narrowed at apex, about as long as distance from base of tergminal struts to apex of lateral lobes (Figs. 98, 101, 102); lateral lobes very short and wide, with distinct lateral margins and angles, and separated from each other by distinct medial gap (Fig. 99); male abdominal segment VIII simple, telesoped into VII (Figs. 98, 100); male IX with ventral strut; female with long ovipositor with short, clavate, densely setose styli and with basal part of coxite broad and shorter than the distal part (Figs. 104-110); hindwing with nearly complete 167. venation; la-2a crossvein incomplete; spurious "R" absent (Fig.91); elytra shortened, exposing one or two complete terga and sometimes part of a third; hypomeron short and narrowed, ending at about metacoxae and apical part of elytra narrowed, lying flat over the abdomen and lacking a tongue-and-groove mechanism to lock them together (Fig. 82); LARVAL CHARACTERS - abdominal segment IX very large, membranous, bulbous and postero-dorsally directed (Fig. 117); trochanter of each leg enlarged, lobate (Fig. 116); tarsungulus bisetose (Fig. 115). DESCRIPTION Body form elongate, parallel-sided and dorso-ven- trally flattened (Fig. 82); pronotum narrower at base than combined width of elytral bases (Figs. 82, 89, 90); total length about 4.8-16.5 mm (males usually do not exceed about 13mm, while females share the lower end of the size range). C olor. Males and females differ in color. Male: Head black; pronotum dark reddish-brown to nearly black with more or less defined nvM-shaped transverse band in front of middle (band of lighter reddish color); elytra yellowish-brown at base including humeral angles, and 1 This description is based on the type-species Lymexylon navale. It is suggested below that the other species in the genus are likely to be synonymous with n avale except for some minor color differences. Therefore, it is assumed that this description is applicable for the genus as a unit as well. See figures 82-116, 134, 222. 168. this pale area narrowing along suture, extending about 7/10 length of elytron; humerai angles, lateral aspects, and apex of elytra dark reddish-brown to nearly black; abdominal terga dark reddish-brown, except tergum VII (the apical, visible tergum) yellowish; antennae, maxillary palpus (and maxillary palp-organ) and mandibles dark, other mouthparts and legs yellowish; thoracic sterna dark, abdominal sterna yellowish. Female: As in male, except antennae less dark and all mouthparts, except apices of mandibles, lighter in color; pronotum paler, reddish-yellow and lacking the "v"-shaped marking of the male; elytra predominantly yellowish, dark color confined to lateral area beginning behind humeral angle, and gradually widening until entire apical area is dark; the only dark ventral parts are the metastemum, metepisternum, and metepimeron. Surface Structures. The body is soft (i-e*» lightly sclerotized). The elytra are sometimes convoluted, with irregularly distributed "bumps" on the surface. The head is darkly colored, and more heavily sclerotized than other parts of the body. The head is covered by very dense, deep, coarse punctures which are sometimes coalesced on the "neck" region, producing a rugose pattern. The pronotum is covered by dense, small, shallow punctures. The punctures of the elytra are similar to those of the pronotum, but less defined. Punctures of the abdominal sterna are dense, very shallow, and poorly defined. The punctures are a little more coarse than on the abdomen. 169. The Head: General D e sc rip tio n . The head is small relative to the body (Fig* 82).; Broad in form (Tigs. 82, 83, 84), widest at the eyes; narrowed behind the eyes forming a distinct "neck" region; with a prominant supra-antennal ridge. Gular region and ventral surface. The gula is incomplete; the gular sutures are short, nearly parallel (slightly convergent anterior ly) ; the small gula between them is shorter than the area anterior to i t (the pregu la of Doyen, 1966, p.106, fig. 3). Dense, long, thin microtrichiae border the ventral edge of the occipital foramen C=foramen magnum) . The area surrounding the eyes ventrally is wrinkled in appearance, otherwise the ventral surface is nearly smooth (there are, however, exceedingly small transverse lines). There are dense setae posterior to the eyes (and on the dorsal surface of the head), but most of the ventral surface lacks these setae. Eyes. The eyes are large, prominant, and constitute the widest point on the head; they are widely separated both dorsally and ventxally and each eye is longest toward the midline dorsally and shortened toward the midline ventrally (Figs. 83, 84); the facets are fine, 5- or 6-sided in shape, and have long, erect setae between individual f a c e ts . Labrum and epiph aryn geal re g io n . The labrum is very small, and transverse. The dorsal surface is covered by very dense, long, stout setae. The epipharyngeal region is about twice the length of the labrum. Near the base of the epipharynx are transverse rows of 170. microtrichiae; the remaining surface is covered by dense, irregularly distributed microtrichiae. M a n d i b l e s . The mandibles are short and broad. Along the mesal edge, near the middle, is a large prostheca bearing dense setae (Fig. 85); the outer margin is covered by sparse, long setae. Some trans verse rows of microtrichiae, oriented perpendicular to the longitudinal axis of the mandible, may represent remnants of the mola. M axilla and m axillary palp-organ. The male and female are discussed separately below. F e m a l e : maxilla of generalized form; lacinia long, narrow, and parallel-sided with many long setae and a patch of long microtrichiae basally; galea short, broad, rounded at apex, covered by moderately dense, long setae, and divided by a broad membrane into a distinct basigalea and distigalea; ventral surface of stipes covered by dense, long microtrichiae, and continuous basally with a membrane which is covered by identical microtrichiae; cardo elongate; palpiger large, lateral; palpus 4-segmented, covered by very long, stout setae; apex of segment IV wide and bearing dense, elongate, blunt sensilla; I is short, little longer than wide; II much larger, clavate; III short, transverse; IV very large, broad,.expanded apically. M a l e : Galea and lacinia distinct, relatively small. Palpus "4- segmented": I short; II abruptly larger in size, widened apically; III small, ringlike; IV relatively short, with dense area of sensilla (Fig. ); maxillary palpus-organ arising from III with primary-secondary 171. branches, and only a few tertiary branches (Fig. ); surface of each branch covered by sparse, short, bristle-like setae on outer surface, and very long, dense setae on inner surface (Fig. ). Labium and hypopharyngeal region. The labium is sm all, and lightly sclerotized. The palpigers are distinctly separated from one another, and very lightly sclerotized medially (an elongate area later ally is heavily sclerotized). The palpus is 3-segmented: I is very short and ringlike with few setae; II is elongate, parallel-sided, bearing several long setae; III elongate (about as long as II), with long sensilla apically. The ligula is very membranous, and covered by sparse microtrichiae and bordered apically by sparse, long setae. Antenna. The antennae are similar in males and female, and are long, narrow, and nearly filiform in structure. Antennomere I is elon gate, constricted near the base; II is short, broad (the bases of both I § II have several very short, erect microtrichiae); III-X are elongate and slightly expanded apically; XI is wider basally and narrowed in a p ic a l h. Prothoracic structures. The pronotum is broad (about 80% of length at midline); the anterior margin broadly rounded; the posterior margin sinuate and the posterior angles sharply produced. The prostem- um is a simple, quadrate sclerite less than % length of pronotum. There is no lateral margin on the notum. The tergo-stemal suture is simple and there is no coxal slit present, and there is no intercoxal process of the prostemum. The hypomeron is not separated from the 172. pronotum, except near the postero-lateral corners which protrude in alm ost e l e t e r i d -like fashion. The procoxae are long and cylindrical, and are not slotted apically. Mesothoracic structures. The mesostemum is small and subtriangu- lar in shape. The mesepisternum is nearly equal-sized to the meso- sternum and is simple (lacking an elevated area set of by a suture as in Melittomma spp.) . The mesepimeron is a rectangular plate, smaller than the mesepisternum and digitiform and posteriorly directed. The mesocoxae are similar to the procoxae (cylindrical), but shorter. Metathoracic structures. The metasternum'is an elongate, nearly quadrangular sclerite with depressions anteriorly which receive the mesocoxae. The metacoxae are obliquely positioned, large, and have deep, transverse grooves which receive the femora. The metepistemum is narrowed posteriorly and separated from the metepimeron by a very deep, distinct suture. Legs. The legs are long and thin in typical lymexylid fashion. The tibial spur formula is 2-2-2 in both sexes. Metendosternite (Figs. 94-96). The lateral parts of the furcal stalk are fused dorsally for the distal half of their length. The heavily sclerotized portion of the stalk is truncate, widening into a very lightly sclerotized plate which has small, triangular, protruding angles basally and which gives rise to the furcal arms. The furcal arms are long and curved. The anterior tendons are further apart basally than in H glecoetu s and arise from the anterior tendon stalks 173. (ats in my figure)_ of lamina. No lateral process is present on the furcal stalk. S cu tellu m . The exposed part of the scutellum (of mesothorax) is a simple lobe, rounded apically; the unexposed part has small, pointed comers posteriorly (lateral and basal to exposed surface). There are no keels and the surface has punctures and setae as on elytra. Elytra. The elytra are long and narrow. The medial margins do not have the usual tongue-and-groove locking mechanism of most genera. The epipleural fold fits into a concave area created at the suture between the metepistemum and metepimeron; the epipleural fold is widest basally and is gradually narrowed until a paint about parallel to the metacoxae, the remaining part of the elytron not folding over the side of the body but lying flat over the back (the apices of the elytra are separated). Several terga are exposed since the elytra are shortened: in females, from 1^-3 tergites are exposed and in males from h - '2 h are exposed. Hindwings (Fig. 91). The hindwings are rather primitive and the venation correspondingly complete. The la-2a crossvein, however, is incomplete between 1A2 and the wedge-cell. Also, the venation is sexually dimorphic: males have an extra lal-la2 crossvein, forming a minute cell, located distally from the wedge-cell. The brachial-cell is feebly defined by a poorly developed vein. M is spurious. 174. Abdominal structure. The sterna are considerably wider than the terga and curve around the side of the abdomen. From a dorsal view, these dorsally-protruding sterna resemble paratergites (e.g., of aleocharine staphylinids, etc.: see, for example, Seevers, 1978). The spiracles lie in the connecting membrane between the terga and sterna, and when the latter sclerites are compressed together dorso-ventrally the spiracles are sealed off. Males and females have the same number of abdominal segments. Sterna III-VII are exposed as is usual. Terga II-VII are well developed. There are evident remnants of sternum II and tergum I and distinct pleural sclerites of segment II lying in the membrane. Male genitalia and associated structures. Abdominal segment VIII is simple in form, consisting of a small tergum and sternum, and is telescoped into VII at rest (Fig* 100). Segment IX forms a small capsule around the aedeagus and consists of a small dorsal, tergal plate, and a sternal plate which has a pair of ventral struts which are fused apically. The median lobe is broad basally, and gradually narrowed apically; its length is about equal to that of the tegminal struts and lateral lobes combined; it is heavily sclerotized dorsally and very lightly sclerotized ventrally. The tegminal plate is large, semi-membranous and without clear borders. The tegminal struts are wide, and in some places indistinguishably fused with the tegminal plate. The lateral lobes are short and bery broad and truncate in form, bearing numerous 175. long setae. The lateral lobes are separated By a wide space medially. The tegminal struts are fused on the ventral side of the median lobe near its base (the median lobe freely slides through this "loop”) (see Figs. 99, 101, 102, 103). Female genitalia and associated structures (F igs. 104-110). Abdomi nal segment VIII is simple in form and semi-membranous, and gives rise to the typical, long spiculum gastrale which lies ventrally. The distal part of the coxite is distinctly longer than the basal part, and the coxital fold is strongly impressed ventrally and slightly visible dorsally (Figs. 104, 105, 106, 108); the apical portion of the distal part is covered by dense setate, and remaining coxite surfaces are covered by sparse setae only. The styli are relatively short and clavate in form, with numerous setae including one exceptionally long setae apically (Fig. 107). The valviferal baculus includes a short, "spurious baculus" arising from a sclerite about h distance from base of valviferal baculus to its apex (Fig. 106). The proctigeral baculi and valviferal baculi are separate for their entire lengths. The median sclerite is rather broad and very long in form (Fig. 106). 176. The Larva: gen eral d e s c r ip tio n . The larva is very long and narrow in form. The pronotum is large and hoodlike. The legs are small, relative to the body size. The color is creamy yellowish to light brown. Abdominal segment IX is in the form of a large, membran ous lobe which is "bustle-like" in form. All thoracic terga and the apex of IX are tuberculate. There is a large, conspicuous epipleural fold on each abdominal segment, and each segment overlaps the segment posterior to it ventrally. The head is broadly oval. Antenna. The segment is very small and 3-segmented: I is short, transverse, ring-like; II stout, longer than I, little longer than wide in shape and with one large, pointed, sensilium, a thin, pointed seta mesally, and two smaller setae; III elongate and narrow bearing one large pointed setae, a smaller pointed seta and about three minute setae; in addition, there are two long, blunt, thin-walled sensilla. Labrum and epipharyngeal region (Fig. 114). The labrum i s a small, simple flap. There are three pairs of large, lateral setae and a dense patch of setae apically (most of which arise on the ventral surface). The epipharyngeal membrane is covered by dense, short microtrichiae and there is a distinct epipharyngeal comb. Table 10, List of described species of Lymexylon, Species Author, date, year Geographic distribution L, navale Linnaeus, 1758, p,4Q3 Europe (Cantharis) L, ru fic o lle * Kurosawa, 1949, p,27 Japan L, oculare* Nalcane, 1963, p .7 Japan L. p ersicu s* Foursow, 1935, p,92 Caucasus *These species are believed to be synonymous with L, navale. Because type-specimens were not availa ble for study synonymy is not proposed here, See text for further discussion. Figs. 83-84. Lymexylon navale, head: fig. 83, dorsal view; fig. 84, ventral view. 5mm NECK Fig. 85. Lymexylon navale, mandible of adult (dorsal). 180. 181. Q2m m Figs. 86-87. Lymexylon navale, male, maxilla: fig. 86, apex of maxillary palpus (palpomere IV); fig. 87, apex of secondary branch of maxillary palp-organ. uuuu r L 8 / V = 3 F 5 vY.i y\ ■£8I Fig. 88. Lymexylon navale, male, apex of labial palpus. 184. LULU g o 00 00 185 186. cT 8 9 Figs. 89-90. Lymexylon navale, pronotum, dorsal view: fig. 89, male: fig. 90, female. Figs. 91-93. Lymexylon navale: fig. 91, hindwing, dashed lines indi cate vein present in male only (illustrated wing is of female); fig. 92, abdomen, basal segments (lateral); fig. 93, abdomen, apical seg ments (lateral). 92 /■ 93 F igs. 94-96. Lymexylon navale. raetendosternite: f ig . 94 dorsal view* trail Ahh*6 °f.furcal i^ entra1); fig. 96, lamiAal region ve^ ’ III' Abbreviations: FA- furcal arm; ATS- anterior tendon "stalk"* SK- furcal stalk; AT- anterior tendon; L- lamina. enaon stallc ’ 189. FA AT ATS FA 96 190 Fig. 97. Lymexylon n a v a l male, maxilla,. 192 16 ------anjgtf Fig. 98. Lymexylon navale, male, apex of abdomen (lateral). 193. S-vii (SP) 9 8 Figs. 99-102. Lymexylon navale, male postabdomen: fig. 99, lateral lobes (ventral); fig. 100, abdominal stem ite VIII (ventral); fig. 101 aedeagus (lateral); fig. 102, aedeagus (dorsal). 196. Q25n*n Q2mm ss -0 2 mm 197. F ig. 103. Lyiaexylon navale, apex of median lobe and endophallus. lateral view. Figs. 104-108. Lymexylon navale, female genitalia: fig. 104, dorsal view; fig. 105, coxital fold (dorsal); fig. 106, ventral view; fig. 107, stylus; fig. 108, coxital fold (ventral). 104 106 Figs. 109-110. Lymexylon navale, female genitalia: fig. 109, apices of valvifers and bases of coxites (dorsal); fig. 110, median sclerite (dorsal). Abbreviations: cx- coxite; ms- median sclerite; pb- procti- geral baculus; v- valvifer. 200. 110 .1mm Figs. 111-113. Lymexylon navale, larva, posterior end of abdomen: fig. Ill, segments VI-X; fig. 112, segments VIII-X; fig. 113, apex of segment IX. All views lateral; not drawn to scale. 202. VII IX VIII 111 Fig. 114. Lymexylon navale, larva, labrum (v en tra l). 204. 205. Fig. 115. Lymexylon navale, mesothoracic leg of larva: fig. 115.1, entire leg; fig. 115.2, apex of tibia and tarsungulus. 115.1 207. Fig. 116. Lymexylon navale, larva: fig. 116.1, thorax and base of abdomen (lateral); fig. 116.2, maxillary palpus; fig. 116.3, mesal surface of mala (of maxilla); fig. 116.4, antenna; fig. 116.5, labium (ventral). Abbreviations: T1-T3 (thoracic segments I-III); A1-A3 (abdominal segments I-III); EPF (epipleural fold); mt (mentum); prm (prementum); lg (ligula). 208. 209. Genus Atractocerus INTRODUCTION Few genera of Coleoptera have ever been as intrigueing to the naturalists studying them as Atractocerus. The b ra c h e ly tro u s, m ostly tropical, bizarre-appearing species belonging to this genus have been a theme for argument and awe for nearly two centuries. As recently as 1955, King advocated making a new and separate suborder of Coleoptera to accomodate these strange beetles, which he believed were the sister group of Adephaga and Polyphaga combined. This suggestion was quickly countered by Forbes (1956) and Selander (1959). Arnett (1971) removes Atractocerus from Lymexylidae, and places it in the Telegeusidae, but Crowson (1955) has demonstrated that it belongs with the lymexylids, a stand which I support wholely. The natural history of Atractocerus is not well known, and an endless series of guesses as to the significance of their strange s tr u c tu r a l co n fig u ra tio n rages on. Gorham (1881), fo r example, said that "From their habit of flying to light, and their largely developed eyes, I think it probable that the Atractoceri are parasitic upon the Lampyridae in some of their stages". Wasmann (1902) suggested that some species may be inhabitants of termite nests. It has been suggested that they might mimic wasps; part of this argument centering on the extremely rapid movement of the hindwings in flight which actually 2 1 0 . Fig. 117, Atractocerus brevicornis, habitus, female, dorsal view. Drawing by J. Nagy. smsa 213. an audible buzz not unlike some hymenopterans. Perhaps one of the most interesting possibilities was discussed by Clark (1926), who observed an Australian species to emerge from their galleries and immediately fly at a rapid rate up to the canopy of the forest. The extremely long front legs of Atractocerus tasm aniensis are similar to the forelegs of arboreal cerambycid beetles, and arboreal habits could certainly account for the relative rarity of adults. Perhaps the most thorough investigation of Atractocerus biology to d a te was th a t undertaken by Simmonds (1956) o f A. b ra silie n sis in Trinidad. His conclusions are discussed above under "Natural History", and include documentation that the adults are short-lived. TAXONOMIC CONSIDERATIONS GENERIC DIAGNOSIS- ADULT CHARACTERS. Body very long and narrow; eyes large and emarginate; head narrowed abruptly behind the eyes; elytra extremely shortened, exposing much of the abdominal tergum (Fig. 117); hindwing venation greatly reduced, wing folds only longitudinally in fan-like fashion; pronotum lacks a distinct lateral margin (Fig. 133 ); antennae very short, compact, usually narrowed gradually toward apex (Fig. 123); maxillary palp-organ present in females and males, although much larger in males (Figs. 125-129; 130-131); the procoxae are not s lo tte d , and the hypomeron o f the prothorax i s not slo tte d ; scutellum with lateral grooves which receive elytra (Fig. 135); tibial spur formula reduced: 0-0-0 in males/ 0-0-0 OR 0-1-1 in females; lateral lobes of male genitalia greatly enlarged (Figs. 145, 164-167); metendosternite greatly reduced (see Figs 176-77 , and description below); female genitalia heavily sclerotized, leathery. LARVAL CHARACTERS. Abdominal segment IX la rg e, membranous, lo b a te, with apical a sp erites (Fig. 148); labrum fused to head capsule; tarsungulus with many setae (Fig. 147). Description o f ATRACTOCERUS BREVICORNIS, type-species (Figs. 117-133, 135-140, 143-153, 172, 173, 176, 177) ADULT CHARACTERS- General Description. A. brevicornis b e e tle s are very elongate and narrow; nearly parallel-sided in form (Fig. 117). The elytra are very short (never reaching the metacoxae), and the fully-developed hindwings are only folded longitudinally over the abdomen in fan-like style. Size ranges from moderate (about 15mm) to gigantic individuals some of which exceed 60mm in body length. Color. The color varies from pale yellowish to very dark reddish-brown. In all individuals, however, there is a pale, yellowish stripe extending down the midline of the vertex of the head, the pronotum , and the scutellum (Tig* 117). Surface structure. The entire body is leathery in texture, being moderately sclerotized. Most of the surface is covered by small, poorly define punctures each bearing a short, fine, suberect seta. The dorsal surface of the head, between and posterior to the eyes is covered by transverse "wrinkles". Male G enitalia- Small, compact, broad (Fig. 145; see also, figs. 164-167). Median lobe is small, broad basally, and gradually narrowed apically. Tegminal plate is large, bilobed apically, and extends from the base of the median lobe to the apex of the medial process of the lateral lobes (see labels on Fig. 167), The lateral lobes are very large and complex: with a truncate and deeply emarginate median process, two pairs of lateral denticuli, and paired, large, cup-like lateral processes. Tegminal struts are short and broad and fused ventro- b a sa lly . Female G enitalia- Elongate, narrow; in-part membranous and in part moderately- to heavily-sclerotized. V alviferal baculus: long, subparallel, thin, heavily sclerotized; laterally expanded and fused with basal sclerotization of valvifer and distally reach the coxital b a c u li. Coxital baculus: paired, elongate, lying along the longi tudinal axis of the basal part of the coxite for most of its length, nearly lined up with the valviferal baculus, and sharply bent near its apex oriented for a short distance transversely toward the midline; a sclerotized, posterior margin on the basal part of the coxite may also be considered part of the coxital baculus. M edial baculus: p a ire d delicate baculi lying mesal to the valviferal baculi and extending distally to about h len g th o f th e c o x ita l b a c u li; th in (much th in n e r than the valviferal baculi), and lying over a membranous, convoluted, internal, elongate area. V a l v i f e r s indistinguishably sclerotized in places making them difficult to delimit; a pair of distinct, relatively 216. long plates which are fused for a short distance with the valviferal baculi and a second, more distal and shorter pair just vefore the junt- ion with with coxites. C o x ite s elongate, narrow, and narrowed at apex; divided into a short basal part and longer distal part; distal part covered by dense, long setate and the basal part with sparse, shorter setae; parts separated by coxital fold ventrally. M edian s c l e r i t e is distinct on the ventral surface between the coxites and extending from about %-point of the distal part (of coxite) slightly beyond the coxital baculi. S t y l i are very long, thin, tubular and very heavily sclerotized; with sparse setae near apex and apical part of outer-surface; with few short, peg-like sensilla near apex; apex with fold in membranous area. Proctigeral baculus paired, separate, long baculi; divergent anteriorly and posteriorly; thickened basally and very narrow apically. Variation in Genitalic Structure. - As noted elsewhere (under the section of cladistics) the male genitalia in the genus Atractocerus vary considerably in structure, some species having the genitalia borad and compact, and others with long, narrow, almost antler-like processes. All of these, however, show the same basic (and homologous) parts in their structure, and I do not think there is any doubt that they share a common origin. Desciption of other structures of Atractocerus follow. 217. The Head. The head is broadly oval (Figs. 118, 119) and strongly flattened dorso-ventrally. The eyes are large and conspicuous, and occupy much of both the dorsal and ventral surfaces of the head. The facets of the eyes are moderate-sized and appear fine under low to moderate magnifications with the dissecting microscope. The gular sutures are short and nearly parallel (slightly convergent anteriorly). The antennae are inserted under a lateral, supra-antennal ridge which is broad and conspicuous,and are little longer than the head. There is no fronto-clypeal suture, and the labrum is immovably fused to the head with no trace of a suture. The epipharynx is simple, small and reduced, consisting only of a row of small setae along the midlin of the ventral surface. The posterior foramen is large. Eyes. The eyes of both sexes are large, ovate, and cover much of the head. The vertex is widest between the posterior part of the eyes, and is narrowed anteriorly until the eyes become proximal. AT the widest point, the eye width/total head width ration is about 42/67 from a ventral view and about 50/80 from a dorsal view. The facets are more or less hexagonal in shape. Mandibles. The mandibles are small, short, and broad. The apex is bidentate. There is no evidence of a molar region or a prosthecal area. Much of the surface bears long, dense setae. Antennae. The antennae are similar in males and females. The spindle-like shape of the antennae and their shortness are distinctive. Antennomere I is elongate and clavate; II much shorter (and shorter 218. than wide); the surfaces of I and II are finely reticulate with a few short, pointed setae near the base and moderately dense, long, fine setae over much of the surface; antennomeres III-XI are covered by dense setae which are shorter than those of I-II, and the entire surface is covered by dense, minute, peg-like sensilla; antennomeres III-VIII are broader than long (V 8 VI being widest of all), IX is about as long as wide, X is slightly shorter than IX, and XI is elongate and pointed a p ic a lly . M axillae. A maxillary palp-organ occurs in both males and females (so far as I know this in only true for Atractocerus spp.). The s tr u c ture of the maxilla is discussed separately for males and females. Maxilla: females. The mala is not divided into a distinct galea and lacinia, and is a single, broad sclerite narrowed to a point apicaDy with a small, tuberculate projection near the middle. The mala is covered by dense setae (Fig. 128). The palpiger is small and ovate. The palpus has 4 palpomeres: I short; II short and very broad, strongly oblique (Fig. 125), exposing a large, membranous apex; III short, ring like, bearing the fourth palpomere and the maxillary palp-organ. The membrane connecting III to both IV and the palp-organ is wrinkled and bears setae and sensilla identical to those of the palp-organ, thus, IV appears at a glance to be simply a branch of the organ (except that its apex bears long setae and narrow sensilla characteristic of the palpus and differing from those of the palp-organ). The palp-organ has only primary-secondary branches (l°-2°). 219. M axilla: male. The male maxillary structure is similar to the female, with the following exceptions. The maxillary palp-organ is much larger and has a more complex branching pattern with primary-ter tiary branches (Fig.130). The fourth palpomere is only recognizable by its apical setae and sensillae and is further removed from III than in the female (apparently because the membranous area separating them is increased in size). The illusion that IV is part of the palp-organ is carried further because its basal surface bears the characteristic wrinkles, setae, and sensilla of the branches of the palp-organ and further it is actually sub-branched itself. The primary branches bear few setae. The sedondary branches have pointed setae on the outer surfaces (which lack visible sockets) while the innter surfaces bear long, blunt sensilla. The tertiary branches which are short, digitiform, and oriented in two parallel rows along the inner surface of the secondary branches are covered entirely by the long, blunt sensilla. Labium (Figs. 172, 173). The labium is greatly reduced. The mentum is elongate and narrowed. The palpigers are represented by a pair of large plates which when viewed laterally resemble an extra, basal palpomere. The palpi consist of two palpomeres: I is elongate, subparallel; II is very robust in form, much larger than I. The ligula is very la rg e and le a th e ry , about equal in len g th to th e mentum and is narrow and expanded apically. The hypopharynx is small, membranous and located near the base of the ligula. The surface of the mentum, 2 2 0 . ventral surface of ligula, and surface of palpi (except for outer part of palpomere II) are covered by dense, long setae. The outer surface of palpomere II bears a large patch of recumbent sensilla. The hypo pharynx is covered by very dense microtrichiae and a few long setae as on much o f th e labium . The Prothorax. The pronotum is short and nearly quadrate; sides are nearly parallel; anterior margin broadly rounded; without lateral margins. The female pronotum is a little broader than in the male, as noted by V illiers Cl969). The prosternum is very short and broad in form; the spiracle is located on a large postpleural sclerite (see Baehr, 1976); and there is a distinct, small spinasternum behind the coxae medially. The coxal cavities are broadly open behind and the procoxae strongly cylindrical and long in form. The Mesosternum and Associated Sclerites. The mesocoxae are large, cylindrical and posteriorly directed. The mesosternum is a small, nearly triangular sclerite between and just anterior to the mesocoxae. The mesepisterna are large sclerites, contiguous medially, broad near the midline of the body and narrowed laterally. The mes- epimeron is a narrow sclerite lying next ot the mesepisternum, which is oblique in position and swollen near the elytral base. The mesepisternum is rather flat, and not elevated anteriorly. The mesepimeron is not grooved. The Metasternum and Associated Sclerites. The metasternum is very long and narrow with shallow depressions anteriorly which receive 2 2 1 . the mesocoxae at rest. Both the metepisternum and metepimeron are long, narrow plates which are broad anteriorly and narrowed posterior ly- Elytra (Fig. 134) . The elytra are extremely shortened. The mesal margin has a broad lobe near the base which fits into a groove on the side of the mesoscutellum. The ventral surface under this lobe is thickened and is pressed against a digitiform process which represents a modification of the alar ridges of the metascutum which cause the elytra to flex, the distal portion being elevated. This may provide room for the hindwing to flex. Abdomen. Abdominal segments II-VII are visible in both males and females. Segment VIII is smaller and is telescoped into VII. The abdomen is very long and robust, and a little flattened dorso- ventrally. Each tergite and sternite is nearly quadrate. There is a minute, transverse carina on each sternite near the posterior angle. Each tergite has a distinct lateral fold, leaving a narrow margin in a nearly vertical orientation. The spiracles are large, oval, and located on these "pleural folds" which are more lightly sclerotized than other parts of the abdomen and covered by dense, short setae except for a small glabrous area around each spiracle. Abdomen: Male segments VIII and IX. VIII is simple, the apex of the sternite narrowed. IX has a nearly complete tergite and a pair of lateral, anteriorly-directed struts which are fused apically to each other and to a pair of ventral struts. Sternite IX is 2 2 2 . narrowed apically and divided anteriorly being prodiced into a pair of long struts which are fused anteriorly with the dorsal struts. Abdomen: Female segment VIII. Segment VIII is simple, and bears a very long, narrow spiculum gastrale. Metendosternite. The metendostenite is extremely reduced. The furcal arms are not widely divergent, and are rather long and narrow. The laminae are fused ventrally into a long, thin flange. The ventral flange of the furcal stalk is very short, lying basad to the modified laminae. I was unable to find the attachment point for the anterior tendons. LARVAL CHARACTERS General Description- The larva is very long and thin in form, and slightly dorso-ventrally flattened with distinct pleural folds on each segment of the thorax and abdomen. The pronotum is very large and hoodlike. The posterior end of the abdomen is in the form of a large, membranous, "bustle-like" lobe with dense asperites over the apical surface. The body is almost entirely cream colored. C ranium - The cranium is oval from a dorsal view, and heavily sclerotized. The epicranial suture is short and strait, extending along the midline from the posterior margin. There is a small, dark, very heavily sclerotized + "w"-shaped area on the frons. The ocelli are absent. There is a large "H"-shaped, yellow-colored pattern on the head, extending from the posterior margin to points above the 223. bases of the mandibles. Labium - The labrum is a small sclerite with dense, large setae at the apex. The epipharyngeal region has a small area of very fine, long setae and a few short sensilla apico-ventrally; a narrow medial line of similar setae; and on each side a lateral, broad, epipharyn geal comb. To each side of the apex is a "fimbrial patch" of very long fine setae. M a n d ib le - The mandible is short and broad, with a single, wide, apical dens. There is a molar area near the mesal base consisting of parallel rows of minute teeth which lie perpendicular to the longitud inal axis of the mandible. M a x illa - The maxilla is compact and moderately sclerotized; much of the surface is covered by very long, stout setae and by shorter, dense setae. The cardo is small and broad, and separated from the stipes by a distinct suture. The stipes is broad basally and widened futher apically. The stipes is not separated from the mala by a suture. The mala is partially divided into a lacinial and galeal area. The lacinial lobe, near the palpus, has an elongate patch of extemely dense setae (Tig* )• The mesal surface of the galeal lobe is flattened and has a broad, elongate membranous region which bears dense setae (the "malar comb" under the morphology section above). Labium - The mentum is broad and more or less triangular shaped. The palpifers are separate, but are not clearly defined. The palpi have two palpomeres: I is broad with several setae and IT much smaller than I with several, short, small, peg-like sensilla apically. The ligula is wide and truncate with a single pair of short, broad sensing, heavily sclerotized and is slightly emarginate apically. The hypo- pharyngeal region consists of a large, membranous area near the apex with dense setae laterally and medially; posterior to this area is a large sclerite, rather "V"-shaped, and covered in part with minute lobes and posteriorly by very dense, broad, truncate setae, Antennae- The antennae are very minute sized, with setae and sensilla as in figure , Leg- The trochanter is slightly produced as a lobe beyond the base of the femur (this is exagerated in larvae of Lymexglonl. The coxa, femur and tibia are all elongate and setose. The tarsungulus bears several setae. Abdominal apex - Abdominal segment X is small and inferior in postition to segments VIII and IX, Segment IX is apical, and is produced as a large, membranous lobe which is "bustle-like” in shape. The apex of this lobe is covered by dense, minute, short, pointed cuticular processes. T a b l e H « The described species of the genus Atractocerus. See alsodiscussion in text, and references to Atractocerus in Appendix A. Species ______Author,date ,page ______Geographic Distribution References ______a ter Kraatz, 1895:163 AFRICA: Congo, Ivory Coast V illiers,1969a, 1969b, Camaroon 1969c bicolor Strohmeyer, 1910:6 NEW GUINEA Schenkling, 1915 bifasciatus Gestro, 1874:544 NEW GUINEA Schenkling, 1915 b la ir i Gardner, 1936:181 INDIA Simmonds, 1956 brasiliensis Lepeletier & Serville, 1825:309 CENTRAL & SOUTH AMERICA: Barber,1952;Blackweld- widely distributed from er,1957; Gorham,1881; Mexico & s.U.S. to Brazil Guerin-Meneville,1844; and in West Indies Sleeper,1964;Spilman, 1971;Vaurie,1957; Vitrac ,1913 brevicornis (Linneaus),1766:643 (N ecydalis) AFRICA & MADAGASCAR: Boheman,1851 ,-Chenu , c o m m on and widespread in1860;Gardner,1935; west Africa Simmonds,1956; V illie rs 1969a,1969b,1969c see synonyms in Schenk lin g ,1915 bruijni Gestro,1874:545 PHILIPPINES, CELEBES, Schenkling,1915; PERAK, HONG KONG Simmonds, 1956 K T a b le 11.Species of Atractocerus (Cont'd .). Species ______Author,date,page Geographic Distribution References crassicornis Clark, 1931:120 AUTRALIA:North-western emarginatus Castelnau, 1836:59 INDIA; SUMATRA; SRI Fulmek,1931;Gardner, LANKA; JAVA; CELEBES; 1935,1936;Kalshoven, BURMA; SIAM; SARAWAC; 1964;Karny,1922; PHILIPPINES, e tc . Simmonds,1956 graci1icornis Schenkling, 1914:320 NORTH AMERICA: C ali Schenkling,1915 fornia kreuslerae Pascoe, 1864:46 AUSTRALIA: New South Clark,1926; Simmonds, Wales, Western Australia 1956 morio Pascoe, 1860:117 NEW GUINEA; MOLUCCAS; Schenkling,1915; PHILIPPINES; CELEBES Simmonds,1956 niger Strohmeyer, 1910:6 INDIA Simmonds,1956 procerus Schenkling,1914:319 BRAZIL; SURINAM; Simmonds,1956 PANAMA (??) quercus Gardner,1935:69 INDIA Simmonds,1956;Gardner, 1936 reversus Walker, 1858:285 INDIA;SUMATRA Bourgeois,1905,1909; Gardner,1935,1936; . 6 2 2 Karny,1922;Roonwal, 1972 T a b le I I .Species of Atractocerus (Cont'd.). S ped es Author,date,page Geographic Distribution References sieb ersi Karny,1922:12 SUMATRA tasmaniensis Lea (ms name?) AUSTRALIA:Tasmania see Clark,1931:122 termiticola Wasmann, 1902:104 BRAZIL tonkineus Pic, 1948:13 Tonkin valdivianus (Philippi),1866:113 (Fusicornis) CHILE Schenkling,1915; Simmonds,1956 victoriensis Blackburn (ms name?) AUSTRALIA: Victoria Clark,1931:122 Figs. 118-119. Atractocerus brevicornis, female, head: fig. 118 dorsal view; fig. 119, ventral view. 228. Fig. 120. Atractocerus bievicornis, male head and pronotum (d o rsa l). 230. 231. 120 Figs. 121-123. Atractocerus brevicornis, female: fig. 121, apical segment of labial palpus; fig. 122, recumbant sensilla on surface of labial palpomere III; fig. 123, antenna. 233. mm ssmm r n m mmm MiW m m 1 Figs. 124-128. Atractocerus brevicornis, female maxilla: fig. 124, left maxilla, dorsal view; fig. 125, maxillary palpus and base of maxillary palp-organ (lateral); fig. 126, base of palpomere IV showing recumbent sensilla; fig. 127, apex of palpomere IV; fig. 128, apex of secondary branch of maxillary palp-organ (2°). 234. 235. 128 126 Fig. 129. Atractocerus brevicornis, female, mala. 236. 237. 129 Figs. 130-132. Atractocerus brevicornis, male maxilla: fig. 130, palpomeres III+IV and maxillary palp-organ (note that apices of most secondary branches are omitted from drawing); fig. 131, apex of palpo mere IV; fig. 132, secondary branch of maxillary palp-organ. 238. 239 Fig. 133. Atractocerus brevicornis, prothorax (ventral view). 240. 241. cervical sclerites trochanter pleural fold -spinasternum pronotum 133 Figs. 134-135. Nota of mesothorax and metathorax: fig. 134, Lymexylon navale; fig. 135, Atractocerus brevicornis. 242. mesoprescutum MESOSCUTELLUM \ MESOSCUTELLUI l—MESOPRESCUTUM METAPRESCUTUM METASCUTUNI ALAR RIDGE GROOVE^ X ‘ALAR PROCESS 'METASCUTUM 'METASCUTELLUM 135 243 Fig. 136. Atractocerus brevicornis , right elytron (dorsal). 244. 245. Fig. 137. Atractocerus brevicom is, mesostemum, metasternum, and associated structures (ventro-lateral view). 246. 247. ..-ELYTRON COXA 2 V- hindwing METEPISTE RNUM METASTERNUM------METEPIMERON COXA 3' — FEMUR 3 .TIBIA 3 137 TROCHANTER 3 Figs. 138-140. Atractocerus brevicornis; fig. 138, male abdominal seg ment VIII (ventral); fig. 139, male abdominal segment IX (ventral); fig. 140, female abdominal segment VIII (ventral). STS § TGS- struts. 249. 'A— SPICULUM GASTRALE MEMB ST 9 140 139 \ Figs. 141-142. Melittomma brasiliense: fig. 141, labium (ventral *s); fig. 142, hypopharyngeal region (dorsal h). 250. 251. Ir’i S ' i V Fig. 143. Atractocerus brevicomis, larva, labrum (ventral). 252. 253. apical BRUSH 1 'N 'l ' i;.V V ? EPIPHARYNGEAL vA SETAE • Fig. 144. Atractocerus brevicornis, female genitalia: fig. 144.1, apex of stylus; fig. 144.2, coxital baculus and associated structures. Abbreviations: cb- coxital baculus; vb- valvifeTal baculus; sb- spurious baculus. Fig. 145. Atractocerus brevicornis, aedeagus, ventral view. Abbrevia tions: LL- lateral lobe; TS- tegrainal strut; TP- tegminal plate; ML- median lobe. ujuug. 01 256 Figs. 146-148. Atractocerus brevicornis, larva: fig. 146, head, thorax, and base of abdomen (lateral); fig. 147.1, metathoracic leg; fig. 147.2, tarsungulus; fig. 148, apex of abdomen (lateral). Abbre v ia tio n s : T1-T3 (th o ra c ic segm ents), A1-A3 § A7-A10 (abdominal seg ments), an (anus), sp (spiracle), trs (tarsungulus), tb (tibia), fm (femur), tro (trochanter), cx (coxa), epf (epipleural fold). Figs. 149-150. Atractocerus brevicornis. larva: fig. 149, antenna; fig. 150, maxilla (mesal view). 259. 260. Fig. 151. Atractocerus brevicornis, larva, maxilla. 261. 262. Fie. 152. Atractocerus Brevicornis, larva, labium: fig. 152,1, hypo- pharyngeal setae; fig. 152,2, median, elevated hypopharyngeal region (dorsal); fig. 152.3, surface of elevated area; fig. 152.4, labium, ventral view. \ mm m m k \ m ' J >^^11 f c s i » nrv 264 Fig. 153. Atractocerus brevicornis, larval head (dorsal view). 266. FRONTAL SLERITE EPICRANIAL 'EPICRANIAL SUTURE MACULA 153 Figs. 154-155. Atractocerus tasmaniensis: fig. 154, antenna (male), antennomeres 2-6; f ig . 155, male maxillary palp-organ (and palpomeres II, III, and IV). 268. 155 Fig. 156. Atractocerus brasiliensis , hindwing. Abbreviations: see table 4. 269. 270. m Fig. 157, Atractocerus procerus , hindwing. Abbreviations: see table 4. 272. 157 Figs. 158-159. Atractocerus procerus, male: fig. 158, maxillary palp- organ; fig. 159, outer surface of maxillary palp-organ secondary branch es. 273. Figs. 160-161. Atractocerus procerus, male: Qfig- 160, inner surface of maxillary palp-organ secondary branches (2 ); fig. 161, surface of antenna. Figs. 162-163. Atractocerus procerus, elytron: fig. 162 (dorsal); fig. 163, "tubercles" at base. 277. Fig. 164. Atractocerus procerus, aedeagus, ventral view. Abbrevia tions: ML- median lobe; LL- lateral lobes; TS- tegminal strut; TG- tegminal plate. 280. 164 Fig. 165. Atractocerus procerus, aedeagus, dorsal view. Abbreviations: ML- median lobe; LL- lateral lobes; TG- tegminal plate. 281. 282. Fig. 166. Atractocerus procerus, aedeagus, lateral view. Abbrevia tions: 11- lateral lobe; ml- median lobe; t - tegminal p late. 16S Fig. 167. Atractocerus sp. (ex Philippines), adeagus (ventral). 285. 286. 167 'MEDIAN LO-BE MEDIAN LO BE-#--: TEGMINAL STRUT la te r a l denticuli MEDIAN- PROCESS l a t e r a l- LATERAL p r o c e s s LOBE TEGMINAL Figs. 168-169. Atractocerus brasiliensis, male, apex of abdomen fig. 168, lateral view; fig. 169, dorsal view. 287. 288. 168 Figs. 170-171. Atractocerus tasmaniensis, metathoracic leg: fig. 170, tibia + tarsus; fig. 171, claw. 289. TIBIA 170 ^TARSUS Figs. 172-173. Atractocerus brevicornis, female, labium: fig. 172, lateral view’; fig. 173, ventral view. Representative setae show rela tive sizes and densities, but many are omitted. Abbreviations: mn- mentum; pig- palpiger, pi- palpomere I, p2- palpomere II, lg- ligula, hyp- hypopharynx. V f VI / /' P1-- 2 % PlQ- -V.‘ 172 m r& \. i. Fig. 174. Atractocerus procerus, female, maxillary palpomere IV. 293. 294. 174 175 Fig. 175. Atractocerus tasmaniensis, head of male, dorsal view. Figs. 176-177. Atractocerus brevicomis. metendostemite: fig. 176, ventral view; fig. 177, lateral view. Abbreviations: vf- ventral flange (of furcal stalk), 1- lamina, fa- furcal arm. Fig. 178. Atractocerus procerus, metendostemite. 298. 299. Genus Australym exylon, NEW GENUS INTRODUCTION I am proposing a new genus, Australymexylon , for two biological species which occur in Australia (Figs. 204, 205), and which account for four nominal species formerly assigned to H ylecoetu s and Lymexylon. Britton (1970: 579) and Lawrence (in litt.) pointed out the incongruence between the reported forms of these genera in Australia, and the north ern Hemisphere species of these two taxa. The species of Australymexylon are th e only Lymexylidae known which do not have a maxillary palp-organ in at least the males. This benchmark familial character is lacking in this genus (all other genera have these structures in males only, except Atractocerus which have these organs in both sexes). These intrigueing beetles occur in east ern Australia (including Queensland, New South Wales, Victoria, South Australia, and Tasmania), seemingly following the Great Dividing Range although geographic and ecological information is too sparse and frag mentary to allow a meaningful analysis of the parameters which affect their distribution in Australia. Perhaps the most interesting structural feature of this genus is the setal pads which are found on the male abdominal sternum VIII (see F ig s. 191- 1 9 3 ). The p o s te rio r end o f th e la rv a l abdomen (fig . 2 01} is very .similar in form to that seen in genera of the Melittomma- complex. F ig. 179. Australymexylon fuscipennis , habitus of male, dorsal view. Drawing by J. Nagy. S s H A f TYPE-SPECIES: Australymexylon australe (Erichson), new combination. DIAGNOSIS OF AUSTRALYMEXYLON. ADULT CHARACTERS. Body e lo n g a te , parallel-sided, dorso-ventrally flattened; head with eyes large, emar- ginate and abruptly constricted behind the eyes forming a distinct "neck"; maxillae of both males and females WITHOUT maxillary palp-organ; antennae slightly to strongly serrate; elytra shortened, exposing one or two terga, narrowed apically, lying flat over abdomen, with epi- pleural fold short and tongue-and-groove locking mechanism ending well before elytral apices; procoxae slotted and hypomeron of prothorax slotted; mesepisternum elevated antero-medially, concave laterally; mesepimeron with sharp carina and concave, lateral "groove” which receives elytra epipleuron; pronotum with sharp lateral margin; metepi- sternum and metepimeron forming a deep groove which receives part of elytral epipleuron; abdomen with segments III-VII visible ventrally; anterior tendons arise on furcal arms of the metendosternite; female genitalia with distal part of the coxite very much longer than the basal part, coxital baculus sometimes (in A. fuscipennis) strait and extending beyond base of the distal part of the coxite; male abdominal sternite VIII with pads of dense setae which are large and circular, or very large and oval in form, and sometimes curving around the sides and occupying much of the dorsal surface as well (in A. australe); lateral lobes of male genitalia large and cup-like, median lobe either very long or quite short. LARVAL CHARACTERS. Posterior segments similar to Melittomma and related genera: cylindrical, blunt (fig.201); tarsungulus bisetose (Fig.203), but with many small spines. 303, 304. GEOGRAPHIC DISTRIBUTION, E astern A u s tra lia , g e n e ra lly follow ing th e Great Dividing Range and including political divisions Queensland, New South Wales, Victoria, South Australia, and Tasmania (Figs. 204,205}. ARBORIAL HOSTS. Associated with various species of Eucalyptus. MYC0- LOGICAL HOSTS. Unknown. SPECIES INCLUDED: See ta b le 12. KEY TO SPECIES OF AUSTRALYMEXYLON 1 Male antennae strongly serrate (Fig, 182}, female antennae serrate (Fig. 183}; lateral lobes very large relative to median lobe CFigs. 196, 197}; abdominal segment VIII of male with large, round, sternal setose-pads (Fig, 191}; color ( u s u a l l y } with an evident, pale spot at base of elytra (Fig. 179} ------,------A , fu sc ip e n n is, new comb. Male antennae serrate (Fig. 18Q}, female antennae scarcely serrate (Fig. 181}; lateral lobes small relative to the median lobe (Figs. 198-200}; abdominal segment VII of male with very large setose-pads covering most of the sternal surface and part of the tergal surface (Figs, 192, 193}; color ( u s u a l l y } uniformly dark, reddish-brown to nearly black ------~A. a u s tr a le , new comb. Australymexylon australe (Erichson), NEW COMBINATION CFigs. 180, 181, 184-186, 189, 190, 192-195, 198-204, 206-216} Lymexylon australe Erichson, 1842: 147 Ctype examined, ZMHB} Hylecoetus australe (Erichson}, Schenkling, 1915: 5; Lea, 1911: 467 Hylecoetus linearis Lea, 1894: 603, NEW SYNONYMY (type examined, South A u stralia n Museum}; Schenkling, 1915: 8 Lymexylon adelaidae Blackburn, 1898: 34, NEW SYNONYMY (type examined, South A u stralia n Museum); Schenkling, 1915: 10 Table 12. List of the species of Australym exylon, NEW GENUS, including new synonymies. Species Author, Date, Page Geographic Distribution a u s tr a le , NEW COMBINATION (Erichson), 1842, p. 147 A u stra lia linearis, NEW SYNONYMY (Lea), 1894, p. 603 Australia adelaidae, NEW SYNONYMY (BlackburnJ, 1898, p. 34 Australia fuscipennis, NEW COMBINATION (Lea), 1911 f p, 466 A u stra lia 305 306. Diagnosis - Color uniform, reddish-brown to nearly black; male antennae moderately serrate (Fig. 180), female antennae very slightly serrate (Fig. 181); male setal-pads of abdominal segment VIII very large, oval (Figs. 192, 193); median lobe very long relative to lateral lobes (Figs. 198-200). Description of the Type-Species A. a u s tra le A relatively full description of the type-species (australe) is included, and only a few diagnostic notes are given for fuscipennis below. General Description. Body elongate, parallel-sided and dorso- ventrally flattened; body only moderately sclerotized - soft bodied; surface covered by short, fine, yellow, suberect setae. Color. Dark reddish-brown, sometimes nearly black; head and pronotum usually a little darker than elytra and ventral surface. Most of the individuals seen were paler in color; presumably teneral to some degree. Head. The head is about as long at the midline as it is wide (at the widest point, the eyes), and is strongly narrowed behind the eyes forming a distinct neck region and convex in form. Eyes large and conspicuous, narrowed toward the midline ventrally and emarginate ant eriorly; eyes widely separated dorsally and ventrally. Fronto-clypeal 307. suture absent, although there is a depressed line in its location. Gular sutures slightly convergent anteriorly and extending little furth er anteriorly than the "neck". Surface covered by dense, round, rela tively deep punctures which become transversely confluent on the neck region. Antennae slender and long, extending beyond the pronotal base when the head is deflexed; antennomere I stout, elongate; II shorter, elongate; III longer than II, slightly to moderately expanded apically; IV-X slightly produced to one side in female, more so in male where it is distinctly serrate; antennomeres I-III with moderately dense, long setae only (the base of I also has some very small, short, erect setae); XI elongate and narrowed at apex; IV-XI covered by very dense, short setae (shorter than setae on I-III). Prothoracic structures. Pronotum quadrate and dorso-ventrally flattened; about 1.25x longer than wide; anteriorly broadly rounded; posterior margin sinuate with lateral angles strongly produced poster iorly. The prosternum is short and transverse, with a small, medial, intercoxal carina; procoxal cavities broadly open behind; pronotum with sharply defined lateral margins; hypomeron with coxal slits (and tergo-stemal suture relatively short, running from the hypomeron slit to the anterior orifice); prosternum with slight depressions anterior to procoxae; procoxae elongate, cylindrical, slotted apically. Mesosternum and associated structures. Mesostenrum small, tri angular, medial. Mesgpisternum paired, each sclerite slightly larger than mesosternum; lateral part concave and antero-medial part elevated. 308. Mesepimeron is smaller, lateral sclerite divided by a carina into a medial, flat area, and a lateral part which is strongly concave and receives part of the elytra epipleuron. Metasternum and associated sclerites. The metastemum is la rg e , elongate and narrowed anteriorly; the area beneath the mesocoxae is very lightly sclerotized and depressed, and also includes a narrow, elongate incision at the midline of the highly sclerotized portion of the metastemum; there is a narrow carina just anterior to the meta coxae on the metastemum. Metepistemum simple, and broader anteriorly. Metepimeron broad at anterior end, but abruptly narrowed for its posterior 3/4 length. Both the metepistemum and metepimeron have sharp carinae, and steeply elevated portions which combine to form a deep groove which receives part of the elytra epipleuron. Scutellum (mesothoracic) . The scutellum is simple, rounded apically and flat; there are no carinae. Tibial spur formula. The tibial spur formula is 2-2-2 in both males and females. E ly tro n . There are four longitudinal costae, plus a short, scutellary one. The elytron is elongate with a distinct epipleural fold basally which fits into deep grooves in the mesepimeron and metepistemum-metepimeron. The epipleural fold narrows along the length of the first visible abdominal sternum, and there is no fold for the remaining length. The tongue-and-groove locking mechanism along the suture ends well before the apex, and the apices of the 309. elytra are narrowed, lie flat over the abdomen, and are separated. The elytra are shortened, leaving one to two terga exposed. The surface of the elytra are covered by very dense, small punctures and dense, short, fine setae. Hindwing. The hindwing is rather narrow in shape, and the venation is primitively complete. A spurious Rs vein is present (basal to the brachial cell). The r-m crossvein (below brachial cell) is indistinct ly sclerotized. Metendosternite. The anterior tendons are inserted on the furcal arm near its base. The laminae are simple, and not exceptionally large in size. The furcal stalk has the sides folded and fused for a short distance ventrally. The furcal arms are not very long. (see Figs. 185-186). The Abdomen. Males and females have the same number of visible abdominal segments. Figure 187 shows a lateral view of the abdomen, with stem ites III-VII fully developed, and tergites 11-VII similarly developed. There is a small pleural sclerite of abdominal segment II, and a small sclerite anterior to stem ite III which I believe is a remnant of II. In females, segment VIII is very lightly sclerotized and simple in form. It is small and telescopes into VII. Ventrally, it gives rise to a very long spiculum gastrale. In males, VIII is moderately large, well-developed and more-or- less heavily sclerotized but is also telescoped into VII. The stem ite 310. of VIII bears large, setose pads of undetermined function. In a u s tr a le they are elongate-shaped, and, in addition to occupying most of the ventral surface they curve around the sides and occupy much of the dorsal surface as well. The tergite VIII is abruptly narrowed along the midline, presumably to allow room for this increase in the size of these setose pads. Although I have been able to examine about a dozen specimens in detail, with regard to this character, I have no firm clues to its function. Even at high magnifications (x450), I cannot find any direct evidence of secretory ducts. The fact that it is sexually dimorphic, however, indicates to me some connection with courtship, female attraction, etc. Barry Valentine (pers. commun.), who is carrying out various studies of grooming behavior in Coleoptera, informs me that he is unaware of even a single instance of grooming functions confined to a single sex. This seemingly provides grounds for dismissal of one possibility.» Female Genitalia (Ovipositor) . The female genitalia are elongate and lightly sclerotized (and, in part, membranous). The valviferal plates are very poorly delimited, and are not drawn in figure 194. The styli are small and slightly clavate with dense setae. The coxites are exceptionally long and narrow (Fig, 195). The coxital fold separ ates a short basal part and a very long distal part (the discrepency in proportions of the basal and distal parts are far greater here than anywhere else in the lymexylids). Both the proctigeral and valviferal baculi are long, simple, and separate throughout their entire lengths. Male genitalia. (Refer to figures 198, 199, and 200). The lateral lobes are broad and concave on the medial surface, with a very dense lining in the concave area of long, fine setae. The median lobe is very long relative to the lateral lobes, and narrow. The recurved basal portion is fuse with the short tegminal struts which are indistinguishably fused with the lateral lobes. The tegminal plate is very large, poorly delimited, and lightly sclerotized. Australymexylon fuscipennis New Combination CFigs. 179, 182, 183, 187, 188, 191, 196, 197, 205) Hylecoetus fuscipennis Lea, 1911: 466 Diagnosis. Elytra with pale area at base (Fig, 179); male antennae strongly serrate (Fig. 182), female antennae moderately serrate (Fig. 183); male setal-pads of abdominal stem ite VIII large and round, but much smaller than in australe (Fig. 191)‘; median lobe small relative to lateral lobes (Figs. 196-197), tegminal struts fused into a small, ventral, "Vn-shaped sclerite. Description. The distinguishing male postabdominal structures are discussed below in detail. Other features are included in the key and d ia g n o sis. Abdominal Segment VIII: sternum. Apex slightly emarginate at middle; baso-medial area membranous; surface covered by fine setae dense near apex and at middle near base; with paired, large, circular baso-lateral setal-pads bearing very dense, long, erect setae. Abdominal Segment VIII: tergum. Bifurcate basally, very lightly sclerotized along midline, surface covered by dense, minute microtrich- iae; apex rounded at middle (very slightly lobed), with moderately dense, long setae. Abdominal Segment IX. Simple, small, lateral plates with sparse 313. setae at apex; with paired spicules which are relatively short and not fused apically. Aedeagus. The general form is short and broad. Lateral lobes elongate, wide, rounded apically with sparse setae over outer (lateral) surface, setae much denser medially (within the cup-like, concave area); lateral lobe about as long as tegminal plate, much longer and more conspicuous than the median lobe. Median lobe relatively small; bifurcate basally and rounded at apex. Tegminal plate large, very lightly sclerotized, ventral in position. Tegminal struts reduced, flattened, forming a "VH-shaped sclerite on ventral surface near base of median lobe. Discussion. The c o lo r o f fuscipennis is highly variable, due in part to various degrees of teneralness among the specimens I have seen. The following description applies to a fully-matured beetle: head dark reddish-borwn; antennomeres IV-XI very dark, nearly black (basal antennomeres paler reddish-brown); pronotum dark reddish-brown anter iorly and medially, very dark (nearly Black) laterally and posteriorly; elytron with basal l/6th pale orangish-color, darker posteriorly (to nearly black); legs dark reddish-brown; exposed tergites dark reddish- brown; ventral surface reddish-brown (variable: from rather pale to chestnut dark reddish-brown); setae on body surface golden color. Figs. 180-181. Australymexylon anstxale, antenna: fig. 180, male; fig. 181, female. 314. 315. w 'J l i f e w • v > %» \ j.Ll]? Figs. 182-183. Australymexylon fusciperm is , antenna: fig. 182, male; fig. 183, female. 316. 318. i m 184 F ig. 184. Australymexylon australe, hindwing. Abbreviations: see ta b le 4. Figs. 185-186. Australymexylon australe, metendostemite: fig. 185, ventral view; fig. 186, base of furcal arm and place of anterior tendon insertion, ventral view. 320. I a n terio r tendon insertion MLP— MLP 185 Figs. 187-188. Australymexylon fuscipennis: fig. 187, lateral view; fig. 188, male postabdomen, dorsal view. Abbreviations. S2-S7 (stem ites II-VII); T2-T7 (tergites II-VII); P2 (pleural scler- ite of abdominal segment II). 187 TERGITEVIl4~ -STRUT OF IX J—SETO SE PAD (SJERNITE VIII) t e r g h t e v m - 4 - a I 'PLEURITE IX ANUS-___ -TEGMINAL PLATE LATERAL LOBEJ-L-'- •^■T-^j-MEDIAN LOBE Figs. 189-190. Australymexylon australe: fig. 189, male abdominal segment IX (lateral); fig. 190, apex of paratergal struts (ventral). Abbreviations: ST-9- ninth stem ite; Paraterg.9- ningth paratergite. 323. 324. paraterg , ST 9 5m m 189 ■STRUT 190 Fig. 191. Australym exylon fuscipennis, male, abdominal ste m ite VIII, ventral view. 326. Fig. 192. Australymexylon australe, male, abdominal ste m ite VIII, ventral view. ] » 1 5 5 mm Fig. 193. Australymexylon australe, male, abdominal ter g ite VIII, dorsal view. 5 5 mm Fig. 194. Australymexylon australe, female genitalia (ventral). 331. 332. Fig. 195. Australym exylon australe-, coxite and stylus of female genitalia (ventral). 195 Figs. 196-197. Australymexylon fuscipennis, aedeagus: fig. 196, lateral view; fig. 197, dorsal view. Abbreviations: ML- median lobe, LL- lateral lobe, TS- tegminal strut, TG- tegminal plate. Figs. 198-199. Australymexylon australe, aedeagus: fig. 198, lateral view; fig. 199, ventro-medial view of lateral lobe. Abbreviations: END- endophallus, LL- lateral lobe, TP- tegminal plate, TS- tegminal stru t, ML- median lobe. 338. 'I* Fig. 200. Australymexylon australe, aedeagus, ventral view. Abbre viations: ML- median lobe, LL- lateral lobe, TP- tegminal plate, TS- tegminal strut. 340. 200 Figs. 201-202. Australymexylon australe, larva: fig. 201, apex of abdomen (lateral); fig. 202, antenna. 342. W W . • . 'K 'j£ -J•..*»■V.-V anus Fig. 203. Australymexylon australe, larva, tarsungulus. 345. 204 Fig. 204. Australymexylon australe, geographic distribution. 346. 205 Fig. 205. Australym exylon fuscipennis, geographic distribution. Fig. 206. Australymexylon australe, prothorax, ventral view. 347. TERGOSTERNAL SUTURt STCRNUM / PRONOTUM— Lv, • COXAL COXAL SLIT-— DEPRESSION ' rSus*' • MARGIN COXA HYPOMERON 206 349. ELEVATED^ MESEPISTERNUM a r e a j r y s . ^-"ELYTRAL GROOVE MESOSTERNUh — MESEPIMERON ^--EPIPLEURAL FOLD —METEPISTERNUM "METERMERON MTST 2 0 7 MTST 2 0 8 Figs. 207-208. Australym exylon australe: fig . 207, mesosternum, meta sternum, and associated structures; fig. 208, anterior area of metastern um with mesocoxa removed. Abbreviations: CX2- mesocoxa, CX3- metacoxa, MTST- metasternum. 350. 209 Fig. 209. Australymexylon australe , apex of procoxa (note slit). Fig. 210. Australymexylon australe , male, maxilla (right, dorsal) 351. m m Fig. 211. Australymexylon australe , labrum (ventral). Epipharyngeal setae omitted. Figs. 212-213. Australymexylon australe, labium; fig, 212, ventral view; fig. 213, dorsal view of'kypopharynx. 356. Fig. 214. Australymexylon australe , larva, maxilla, apex. 357. A A 214 Figs. 215-216. Australymexylon australe, larva: fig. 215, metathoracic leg; fig. 216, labium, left half (ventral). 359. 360. 216 MELITTOMMA-COMPLEX Within the subfamily Melittonuninae, there is the singular genus Australymexylon (defined by a number of autapomorphies) and a complex of genera which I refer to as the Melittomma-complex. Below, I propose a system using three generic taxa to handle the species of this group. My rationale is to preserve what has already been done taxonomically in this complex, without loosing sight of information about "natural" relationships (i.e., cladistic relationships) which has been found. Melittomma sensu stricto (as defined here) is a compact, well- defined, monophyletic group which includes those (mostly Pantropical) species with appendiculate paratergites on male abdominal segment nine. I believe it is worthwhile to recognize this distinctive lineage nomenclaturally (as Melittomma), and not to dissipate its integrity through inclusion of other taxa which are, overall, more primitive in morphological structure (e.g., insulare and Melittommopsis spp.). Males are not known for species of Melittommopsis Lane. All of these heavily-sclerotized, melandryid-like beetles occur in South America and closely resemble the North American species Melittomma sericeum in appearance (as well as Melanomelittomma). In the absence of males, it is difficult to determine if these South American forms are closely allied to other Melittomma (i.e., do the males have the ninth paratergites appendiculate?) or to Melanomelittomma (are the lateral lobes large?). The females do, however, have some 361. Fig. 217. Melittomma brasiliense, male, habitus, dorsal view. Drawing by J. Nagy. 362. 217 364. peculiarities of their own, such as the expanded apical palpomere of the female maxillary palpus. How can this diversity best be dealt with? There are, of course, several possibilities. First, all of these taxa could be "lumped" under Murray's Melittomma (the case until Lane's publication of 1955). This, however, would dilute and de-emphasize the unique structural attributes of Melittomma in a strictly-defined sense. I am confident of the monophyletic status of this genus (strictly defined), but less so for the complex as a single lineage. Therefore, I would not favor loss of what little firm, cladistic information is now in hand. It would also be unfair to supress Lane's genus (Melittommopsis) until it is given sufficiently complete investigation to determine its closest relatives (a job which awaits discovery of males). Melittomma insulare, a serious pest of coconut palms in the Seychelles and Madagascar presents yet another problem. This beetle is definately not closely allied to Melittomma, sensu stricto, and has some apomorphies of its own. Are the South American forms closer to insulare or Melittomma, s.s.? I do not know. We cannot satisfactor ily answer this until males are collected or perhaps other female and/ or larval characters are elucidated. It would be too presumptuous to combine these South American species with ither taxon without a complete understanding of the overall pattern of relationships. An immediate solution, and the one I follow, is to recognize a third genus for insulare until this complex is unravelled. This is indicative of the actual situation and preserves what we do understand regarding 365. relationships, while emphasizing where the major problems lie. Per haps a more intensive review of world species for all three taxa will point to additional characters or additional taxa which have bearing on these questions. Although these genera were keyed out above in the key to sub families and genera, I present a brief key here for reference in the discussions which follow. KEY TO THE GENERA OF THE MELITTOMMA-COMPLEX 1 Body heavily sclerotized OR lightly sclerotized; female maxillary palpus with apical segment narrow; male with paratergites of abdominal segment IX long, appendiculate; aedeagus with lateral lobes small, setose; larval abdo minal segment IX truncate and rimmed by teeth; d istri bution Pantropical + North America ------Melittomma Murray Body heavily sclerotized; female maxillary palpus variable; some males without appendiculate ninth paratergites (other males unknown) and with very large lateral lobes bearing rows of stout teeth mesally; larva with ninth abdominal segment crenulate around rim (or larva unknown); d istri bution: S.America OR Madagascar-Seychelles + Sumatra (?)—2 2 Female maxillary palpus with apex strongly widened; males and larvae unknwon; S.America Melittommopsis Lane Female maxillary palpus with apical segment not widened; male ninth paratergites not appendiculate; lateral lobes very large, elongate, with rows of stout teeth medially; Madagascar, Seychelles, and (?) Sumatra ------Melanomelittomma, new genus 366. Genus M elittom m a Melittomma Murray, 1868: TYPE SPECIES- Hylecoetus brasiliensis Laporte, 1832 INTRODUCTION I am using Melittomma in a new and restricted sense, to include those Pantropical species listed in Table 12, This tabular list is, incidentally tentative and subject to alterations when all of the types have been examined. From descriptions, illustrations, geographic distributions, identified specimens, and other clues I am relatively satisfied with the accuracy of their inclusion here. Nonetheless, it is imperative that the types be examined carefully, and it might also be noted that a revisional study of the species is very badly called fo r. While most of the species included here are pale colored, lightly- sclerotized, and are tropical in occurance there is one North American species which is more like Melittommopsis and Melanomelittomma in appearance, being darkly colored and heavily sclerotized. Because of this atypical species, I am including a description of both Melittomma S brasiliense (type-species for the genus) and Melittomma sericeum (th e unusual, North American species). T a b le 13 . L ist of species assigned to MELITTOMMA, sens u n o v o . SPECIES______AUTHOR, DATE, PAGE______GEOGRAPHIC DISTRIBUTION pervagus NEW COMBINATION (Olliff), 1889, p. 87 AUSTRALIA vigilan s NEW COMBINATION (Lea), 1911, p. 467 AUSTRALIA africanum (Thomson), 1858, p .8 2 ;V illie r s, AFRICA 1969a: 78 auberti Fairmaire, 1891, p. 246; Villiers, AFRICA * 1969a: 81 benitonum (Fairmaire),1901, p. 181; Villiers, AFRICA 1969a: 80 brasiliensis (Laporte), 1832, p.398 CENTRAL & SOUTH AMERICA coomani Pic, 1945, p.l TONKIN (Hoa Binh) curtipenne Pic, 1939, p. 32 MADAGASCAR javanicum (Chevrolat), 1829, p. 57 JAVA, SUMATRA, NEW GUINEA la teritiu m (Fairmaire), 1887, p.156; Lane, SURINAM 1955, p .150 marginellum (Schenkling), 1914, p .320; Lane, ECUADOR 367 P .150 T a b le 1 3 . L i s t o f s p e c i e s o f MELITTOMMA (Cont'd.f. SPECIES______' ' '■ AUTHOR, DATE, PAGE GEOGRAPHIC DISTRIBUTION p errieri Fairmaire, 1901, p.181 MADAGASCAR pubicolle Pic, 1944 f p ,l BRAZIL sicardi Pic, 1939, p.32 MADAGASCAR sericeum (HarrisI, 1833, p.52 NORTH AMERICA javanus NEW COMBINATION ■(Kurosawa1, 1956, p. 81 JAVA ph ilippen sis NEW COMBINATION (KurosawaJ, 19.56, p . 82 PHILIPPINES ablbitarsis Blair, 1936, p . 154 INDIA Melittomma brasiliense (Figs. 217-221, 223-238, 274, 275, 279) Hylecoetus brasiliensis Laporte, 1832: 398 Melittomma b ra silien se (L ap o rte), Murray, 1868: 314 GENERAL DESCRIPTION- Body elongate, nearly parallel-sided, and cylindrical in cross-section. Lightly sclerotized throughout (a little more heavily sclerotized than Hylecoetus, but much less so than Melittomma sericeum, Melittommopsis s p p ., or Melanomelittomma spp. ). Surface covered by dense, small, punctures each bearing a small, fine s e ta . COLOR- Females almost entirely testaceous to reddish-yellow; often with a slight darkened area on humeral angles of elytra and along the lateral edge of the pronotum. Males variable: with more or less exten sive black markings along the edge of the pronotum and elytra. These variations in the color of the males have apparently been responsible for several synonymies ( lateritium, marginellum: see Lane, 1955) which have not yet been resolved. HEAD- Males with vertex small, triangular, located between and posterior to the very large eyes. Eyes widely separated ventrally, and nearly touching along the midline dorsally. Females with eyes more widely separated dorsally than in males. The head is abruptly narrowed behind the eyes forming a distinct neck-region. Gular sutures nearly 370. parallel, slightly divergent posteriorly. Eyes. Very large (see above). Facets large, "coarse", more or less hexagonal in shape. Antennae. The antennae are sexually dimorphic. In males they are produced asymmetrically (ie., slightly serrate). In females, antenno- meres III-X are each bifurcate. (It is interesting that many museum specimens are arranged such that brasiliense is represented only by these pale, bifurcated-antennal females; males, with the dark colored margins are often under other names, lateritium or marginellum). Labrum and epipharyngeal region. Labrum small, subquadrate, with dense setae on dorsal surface. Ventral surface along anterior margin and for a short distance along the midline there are dense, long, stout setae which converge with those overhanging the anterior margin from the dorsal surface to form a dense comb of marginal setae (which are usually broken off, giving them a more robust appearance than if they were tapered apically). Epipharyngeal region is membranous and rather simple; covered by very dense microtrichiae which also meet along the midline forming a poorly defined medial "ridge". Mandibles. Short, broad, "typical": outer margin with long setae, molar area with patch of microtrichiae, no prosthecal elements visible, and with two dens (one apical, one subapical). 371. Maxilla. The structure of the maxillae is sexually dimorphic: males have a maxillary palp-organ, females do not. The form of the male maxilla is described first, and the female is then contrasted below. Maxilla: male. The maxilla is primitive in structure. Basigalea small, distinctly separated from large, rounded distigalea which is covered by dense setae and bordered by very dense, long setae. A small, narrow, triangular basistipes is separated from a larger, ovate dististipes by a suture. The palpifer is narrow, almost ringlike. The palpus has four palpomeres. Palpomere I is small, short, and broad in form; II abruptly larger and transverse; III much larger than II, very transverse and broadly oval; IV arises from III and is very long, and narrow and expanded apically; the apex of IV bears several minute, long, blunt, thin-walled sensilla; the surface of the basal "stalk" of IV is wrinkled in appearance much like that of the surface of the various branches of the maxillary palp-organ, but the clavate apical portion is not and has transverse lines to hexagonal impressions and setae which are similar to those of basal palpomeres. The maxillary palp-organ is large and multi-branched, and arises laterally from the apex of palpomere III; it is connected by a wrinkled membranous area. The base of the primary branch has a small, highly-sclerotized area which I believe is homologous with the auxiliary sclerite of Hylecoetus. I The maxillary palp-organ has 1°- 5° branches, and the surface has the 372. characteristic wrinkled appearance seen in all other lymexylids which have a maxillary palp-organ. The outer surface is covered by more-or- less well defined rows of stout, tactile setae, and the inner surface by very long, thin-walled sensilla (see also Slifer et al, 1975, who discuss this structure in M. sericeum\. My figure illustrates only % of the branches of the maxillary palp-organ (i.e., there is another, parallel set of branches!). Maxilla: female. The female maxilla is very similar to that of the male, except for the form of the palpus and the lack of the maxillary palp-organ. The palpus has four palpomeres: I is short and broad; II very long, narrow at the base and widened apically; III is shorter than II but also gradually widened apically; IV is longer than III, gradually widened apically and the apex is membranous and bears many minute sensilla similar to the sensilla at the apex of male palpomere IV. Labium and hypopharyngeal region. The labium is simple, similar to that of M. sericeum (described below), except the long, marginal hypopharyngeal setae are simple (not forked) and the ligula does not have a medial notch. THORAX. The Prothorax. The pronotum (Figs. 223-224) is elongate in,males, rounded anteriorly with the posterior angles slightly pro duced. In females the pronotum is shorter than in males (i.e., broader in form). The pronotum has sharp lateral margins. The hypomeron does not have the slots which characterize related taxa of the genus-complex. The procoxae are slotted apically. 373. Pterothoracic structures. The mesosternum is small, triangular, rounded posteriorly without an intercoxal process. Mesepisternum is elevated anteriorly. Mesepimerion elongate, without a groove for the elytral epimeron. The raetepimeron has a deep groove which receives the elytral epimeron. The scutellum Gneso-} is simple, narrowed posteriorly without any carina or "keel”. The elytron is long and narrow; a tongue-and-groove mechanism is present, but ends before the apex; the elytra nearly cover the abdomen, about h . of the apical tergite is exposed in males and up to 2 tergites in the female (espe cially when the abdomen is distended with eggs]; the surface of each elytron has 4 costae and a short, sutural costa. The metendosternite has a narrow lamina. The hindwing has relatively complete venation. ABDOMEN. Both males and females have five visible sternites: III- VII. The abdomen is robust, and more-or-less cylindrical in cross- se c tio n . female GENITALIA AND POSTABDOMEN. Abdominal segment VIII is small, lightly sclerotized, and not clearly defined. There is a long spiculum gastrale, and a long connecting membrane leading to the genitalia. The genitalia have the distal parts of the coxites differentiated, more highly sclerotized and covered by long, fine setae. An unusual feature is that the proctigeral baculi are fused for most of their distance, narrowly divergent apically and more widely divergent basally. MALE GENITALIA AND POSTABDOMEN. (Figs. 231-235]. Segment VIII is small and not clearly defined, but does support a pair of long, ventral 374. struts which are broad and fused apically (anteriorly). The parater gites of abdominal segment IX are very large, appendiculate, and cover ed by setae and dense, short, broad spines. The aedeagus consists of a very small, transverse tegminal plate which is immovably fused to the two small, medial, proximal lateral lobes which are covered by fine setae. The median lobe is enlarged basally and much narrowed apically, and curves forward at the base and is attached to the tegminal struts. The struts are long and wide. DISCUSSION. Melittomma b ra silien se is very representative to most species placed in my restricted concept of the genus. Most of the species are similarly soft-bodied, with most of the features discussed here including those pertaining to tie aedeagus and the bizarre para tergites of abdominal segment IX. Villiers (1969) discusses the species of Africa, and presents the only modern systematic considera tion of any Melittomma. Table 12 lists the described species which I place in Melittomma. Lane (1955) realized that several names seem to apply to brasiliense. Variations in color and form (discussed above in the description) and very broad geographic ranges ( brasiliense occurs in Central and South America, including the West Indies and perhaps an occassional occurance near the United States border), have served to encourage synonymy form a tio n . Two species from Australia (Hylecoetus pervagus & Hylecoetus vigilan s) are transferred into Melittomma, and a genus described by 375. Kurosawa in 1956 ( Neohylecoetus) is synonymized with Melittomma and its species transferred accordingly (javanus: JAVA; philippensis: PHILIPPINES) (See Appendix A: Item 19). All of the species of Melittomma need to be critically revised, including careful comparisons of the forms which occur in Southeast Asia extending to Australia. 376. Melittomma sericeum (Figs. 239, 241-253, 276, 286-289) Lymexylon serciceum Harris, 1833: 52 Melittomma sericeum (Harris); Leng, 1920: DESCRIPTION: GENERAL. Body elongate, broad, + parallel-sided; head flexed under pronotal "hood"; dark reddish-brown to nearly black, covered with very fine, golden setae; body heavily sclerotized through out; head and pronotum with dense, coarse punctures; elytra with dense, round, raised (almost bubble-like in appearance) bumps, sometimes aligned into more or less distinct, transverse rows. See dorsal and lateral habitus drawings (Figs. 241-242). Maxilla: Male. The stipes is divided by a distinct suture into a narrow basistipes and large dististipes. The lacinia is long, narrow, separated from dististipes by suture; median side with very long, dense setae. Galea divided into small, transverse basigalea and large, broad distigalea; distigalea covered apically by very long, dense setae. Palpifer is large; mesal margin is indistinct ventrally, and the palpifer curves around the side and onto the dorsal surface (ie., ringlike in form). Palpus with four palpomeres: I small; II longer; III largest (though not longest) palpomere, broad; IV very long, parallel-sided, narrow, with several apical short s e n s illa . 377. The maxillary palp-organ is very large, complex and conspicuous. This basket-like structure has two parallel rows of 2°-branches (only one of these rows is illustrated in Fig. 250). Each secondary branch bears two or three (usually three) 3°-branches. The tertiaries have only long, fine setae. Inner margins of primaries and secondaries also have these long, fine setae; the outer surfaces of these (but not of tertiaries) are covered by dense, shorter, stout setae. The inner, thin setae are chemoreceptive in function, while the thick, fluted, outer setae are tactile as discussed by Slifer et al (1975) in a very detailed morphological study of the maxillary palp-organ for this species. Slifer et al (1975) also includes illustrations and electron micrographs of interest. Labium. Mentum small, transverse with a few setae. Palpigers distinctly separated; each consists of a small sclerite with long, heavily-sclerotized lateral areas; medially, they become gradually membranous, and are separated by an emargination. Palpi with three palpomeres: I short, transverse; II longer, widened apically; III, large, broad, nearly parallel-sided, apex with many small sensilla. Ligula simple, transverse, with a deep, medial emargination. Hypo- pharynx large and membranous; setae over most of surface simple, however, those along margin are forked (Fig. 244). 378. Mesosternum and associated structures. The anterior area of the mesosternum is elevated, broad, and emarginate posteriorly. The post erior part of the mesosternum is continued medially as a very long, pointed inter-coxal process. The large coxal depressions (for the mesocoxae) are margined by a sharp carina. The mesepistemum has the anterior area elevated and separated fromthe posterior part by a sharp carina. The mesepimeron has the lateral portion depressed (to receive the epipleural fold of the elytron in repose), and separated from the remaining (postero-medial) portion by a sharp carina. Male postabdomen and genitalia. The eighth abdominal sternite of the male is produced medially into a long, "clavate" process (Fig. 286). The paratergites of abdominal segment IX are short and very broad, but are distinctly appendiculate (Fig. 248). The surface is covered by dense, stout spinose setae (Figs. 248, 246) and the mesal margin bears a row of dense, stout setae as well (Figs. 247, 248). The aedeagus consists of an elongate and robust median lobe which is large relative to sizes of other genitalic parts. The tegminal plate is largely membranous and is poorly delimited; the struts simple. The lateral lobes are long, and covered by very dense, long setae. There is a heavily sclerotized, internal, concave "receptacle"-like structure under the base of each lateral lobe (Fig. 245). The female genitalia. The distal part of each coxite of the female genitalia is heavily sclerotized (i.e., highly differentiated from the 379. membranous basal part of the coxite), separated by a well developed coxital fold. The larva. The abdominal segment IX of the larva is circular and truncate and the margin is bordered by very wide, truncate teeth (Fig. 251). The legs have many stout spines, and the tarsungulus is bisetose (Figs. 252, 253). Discussion. I have included the above descriptive remarks on M. sericeum (the single North American species of this complex) because it differs so much from the type-species of •Melittomma, sensu stricto (M. brasiliense). Most members of Melittomma are soft-bodied, and to a large extent very similar to brasiliense (including the long, thin form of the ninth paratergites of the male abdomen), but sericeum is the exception. This species looks very much like species of the related genera Melittommopsis and Melanomelittomma. For this reason I have pointed out some interesting features, although in less detail than that given for brasiliense. Genus Melanomelittomma, new genus Type-species: Melittomma insulare Fairmaire (here designated! Melanomelittomma in s u la re , NEW COMBINATION (Figs. 256-268, 270-273, 277, 278] General Description Body Form. Body elongate, but robust; elytra a little wider than pronotum, broad, attenuate posteriorly. Habitus almost melandryid- lik e . Color. Dark reddish-borwn to black; color uniform. The Head. The head is relatively small; widest at eyes and sharp ly narrowed behind eyes forming a narrow, constricted neck. Eyes very large, nearly touching dorsally and actually touching ventrally. Gular sutures short (confined to lengh of neck region) and slightly conver gent anteriorly. Antenna. Long and narrow; antennomere I large, narrowed basally; II much smaller than I; III elongate and widened apically; IV-X serrate (asymmetrically produced toward body midline); XI very long, narrowed in apical half. Antennomeres I-III with long, fine setae only; IV-XI with both long, fine setae and short sensilla. Labrum and epipharyngeal region. Labrum small, rounded, lobate. Epipharynx simple (see M. brasiliense description). 380. Table 14. List of species of Melanomelittomma, new genus. Species______Author, date, page Geographic distribution insulare Fairmaire, 1893, p.323 Madagascar; Seychelles undescribed species* Sumatra *probable placement, based on single female specimen which is elongate in shape and has biramous antennae, similar to those of female Melittomma brasiliense. 382. MANDIBLE. Short and broad. Apex with two dens: one apical, one subapical. Prostheca present, consisting of a dense, long patch of setae. .Molar region marked by short, fine, more-or-less parallel rows of setae. A small patch of setae is present near the point of muscle attachment at the mesal base of the mandible. MAXILLA. The maxilla is primitive and complete in structure, including separate basigalea/distigalea and basistipes/dististipes. The lacinia and distigalea are relatively small. The lacinia is long, narrow, and covered by long setae on apical half. The distigalea is broadly rounded, with very dense setae apically. The male palpus bears a small, simple maxillary palp-organ on palpomere III, and has four palpomeres: I small, little longer than wide; II, much larger than I, broad; III, larger than II, broadly rounded; IV, elongate and nearly parallel-sided with minute apical sensilla. Maxillary palp-organ with primary to secondary branching pattern (1°~ 2°); outer surface of each branch covered by short, broad setae and inner surface by dense, long, very fine sensilla. Female maxilla as in male, except palpomeres II and III are more elongate in shape, and there is no maxillary palp- organ. LABIUM AND HYPOPHARYNGEAL REGION. Sim ple, com plete. Very similar to M. brasiliense. 3.0 THE THORAX PRONOTUM. Wide; broadly rounded anteriorly; posterior angles only slightly produced. Lateral margin sharply defined; margin 383. extending around entire parimeter of pronotum. Hypomeron wide, with supracoxal slot. Tergo-sternal suture unusually deeply impressed. PROSTERNUM. Relatively long, with a short, pointed, inter- coxal process. PROCOXAE. S lo tte d a p ic a lly . TIBIAL SPUR FORMULA. Male: 2-2-2; fem ale: 2-2-2. MESOSTENRUM AND ASSOCIATED SCLERITES. Mesosternum broad, triangular, with small, anterior, triangular area heavily sclerotized. Mesepisternum with anterior area raised and set off from the posterior area by a sharp carina. Mesepimeron with lateral groove for epipleural fold of elytron; set off by sharp carina. METASTERNUM AND ASSOCIATED SCLERITES.Metasternum r e la tiv e ly short and broad, with longitudinal suture along midline; mesocoxal depression large, separated from metasternal plate by sharp carina; with narrow, short emargination at middle, behind the coxal depressions. Metepistemum braod, with tongue-and-groove-like locking mechanism with lateral edge of metasternum. Metepimeron narrow with deep groove which receives elytral epipleural fold at rest. SCUTELLUM. Simple, rounded apically, without carinae. ELYTRON. Elongate, broad and nearly covering abdomen; narrowed apically; tongue-and-groove locking mechanism complete to apex; surface with four longitudinal carinae and a short, oblique sutural carina (nearly traceable along suture longitudinally); a narrow epipleural area is separated by a sharp carina. 384. HINDWING. As in other members of the genus complex, the venation is primitive and nearly complete, (see sericeum) METENDOSTERNITE (Figs. 262-263). Lateral parts of the furcal stalk fused in distal half. Lamina short, transverse. Anterior tendons arise on short stalks on the lamina near the middle. Furcal arms long, twisted, anteriorly-directed. 4 .0 - THE ABDOMEN GENERAL. Both males and females ahve stem itesiH -V II visi ble. The abdomen is broad, short relative to body length, and cylin drical in shape to a greater or lesser degree. EXPOSURE. The elytra nearly cover the abdomen; only the tip of tergite VII is visible from a dorsal aspect. MALE POSTABDOMEN. Abdominal segment VIII is lig htly sclero tized and telesoped into VII; the stem ite is emarginate (deeply) medially. Segment IX is greatly reduced, and the paired (anteriorly- directed) struts are fused apcially. MALE GENITALIA. The lateral lobes are very large, long, and widened apically. Apex of lateral lobe curved toward midline, and bearing a row of very large, broad spines; the surface of the lateral lobe is covered by very long, fine setae. The median lobe is very long (nearly as long as tegminal plate + lateral lobe lengths combined), not very wide, and narrowed apically (the apex is small, very slightly enlarged, and rounded). The tegminal struts are short, broad, and fused apically. The tegminal plate is small, transverse and basal. 385. • FEMALE POSTABDOMEN. Abdominal segment V III sim ple, very lightly sclerotized, narrowed apically (posteriorly). FEMALE GENITALIA. Long, narrow, lightly sclerotized. Proc- tigeral baculi narrow and only moderately sclerotized; separate and nearly parallel apically, proximal along their middle lengths, and widely divergent basally. Valviferal baculi widely separated and nearly parallel throughout entire lengths; heavily sclerotized. Coxital baculi narrow, oblique. Basal part of coxite short and wide; distal part of coxite very long and narrow, attenuate apically, membranous, and covered by long setae; mesal margin on dorsal surface heavily sclerotized (this sclerotization becomes very narrow distally). Styli long, narrow, membranous, covered on apical half by dense setae, and bearing one very long seta apically. Genus Melittommopsis Melittommopsis Lane, 1955: 152 TYPE-SPECIES: M elittommopsis ju q u ie n sis Lane, 1955: 152 The following descriptive remarks are based on specimens of Melittommopsis sp. from Brazil. As Lane (1955) discussed the notion, it is feasible that one of his species could be synonymous with the Schenkling species validum; this, of course, makes application of a credible name difficult. Among all of the museum collections which I have studied, I have not seen even a single male specimen for this genus. This is the source of confusion most pressing in the Melittomma- complex. Except as noted, the species before me is very similar to Melanomelittomma insulare in structure. THE HEAD General form. Eyes large, but widely separated dorsally and narrowly separated ventrally. Antenna. Subserrate; XI less elongate than in insulare. Labrum and epipharyngeal region. Small; dense patch of long setae medially, and dense microtrichiae posterior and lateral to this patch of longer setae. Mandible. Similar to insulare, with two apical dens. Prostheca present, but reduced to a small patch of microtrichia. T a b le 15. L i s t o f s p e c i e s o f MELITTOMMOPSIS. SPECIES AUTHOR, DATE, PAGE GEOGRAPHIC DISTRIBUTION juquiensis Lane, 1955, p. 152 BRAZIL nigra Lane, 1955, p. 156 BRAZIL validum NEW COMBINATION Schenkling, 1914, p. 321 BRAZIL 387 388. Maxilla (Figs. 280, 281). Primitive, complete. Lacinia broad; galea very broad. Female palpus long: I short, but elongate; II, very long and widened apically; III, much shorter than II; IV large, securi form, apex membranous and covered by short, pointed sensilla. Labium and hypopharyngeal region. Ligula without medial notch; hypopharyngeal region simple; palpus with palpomere III securiform; hypopharyngeal setae very long and dense. THORAX Pronotum. Elongate, slightly narrower than combined elytral width; rounded anteriorly; slightly sinuate posteriorly; postero-medial area raised (with "hump"); with sharp margin which extends entirely around the pronotum; hypomeron with supracoxal slot; tergo-sternal suture obscure (but complete). Procoxae. Slotted apically. Tibial spur formula. Male: unknown; Female: 1-2-2. Mesosternum and Associated Sclerites. Mesosternum broad, triangu lar, with small, elongate (and posteriorly-pointed), elevated area anteriorly (Fig. 283). Mesepistemum with raised, elevated anterior area, separated by sharp carina. Mesepimeron with depressed lateral area which receives elytral epipleuron, but which is NOT set off by a carina (as in insulare). Metastemum and Associated Sclerites. Without medial emargination of metasternum posterior to the (meso-) coxal depressions. These coxal depressions are separated from the metasternum by carinae. 389. Elytron as in insulare, but carina three (see Fig. 45 for term explanation) and sutural carina not evident. ABDOMEN Female Genitalia (Figs. 284, 285). Long and narrow; lightly sclerotized in large part. Proctigeral baculi long; narrow anteriorly and broader posteriorly (i.e., distally); separate and more-or-less parallel. Valviferal baculi long and parallel (except slightly diver gent near middle). Coxital baculi transverse; oblique laterally. Basal part of coxite short and broad, sclerotized with a few short setae near apex. Coxital fold wide, very conspicuous. Narrow area at base of distal part of coxite sclerotized, but most of the long distal parts are membranous and without pigment and with many long setae. Styli long, slightly clavate in form; basal half sclerotized, without setae; apical half membranous, covered by setae, and bearing one very long seta at the apex (Fig. 285). Genus Hylecoetopsis, INCERTAE SEDIS (Fig. 290 ) In 1955, Lane proposed a genus, Hylecoetopsis, for the single sp ecies Hylecoetus cylindricus Germer (1912); Germer's publication served to validate a Dejean checklist name. Lane did not see the only known specimen of this bizarre species, and based his genus on the one drawing of the antenna published by Germer. I have written many museums in Europe, and have been unable to determine where the type is located. Until it is found, it is doubtful that the significance of the species will be known. The antennal structure is indeed unusual. I would judge unusual enough to seriously question the familial placement of this taxon. What this species might be is an intrigueing question, but the antennal structure resembles nothing else in Lymexylidae that I have seen. Some Neotropical species of the tropical family Mycteridae (see Crowson, 1955) which I have seen look somewhat like lymexylids (I have received them among unidentified Lymexylidae from various museums), but the antenna figured by Germer would be atypical for myceterids as well. The search for the type specimen, and intensive collecting in South America (particularly around Cayen.ne, French Guiana- the type locality) are badly needed so that the systematic position of this species can be settled. I place the genus in Incertae Sedis, and do not consider it further in this study of Lymexylidae. 390. Fig. 218. Melittomma brasiliense, male, head (dorsal). 391. 392. o D Fig. 219. Melittomma brasiliense, female, head (dorsal). 394. 219 Fig. 220. Melittomma brasiliense, male, antenna. 395. 396. X I IX VIII VII VI III PEDICE L 220 SCAPE Figs. 221-222. Antennae; fig. 221, Melittomma bramHense, female; fig. 222, Lymexylon navale, male. 397. 398. Figs. 223-225. Melittomma brasiliense; fig. 223, prothorax (lateral); fig. 224, pronotum (dorsal); fig. 225, elytron, right, dorsal view. * 399. 400. Figs. 226-228. Melittomma brasiliense, male, maxilla: fig. 226, apex of palpomere IV, representative setae, sensilla, and surface detail only (other omitted); fig. 227, maxilla with maxillary palp-organ; fig. 228, portion of tertiary branch (3 ) of maxillary palp-organ (note short tactile setae, and long sensilla). la c in ia Figs. 229-230. Melittomma brasiliense, female genitalia: fig. 229, dorsal view; fig. 230, sclerotized process located between basal parts of coxites. 1230 Figs. 231-235. Melittomma brasiliense, male: fig. 231, aedeagus (lateral); fig. 232, apex of paratergite of abdominal segment nine; fig. 233, aedeagus (ventral); fig. 234, aedeagus (dorsal); fig. 235, abdominal segments VIII and IX internested (ventral), note large, appendiculate paratergites of segment IX. 231 232 Fig. 236. Melittomma brasiliense, metendosternite. 407. Fig. 237. Melittomma brasiliense, metepimeron and base of metacoxa (note groove which receives epipleural fold of e ly tr o n ) . 409. 410. Fig. 238. Melittomma brasiliense, hindwing. Abbreviations: see Table 4. 411. 412. Fig. 239. Melittomma sericeum, hindwing. 413. 414. Fig. 240. Melittomma sp. CBrazil), hindwing. 415. 416. 2 4 0 Figs. 241-242. Melittomma sericeum, male, habitus: fig. 241 (dorsal) fig. 242 (lateral view of forebody). Drawings by J. Nagy. Fig. 243. Melittomma sericeum, mesosternum, metastemum, and associa ted structures (ventro-lateral). Abbreviations: st2a- anterior part of mesosternum; st2b- posterior part of mesosternum; eps2a- elevated, anterior part of mesepisternum; eps2b- posterior part of mesepisternum; epra2a- posterior part of mesepimeron; epm2b- depressed part of mesepi- meron which receives elytral epipleural fold; tn2- mesotrochantor; icp- intercoxal process; cx2- mesocoxa; eps3- metepistemum; epm3- metepi- meron; cx3- metacoxa; st3- metastemum; tr3- metatrochantor; fm3- meta femur . CPS 3 GPFP3 CX3 243 Fig. 244. Melittomma sericeum, labium (ventral). 421. 422. tt?an1^ sericeum, male: fig. 245, aedeagus j-J T f l &‘ 246» sPme on surface of paratergite IX: fig 247 row of spines on mesal margin of paratergite IX; fig. 248, paratergite IX. 424. Figs. 249-250. Melittomma sericeum, male: fig. 249, base of maxilla (NOTE: cardo removed, and all setae omitted); fig. 250, maxilla, in cluding m ax illary p alp -o rg an . A bbreviations: ST- s tip e s , PLF- p a lp i- fer, LC- lacinia, GA- galea, PI- palpomere I, P2- palgomere II..., AXS- auxiliary sclerite, MPO- maxillary palp-organ, 1 -primary branch (MPO), 2°- secondary branch... 425. DISTIGALEA MPO PALPUS — LACINIA \ PALPI FE R -- 250 BASISTIPES Fig. 251. Melittomma sericeum, larva, habitus, lateral view. 427. Figs. 252-253. Melittomma sericeum, larva, proleg: fig. 252, mesal surface; fig. 253, lateral surface. 429. 430. a Figs. 254-255. Melittomma sp. (New Guinea): fig. 254, antenna; fig. 255, apex of antennomere VIII. •I 431. 432. M i Fig. 256. Melanomelittomma insulare, new combination, female, habitus (lateral view of forebody). Drawing by J. Nagy. 434. Fig. 257. Melanomelittomma insulare, female, head (dorsal). 435. 257 Fig. 258. Melanomelittomma insulare, male, aedeagus, lateral view. 437. 258 Figs. 259-261. Melanomelittomma insulare, male genitalia: fig. 259, aedeagus (ventral view); fig. 260, apex of lateral lobe (note row of spines); fig. 261, apex of endophallus. Abbreviations: LL- lateral lobe, ML- median lobe, TS- tegminal strut, TP- tegminal plate. 439. 261 440 Figs. 262-263. Melanomelittomma insulare, metendosternite: fig. 262, apical portion; fig. 263, base. Abbreviations: VLF- ventral flange, SK- furcal stalk, L- lamina, FA- furcal arm, AT- anterior tendon, ATS- anterior tendon stalk. 441. 442. 263 —VLF Fig. 264. Melanomelittomma insulare, male, antenna. 443, EL V 2 6 4 Figs. 265-266. Melanomelittomma insulare: fig. 265, mesal part of mandible (dorsal); fig. 266, apex of mandible. 445. 446. Figs. 267-268. Melanomelittomma insulare: fig. 267, maxilla, male; fig. 268, maxillary palpus, temale. 447. i i [ Fig. 269. Atractocerus brevicornis, female genitalia. 449. 2 6 9 cFo g t e ^ 0 ;g L ;^ n-°;;iu “ t°^ s^ ^ - .fr ie *enitaiia: ««• lus; CX8- coxital bacillus! CF- coxital S valviferaI bac“- 451. 270 Fig. 272. Melanomelittomma insulare, male, abdominal segment IX. Abbreviation: P9- paratergite of abdominal segment IX. 453. 454. RUT 455. SLOT 273 Fig. 273. Melanomelittomma insulare, procoxa (apex). Fies. 274-275. Melanomelittomma insulare, female genitalia, procti- geral baculi (dorsal): fig. 274, basal ends (anterior); fig. 275, distal ends (posterior). 456. Fig. 276. Melanomelittomma insulare, apex of protibia. 458. 459. 2 76 Fig. 277. Melanomelittomma in su la re, pupa, head and pronotum (la te r a l). 460. 461 • \ 277 Figs. 278-279. Apex of larval abdomen: fig. 278, Melanomelittomma insulare; fig. 279, Melittomma brasiliense. st§8S Figs. 280-282. Melittommopsis sp.. female: fig. 280, maxillary palpus; fig. 281, apex of maxillary palpomere IV (corner); fig. 282, labial p a lp u s. '464. Fig. 283. Melittommopsis sp. (Brazil), mesostemum and associated sclerites. Abbreviations: ST2a- anterior part of mesostemum, ST2b- posterior part of mesostemum, EP2a- anterior part (elevated) of mesepisternum, EP2b- posterior part of mesepistemum. 466. 467. ST 2a EP2a ST2b \E P 2 b 283 Figs. 284-285. Melittommopsis sp., female: fig. 284, h apex of female genitalia; fig. 285, stylus. 468. 469. Fig. 286. Melittomma sericeum, male, apex o£ abdominal sternite VIII. 470. 471, Figs. 287-288. Melittomma sericeum. female genitalia: fie 287 coxites; fig. 288, distal part of coxite. 472. Fig. 289. Melittomma sericeum, insertion of anterior tendon of metendostemite. MLP Fig. 290. Hylecoetopsis cylindricus, antenna. Redrawn from Germer. 1912. . ------CLADISTIC ANALYSIS Under "Philosophical methods" above I outlined some of my views on cladistics, and a more detailed discussion of some points is found in Watrous § Wheeler (Systematic Zoology: in press). What follows is my analysis of the cladistic relationships among the genera of Lymexyli- dae. That is, the sequence and pattern of branching in the history of the family constructed on the basis of shared, apomorphic features ("characters" in the sense of Platnick, 1979 and Watrous § Wheeler, in press). My analysis consists of two parts: (1) a discussion of the characters, including hypotheses of polarities; and, ( 2) a discussion of the monophyletic groups indicated by these characters (and my interpretations of them). The end result is an hypothesis of the relationships among the genera. Character State Analysis «- My explicit purpose for this section of the paper is to present a clear discussion of the polarity of character states which I use as the basis for hypothesis-formation with regard to cladistic relation ships among the genera of Lymexylidae. Because it is my plan that this study also serve as a basis for further research at the species level and also at the superfamilial level in the Cucujiformia I think that it 477. 478. is desirable to include comments on structures which I believe will be useful to these other studies, but which are not of direct importance to my cladistic analysis of the genera. To achieve this broader goal in the following section of the paper, I have arranged it "morphologic ally". That is, it proceeds from structures of the head to those of the postabdomen, with some general features of body-form first and the characters of larvae last (this, incidentally, is the same sequence followed in the section of external morphology and in each section which discusses an individual genus above). This arrangement gives a logical order which augments retrieval of information. Coleoptera Cucujiformia is so massive (in excess of sixty families of beetles, including some of the largest) that a comprehensive, compara tive study is far beyond the bounds of my study of the Lymexylidae. As will be discussed more fully below, the relationships of the lymexylids are unclear and as of now the family is curiously poised between the Cleroidea and Cucujoidea (Heteromera + ciavicornia). Thus, the logical out-group for the lymexylids would include these two superfamilies. If the available out-group is too large, how can we polarize the characters in order to hypothesize relationships of the genera? The answer was given in abstract terms by Watrous 8 Wheeler (in press), and in practice the following actions are taken. Some characters which are apparently unique to the Lymexylidae (i.e., autapomorphic), based on a broad but fragmentary sample of related superfamilies, are used to defend the hypothesis that Lymexylidae is a monophyletic unit. Next, 479. a monophyletic group within the family is recognized, based on several unique characters (again, autapomorphies). This intrafam ilial, mono phyletic group is then used as a "functional out-group" for polarization of other characters used in the generic analysis. Monophyly of Lymexylidae. A more exhaustive discussion of this topic is included in the next major section ("Monophyletic Groups") be low, but a few characters are mentioned here to defend my following polarity arguments. The partial division of the coxites of the female genitalia by the coxital folds(Figs. 53, 54) has not been seen in any other family of Coleoptera, and is believed autapomorphic for lymexylids (it occurs in all lymexylids examined). The maxillary palp-organ of the male is present in all genera except one (Australymexylon, new genus), and appears to be homologous in form and placement at the apex of maxillary palpomere III. Larval maxillae have the mesal surface membranous and bordered by long setae: I am not aware of this structure in other beetle larvae. Larval abdominal segment IX is very large, and superior to X; it is always produced posteriorly to a degree greater than that seen in most other larvae. Based on these considerations I hypothesize that the Lymexylidae are monophyletic. Functional Out-Group. Finding a monophyletic unit within the family was not difficult. The Hylecoetinae (subfamily) was recognized as unique and "monophyletic" by Boving § Craighead (1931) and by Crowson 480. (1955), based on both larval and adult structures, only a few of which need be cited here (others are also discussed below under "Monophyletic Groups"): epicranial pit, asymmetrical lateral lobes of the male genitalia, very long, "sword-like" ninth abdominal segment of the la rv a . Recognition of Hylecoetinae as monophyletic means that it can be used to determine polarities of characters by out-group comparison. In short, where two states of a character occur in other groups of the family, that state which occurs also in Hylecoetinae is plesiomorphic and the alternative state is an apomorphy (character sensu Platnick, 1979; Watrous 8 Wheeler, in press) which can be used to group together those taxa which share it. A Discussion of the Characters and Character States. Numbers which appear in the following discussion of the characters and states refer to apomorphies listed in table 16, and which appear in the cladograms. Pertinent illustrations are cited in table 16. Sometimes I refer to particular plesiomorphic states of characters which are the analogues of the apomorphies defined in the table. Where apomorphies are referred to the number appears in parentheses, e.g., ( 1) , and where analogous plesiomorphic states are referred to the number in parentheses is preceeded by a negative sign, e.g., (-1). The same numbers occur in text, table, and cladograms. BODY FORM AND GENERAL CHARACTERS.- Very elongate forms ( e .g ., Atractocerus, and to a lesser degree Australymexylon, and Melittomma sensu stricto, excluding M. sericeum) are apomorphic relative to shorter, broader body forms (e.g., Hylecoetus, Melittommopsis, Melanomelittomma). Because this feature is difficult to quantify, it is not used in the cladistic analysis. Forms which are highly compressed, or flattened, dorso-ventrally are also considered apomorphic relative to more cylindrical forms. Examples of such compressed forms would be Lymexylon or Australymexylon. But, Hylecoetus is very slightly compressed, and some Atractocerus could be interpreted thusly (to a limited degree). While this character is not used here, because it is difficult to define in unambiguous terms, it does indicate a variation in the family which should be noted. SCLEROTIZATION.- Hylecoetus, Australymexylon, Lymexylon, and most species of Melittomma (and many species of Atractocerus) are all soft-bodied (i.e., lightly sclerotized) and often covered by long, fine setae. The species of Melanomelittomma and Melittommopsis, and at least one species of Melittomma are all heavily sclerotized. It seems reasonable to argue that heavy sclerotization is plesiomorphic for the cucujiform series of beetles (e.g., primitive families of both Clavicornia and Heteromera are more-or-less heavily sclerotized and primitive Cleroidea, such as Peltidae, Trogositidae, Chaetosomatidae, 482. and Phycosecidae, are also heavily sclerotized), What then is the explanation for soft-bodied forms in most genera of Lymexylidae, and * in the "higher" Cleroidea (e.g., Melyridae and Cleridae]? I believe that Crowson gives us the answer in his study of the higher classification of Cleroidea (Crowson, 1964: 277): "In adult Coleoptera generally, 'Malacoderm' features are doubtless degenerative rather than primitive, associated with short-lived and free-living adults and often with some form of chemical defense". In all cases where we know something of adult lymexylids they are very short-lived, u su a lly th re e days to a week ( e .g ., Simmonds, 1956; Brown, 1954). During my field investigations at the Barro Colorado Island laboratory of the Smithsonian Tropical Research Institute in Panama (in 1978) I was able to keep adults of Atractocerus brasiliensis and Melittomma brasiliense alive in my laboratory, but always for no more than two or three days. Our understanding of the biochemical lives of the lymexyl ids is far more fragmentary. Certainly the elaborate increases in surface area of the palpi and antennae point to some kind of heightened sensory ability. Functions of defence, pheromone production-emission, * You may recall the historical significance of this question of soft- bodied forms. As discussed under my historical section the lymexyl- id beetles have often been classified with cleroid families or canthar- oid families because of the general sim ilarities among adult beetles in soft-bodiedness and clothing of long setae. 483. or other chemical activities are only speculative today, and no conclu sions can be made. The significant point, however, is that this soft- bodied condition is likely to be degenerative and as such easily subject to convergence (that is, degenerative features may arise any number of times and yet appear almost or quite identical}. Thus we can probably argue effectively that soft-bodied cleroids and lymexyloids (not to mention cantharoids, lampyroids and others) could easily have had independent development of these "malacoderm", apomorphic characters ( 1). THE HEAD.- The most obvious character of the head capsule is the presence (2) or absence (- 2) of a "neck" (a condition resultant of abrupt narrowing of the head behind the eyes). Only Hylecoetus and a few species of Atractocerus do not have the head abuptly narrowed behind th e eyes. Increased size of the eyes (3) and emargination of the anterior margin of the eyes (4) are also apomorphic characters (Hylecoetus has small, entire eyes). The epicranial pit is a unique feature of Hylecoetus (5) which I have not seen in any other lymexylid (or beetle). This strange structure (Figs. 2, 30) is of unknown function. Crowson (1955) guessed that it might be used to carry spores of the host fungus.' This seems superfluous, however, when we consider the elaborate fungal-pouches which are present on the female genitalia for just this function. This pit occurs in both sexes. 484. The size of the facets of the eyes vary, giving imprecise im pressions of "coarse" or "fine" facets with the dissecting microscope. Such differences will be useful interspecifically in Atractocerus and the Melittomma-complex, but are not useful at the generic level. Simi larly, the shapes of facets may be of value as well. Some are round while others are rather well-defined hexagons. I have not used microsculpturing of the head (or other parts of the surface) because they vary widely within the larger genera. Some interesting variations occur, however, including very deep punctures, quite shallow ones, and some cases of confluent punctures. A detailed stuyd of the integument by scanning electron microscopy promises to yield additional characters of value in comparative study of species. Filiform antennae occur in Lymexylon, while all other lymexylids have some modification in antennal structure. Slight to strong serra tio n ( 6 ) occurs in Hylecoetus, Australymexylon, and the Melittomma- complex. Short, compact, pointed, spindle-shaped antennae (7) occur in Atractocerus beetles. Because the surface of the antennae (and the surface of the apex of each serrate antennomere) is covered by sensilla it would seem reasonable to guess that this modification accomplishes increased surface area so that more sensilla can be "accumulated" there. Even the antennae of Hylecoetus (which are only slightly expanded) have sensilla over most of the surface of the flagellar segments (e.g., Fig. 18). 485. The only characters found on the mandibles were reductions. At times there is a single apical dens, rather than the plesiomorphic two. The prostheca is sometimes represented by a patch of setae, and is often missing entirely. Similarly, the molar area of the mandible has at times various small-sized spines and/or setae in transverse rows (relative to the longitudinal axis of the mandible), but these too are often lacking. Reduction characters of the maxillae (such as loss of distinct suture between dististipes and basistipes,§ suture between distigalea and basigalea), similarly, were not used. Derivative features are primarily associated with the bizarre maxillary palp-organs. These structures are present in all Lymexylidae ( 8) except Australymexylon, new genus, in the male, and also present in the female in the large, tropical genus Atractocerus (9). Whether the absence of these struct ures in Australymexylon is primitive or derived is a question of much intrigue and importance, and will be raised again under my discussion of monophyletic groups below. The structure of the palp-organ in Hylecoetus is fairly primitive. The organ arises from the apex of palpomere III, as does palpomere IV. The organ has one long and one short primary branch, each of which support two parallel rows of secondary branches. The net effect is a large structure which is basket-like in form. The outer sides of the "basket" are covered by fluted, tactile setae (Fig. 13) and the inner surfaces by long, thin-walled sensilla (Fig- 14). The tactile setae 486. are set in distinct sockets, while the sensilla are not. A similar (primary-secondary) branching pattern is found in male Melanomelittomma insulare. A few tertiary branches are seen in male Lymexylon navale (10); in this case, however, the surface of the tertiary branches is identical to that of the secondaries (this is not true in Atractocers: see below). In Atractocerus there is a regular and elaborate system of paired secondary branches, each one of which is lined with parallel rows (ie, paired) of tertiary branches ( 11) ; th ese tertiaries are covered by long, thin sensilla, and none of the shorter, tactile setae (present on the outer surfaces of the secondary branches) are present (the inner surfaces of the 2°-branches have similar sen silla ). Melittomma sericeum has a branching pattern similar to that just described for Atractocerus, but M. brasiliense has a very complex maxillary palp-organ which has as many as l°-5°-branches; all branches have tactile setae on the outer surface, and only sensilla on the inner su rfa c e . Several authors have thought that Atractocerus had only three maxillary palpomeres (e.g., King, 1955). This, however is not the case. Close examination of the palps clearly show palpomere IV to be present, albeit in a modified form. The sensilla at the apex of IV are shorter and broader in form than any seen on the palp-organ, and the typical, long setae characteristic of palpomere IV are also present (but not on surface of palp-organ). Why the confusion? Simply stated, the membranous area connecting the fourth palpomere to the third has 487. become enlarged, and has acquired the setae and sensilla of the palp- organ. The result is that palpomere IV appears to be part of the maxillary palp-organ. Figure 124 shows the maxilla of Atractocerus brevicornis (female). The base of palpomere IV looks the same as the branches of the maxillary palp-organ (Fig. 125), but its apex bears unusually long setae and typical, short sensilla (Fig. 127), and several recumbent sensilla (Fig. 126) none of which is ever seen on branches of the palp-organ (Fig. 128). The same situation is illustra ted by Atractocerus procerus (South America), where IV is perched upon the maxillary palp-organ (Fig. 174). Because these maxillary palp-organs have become so important in distinguishing the lymexylid beetles from other beetle families, I think it would be justifiable to re-examine some long-standing questions in light of my comparative studies. I am referring specifically to points raised by Germer (1912), Germer 8 Stech (1912), and Slifer et al (1975). Slifer et al (1975) provide an excellent, detailed morphological study of the structure of the male maxillary palp-organ of the North American species Melittomma sericeum, and many of their conclusions can, I believe, be applied to the lymexylids in general. They describe the soft, very lightly sclerotized integument which comprises the palp-organ. This is obviously (as they state) to allow expansion of the organ by fluid-pressure so as to expose the maximum amount of sensilla. Through careful sectioning techniques they clearly show that the stout, fluted setae on the outer surface of the organ 488. are tactile, and that the long, tapered but blunt, inner sensilla are thin-walled chemoreceptors which have minute pores (too small to see with light microscopy) covering their surface. I have not attempted to make sections and examine the innervation of the setae to determine their functions because it was beyond the set of techniques I had chosen to study these beetles and because no fresh material could be obtained for most taxa. I think it is very reasonable to infer similar functions for similar-appearing setae seen in all lymexylids. Slifer et al (1975: 126) said that "It would be interesting to know whether the thin-walled chemoreceptors on the antennae fof Hyle coetus flabellicom is) are similar to those found on the palp organs of H. dermestoides and the species studied here". To investigate their question I prepared an antenna from flabellicornis and dermestoi des (and also maxillae from each) for scanning electron microscopy. The results are seen in figures 13 and 14 (dermestoides) and figures 23 and 24 (flabellicornis), comparing palp-organ and antenna respective ly. Although different in shape to a degree, the outer and inner structures of the palp-organ of dermestoides are comparable to those studied for M. sericeum by Slifer and colleagues. The enlarged, flabe- llate antennae of flabellicornis, however, are quite different. Tactile setae are present on the outer surface, as are scales and short, thin- walled processes. The inner surface has only these processes, and not the very long chemoreceptors seen elsewhere. The function of these two thin-walled structures may be the same, but I do not believe that 489. they are homologous. Slifer et al (1975) also state that some female lymexylids have maxillary palp-organs while corresponding males do not. I am unaware of any instance when females have organs and males lack them. In all species of Atractocerus examined both males and females have maxillary palp-organs, and they are always more highly developed in males (the presence of MPO's in females is apomorphic for this genus ( 9 ) . A lso, they say that in no case are both antennae and palpi modified in a single form. This too is not true. In flabellicornis the maxillary palp-organ is present, but admittedly reduced greatly (Fig. 26). And, in the Australian species Atractocerus tasmaniensis the maxillary palp-organ is rather small and simple, but the antennae bear very large thin-walled sensilla (Figs. 154, 155). I have found no very useful characters on either the labrum or labium. There are some v a ria tio n s which w ill be o f im portance a t some level, and we can realistically comprehend a generalized plan for the structure of each of these for the Lymexylidae. The labrum is small, rounded, and separated from the clypeus by a broad membrane (except in Atractocerus spp., in which it is immovably fused to th e cranium ). The mentum i s u su a lly q u ite sm all, tra n s v e rs e , and quadrate. The palpigers are often widely separated, and almost appear to be extra basal palpomeres. The ligula is usually small, and transverse, sometimes with a medial emargination (e.g., M. sericeum, fig. 244). The most unusual modification of the ligula occurs in 490. some species of Atractocerus (e*g*» A. brevicomis) in which it is very large and elongate (Figs- 172-173). Other species of the same genus, however, have typical ligulae. The labial palpus varies in sizes of the palpomeres, but always has three palpomeres. The hypopharynx is usually rather small, membranous and covered by dense setae which may be long or short, and the margin sometimes has very long setae which are rarely forked. The epipharynx is covered by microtrichiae, and there are various patches of long setae, and almost invariably long marginal setae. THE PROTHORAX. The shape of the pronotum varies widely among the genera, and between species (and sometimes sexes of a single species) within many genera, from nearly quadrate and transverse in Hylecoetus to very elongate and narrow in some Melittomma-complex members. Hylecoetus, AustralymexyIon, and the Melittomma complex all have sharp lateral margins on the pronotum, while Lymexylon and Atractocerus lack this marginal feature (apomorph 12). The hypomeron of the pronotum sometimes has a distinct slot above the base of the procoxa ("supracoxal slot": apomorph 13), which is pre sent in the genera Australymexylon, Melanomelittomma, and some species of Melittomma (e.g., sericeum has well defined slots, while brasiliense lacks them; I interpret the lateral structure as secondary); Melittomm- opsis has a supracoxal emargination, but this is less well defined than the slots seen in the genera mentioned above. 491. The apex of the procoxa of Australymexylon and all genera of the Melittomma-complex has a narrow, elongate slot (see Fig. 273: apomorph 14); all other lymexylid taxa have the apex simple, with no evidence of a s lo t. THE LEGS. Although there are some special cases of modifications in leg structure (e.g., very elongate prolegs of Atractocerus tasman- iensis; see Fig. 170), these are atypical and most lymexylids have long, thin legs similar to those described for Hylecoetus under the morphology chapter. The tarsi are invariably 5-5-5, but the number of spurs at the apex of the tibiae vary. The basic number of tibial spurs, expressed as a tibial-spur-formula (just like the tarsal-formula), is c e rta in ly 2- 2-2 (this is seen in most families of beetles, including the cleroids § cucujoids). Reductions found in the lymexylids include the following combinations: 0-1-1 (male)/0-l-l (female) in Hylecoetus (15); 0-0-0 (male)/0-0-0 or 0-1-1 (female) in Atractocerus (16); 0-2-2 (male)/ 1-2-2 (female) in some Melittomma species (e.g., brasiliense)(17); and ?-?-? (male unknown)/ 1-2-2 (female) in Melittommopsis sp. THE MES0STERNUM AND ASSOCIATED SCLERITES. The mesosternum is primitively a quadrate (and narrowed anteriorly) to nearly triangular sclerite. In Australymexylon and the Melittomma-complex genera, a small, anterior area is elevated (18), but in M. brasiliense this raised area is missing. This elevated area is sometimes broadly triangular (M. sericeum), deeply emarginate posteriorly (M. insulare), or very long and narrow (Melittommopsis) (apomorphies 19 8 20, respectively). 492. The mesepistemum sometines has the anterior part elevated ( 21): Australymexylon and Melittomma-complex (except Melittomma brasiliense). This area is also sometimes separated from the posterior part of the mesepisternum by a sharp carina (22). The lateral portion of the mesepimeron is sometimes depressed (to deeply grooved) and receives the epipleural part of the elytra ( 23). This depressed area is sometimes separated from the remaining part of the mesepimeron by a sharp carina (24). Thoracic Terga. The primitive structure of the terga include simple alar ridges (Fig. 134), and a simple mesoscutellum (except in Hylecoetus which has a median scutellar keel: 25). In Atractocerus, the scutellum has lateral grooves into which the elytron fits (26; see Fig. 135) and the alar ridge is modified into an elevated process which fits under the elytra (Fig. 135; apomorph 27). Elytra. Shortened elytra occur in several lymexylid genera (Hyle coetus , Lymexylon, Australymexylon, and some species of Melittomma),but this phenomenon is nonpereil in Atractocerus which is brachelytrous (Figs. 117, 136), in which the elytra never surpass the metacoxae (and rarely approach them) (28). The paired elytra are usually united along the midline (the suture) by a tongue-and-groove locking mechanism (one elytron has a groove,into which the oposing elytron's narrow sutural flange fits). This mechanism is present in most lymexylids, but sometimes ends before the apex 493. (examples of this situation include Hylecoetus and some species of Melittomma]. This tongue-and-groove, however, is lost in Lymexylon and Atractocerus (29]. The Hindwing. The terms applied to the veins and cells of the hindwings of beetles are bewildering. I have not tried to make the terms used here rigorously correct in a strict anatomical sense, but rather I have stressed recognition of homologous structures in accord ance with other recent usage of terms (such as Doyen, 1966; Crowson, 1955; and others). The basic system is that of Forbes (1922), with several modifications. I have summarized the terms which I use, and compared them to other prominant systems in Table 4. The brachial cell is usually present (Fig. 46), but is absent in Atractocerus (30). Similarly, the wedge cell is absent in Atractocerus (31). The medial vein is usually present, and arises spuriously about half-way out the wing (Fig. 46); the M, however, is absent in Atracto cerus (32). The la-2a crossvein (which arises distally from the wedge cell) is incomplete (33) in both Lymexylon and Atractocerus. It is present, but broken in Lymexylon, and entirely absent in Atractocerus ( 38)• Vein 1A, which primitively arises spuriously, is fused with 2A in Atracto cerus (34). A spurious Rs is retained in Australymexylon and some Melittomma- complex taxa* but is absent in Hylecoetus, Lymexylon, and Atractocerus (35) . (*present in M. sericeum only) 494. Crossvein Cu-IA is sometimes (and apomorphically) lost (36) (in Atractocerus and some MeHttomma-complex taxa; present in M. sericeum). In Lymexylon, the male has an extra crossvein connecting 1A1 § 1A2 (Fig. 91; apomorph 37). In Atractocerus la-2a (crossvein) is lost (38), and the wings are folded longitudinally only (i.e., fanlike) (39). THE METENDOSTERNITE Crowson (1955; also, 1938, 1944) relied heavily on the structure of the metendosternite in the formation of his ideas about relationships among families of beetles. His broad base of comparative studies makes his judgements of polarities very important. Although he does not state polarities explicitly, a reading of his discussion soon reveals a number of his hypotheses. I have attempted to interpret these, and present my results in tabular form (Table 17). For further details, and examples, consult Crowson (1944, p. 305). Briefly, a primitive metendosternite might have the following description: stalk narrow*; anterior tendons nearly contiguous*, narrowly separated*; median projection present (= median laminal process in my figures 48, 49)*; no posterior sclerotization**; furcal arms pointed laterally*. (* denotes that Hylecoetoid forms have the primitive condition of this character; ** that they have the derived s ta te ) 495. My terminology varies somewhat from that used by Crowson (1938, 1944, 1955), and can be surmized from a perusal of my figures 47, 48, and 49. Crowson was not consistant in the terms he used, and I have tried to be a little more consistant here. I have found only a few characters of use to my study of the genera of Lymexylidae in the metendosternite, but a consideration of this structure is particularly important to further analysis of the relationship of lymexylids to other families of beetles. Especially in light of the historical importance of the metendosternite in higher studies of families this century. The furcal arms are laterally to (more strongly) antero-later- ally oriented. They are clearly, anteriorly directed in Atractocerus (40). The anterior tendons are very proximal in Hylecoetus (note summary of Crowson's interpretations of the polarities of metendostern ite structures in table 17), and (more) widely separated in other genera (41). In Australymexylon, the anterior tendons actually arise on the furcal arms (42), rather than between them. The laminae of the Melittomma-complex are very short (anterior to posterior) and wide (mesal to lateral) (43), in contrast to Australymexylon, Lymexylon, Hylecoetus, and Atractocerus structure, which involves longer and less-wide shapes. 496. THE ABDOMEN ’ The following characters refer to the visible part of the abdomen. The apical segments which are telescoped into the abdomen (along with the male/female genitalia} are discussed below as the "postabdomen", Hylecoetus is the most unusual of all lymexylid genera with regard to abdominal structure. Males have seven visible stem ites (XI-VIII), segment IX forming a genital capsule, and females have six visible sternites (II-VII) (see figures 50 § 511. In aH other genera segment VII is the apical one (VIII being telescoped inside}( 4 4 1 , and i n many taxa Ce.g., Australymexylon, most Melittomma~complex excluding Meli~ ttomma pervagus from Australial sternite II is greatly reduced or completely lost (45) making the abdomen appear 5-segmented (it appears, conversely, 6 -segmented in Atractocerus spp., Melittomma pervagus). The sexual dimorphism only occurs in Hylecoetus (46). THE FEMALE POSTABDOMEN AND GENITALIA The coxital fold (47) which divides each coxite into a basal part and distal part in believed to be autapomorphic for Lymexylidae. It is present in all taxa which I have seen, and I have not seen it in any other family of Coleoptera. The fungus pouch (see Fig. 53) is certainly a derived feature, but its distribution among the genera is not firmly established. In 497. Hylecoetus this structure is highly differentiated and obvious. But, in various other species in the family it may be present, but developed to lesser degrees (that is, there are not such distinct membranous folds). I suspect, however, that it is present in all taxa. A combi nation of ecological, ethological, and perhaps cross-sectioning tissue studies from fresh material will probably be required before a satisfactory answer is had to the question of the distribution of this fungal pouch. In Atractocerus there is sometimes a spurious baculus which is mesal and parallel to the valviferal baculus (48); and, at least the distal part of the coxite is thickened, and leathery (49), although the overall shape and configuration varies greatly. The proctigeral baculi are sometimes proximal or actually fused in the Melittomma complex (50): they are proximal, at least near the middle, in Melanomelittomma and Melittomma spp; they are fused (51) in some Melittomma sp e c ie s. 498. MALE POSTABDOMEN AND GENITALIA The unusual asymmetrical lateral lobes (52) and asymmetrical abdo minal segment IX (53) of Hylecoetus were discussed under morphology above. Australymexylon has large, setose pads on the sternite of abdominal segment VIII in the male (54), which have no parallel anywhere in the family (or elsewhere to my knowledge). Close examination of several preparations (up to x450) failed to show.any sign of secretory cells, but I had a limited number of specimens. Thus, both secretory and sensory functions seem to be viable guesses as to the use of these structures. Field investigations in Australia are badly needed to clear up a number of interesting points regarding Australymexylon species. (This is the only genus which lacks maxillary palp-organs in th e male) Species of Melittomma, sensu stricto, have the paratergites of abdominal segment IX appendiculate (55). These "appendages" range from short, broad forms (e.g., M. sericeum, fig. 248) to the more common long, narrow forms (e.g., M. brasiliense, figs. 232, 235). It is interesting that the lateral lobes of some species of Atractocerus are very elongate, and resemble these paratergites (e.g., A. procerus, figs. 164, 165, 166). 499. The lateral lobes are quite small in Hylecoetus, and in most other genera of the family. In Atractocerus, however, the lateral lobes are very much enlarged (56), always more conspicuous than the (relatively) small median lobe. In Australymexylon fuscipennis the median lobe is greatly reduced (57), the result being that the lateral lobes are (relatively) much larger than the median lobe. By contrast, the elongate median lobe of sister-species A. australe makes the lateral lobes seem much sm aller. Primitively, the tegminal struts are elongate, conspicuous struct ures which connect the tegminal plate (usually from about the base of the lateral lobes) to the base of the median lobe. The tegminal struts are, however, greatly reduced in Australymexylon spp. and(even more so in) certain species of Atractocerus (e.g., A. procerus , see fig. 164). The condition of hav ing reduced struts (58) is interpreted as apomorphic, but I would argue that it has been independently acquired in these two genera since other species of Atractocerus have relatively large tegminal struts. While the tegminal plate is typically simple (and often semi- membranous and not clearly deliminted), in certain Atractocerus it is strongly bifid and resembles a pair of lateral lobes (59) (see, e.g., fig. 167). Also, the tegminal plate is usually relatively large, but in the Melittomma-complex it is reduced in size (60) (see, e.g., fig. 233) . 500. The elongate, large lateral lobes of MelanomelittOmma Q51), with a row of stout spines medially near the apex, is unique among male genital structure in the family. STRUCTURES OF THE LARVAE Although a broader outgroup is needed, the membranous mesal margin of the larval maxilla which is bordered by long setae may be apomorphic for the family as a unit C62). The partial division of the mala which Crowson (1955) discusses is a plesiomorphy, and this division is subsequently lost in various taxa. Although ocelli are described by Grandi (1961) in neonatal larvae of Hylecoetus dermestoides, and I have seen ocelli in neonatal (i.e., 1st instar) larvae of both Atractocerus brasiliensis and Melittomma sericeum, the ocelli seem to be lost in late-instars of all lymexylid larvae. This loss of the ocelli may be an apomorphic feature of the Lymexylidae (63). This is an interesting development in light of what ethological information has been published. According to Francke- Grosmann (1967), the first-instar larvae of Hylecoetus wiggle about (brushing against their sibling larvae and egg-shells) to insure innoculation with spores and then seek out a place at which they bore into the wood. Once inside the arboreal host, there would be no apparent need for the ocelli. 501. The tarsungulus of the larval leg almost invariably has two setae in the beetles, and this is true for most lymexylid genera as well. Crowson (1955; 1964) pointed out that certain lymexylids have many setae on the tarsungulus. To be precise, Hylecoetus and Atractocerus have pleurisetose tarsunguli (64). It should be noted that the tarsungulus of Australymexylon is bisetose, but is covered by many short spines (65). The epipleural folds of the abdominal segments are characteristic of lymexylid larvae, but this structure is not clearly defined (i.e., some folds are more distinct than others) and it has not been compared with other families. The posterior end of the larva has the ninth segment greatly enlarged, and the tenth very small and inferior in position. The protruding (and therefore apical) ninth segment is commonplace among the cucujoids (see, e.g., illustrations in various papers on cucujoids by Hayashi listed in References below), and this is true also for cleroids (see illustrations in Crowson, 1964), but rarely approaching the extreme development seen in the lymexylids. In Hylecoetus, the ninth segment is very long, narrow, heavily sclerotized and sword-like in appearance ( 66 ). In Lymexylon and Atractocerus the ninth segment is membranous and bulbous in shape ("bustle-like") (67), and in most Atractocerus there are various configurations of asperites apically on this segment ( 68 ). 502. In Melittomma-complex taxa and Australymexylon the apex of segment nine is truncate and very heavily sclerotized (69). The border of the truncate ninth segment is composed of small to deeply excised teeth in Melittomma, s.s., and Me1anomelittomma. This is not true for Australymexylon. The teeth are apomorphic (70). Monophyletic Groups Autapomorphies, or uniquely derived characters, are cited in support of monophyletic status for each genus. Then, groups of genera which are believed to be monophyletic are justified by brief discussions of the synapomorphies which indicate these groupings. Genus Hylecoetus (autapomorphies): Soft-bodied (1)*; head with epicranial pit (5); antennae serrate ( 6 )*; tibial spur formula 0-1-1/ 0-1-1 (15); scutellum with sharp, medial keel (25); male with one more visible abdominal segment than female (i.e., female has viii retracted, male does not)(46); lateral lobes of male genitalia asymmetrical (52); abdominal segment IX of male asymmetrical (53); tarsungulus of larva pleurisetose (64); larval abdominal segment IX very long, narrow and heavily sclerotized ( 66 ). Genus Lymexylon (autapomorphies): hindwing of male with extra lal-la2 crossvein (37); soft-bodied (1)*; head narrowed behind eyes, forming "neck" ( 2)*; maxillary palp-organ of males with at least a few tertiary branches (10)*; lateral lobes very short, broad, and truncate (separated medially by wide gap)(71); apex of larval abdomi nal segment IX with tooth-like process (Fig. 113)(72); ventral surface of labrum (larva) with three lateral-setae (Fig. 114)(73). *These characters have analogous structures in other genera. See polarity discussion. 504. Genus Atractocerus (autapomorphies): Soft-bodied (1)*; head narrowed behind eyes (2)*(in some spp.); eyes large and emarginate (3) (4); antenna short, spindle-shaped (7); maxillary palp-organ present in female (9); maxillary palp-organ of male with tertiary branches (10)*, tertiaries arranged in paired, parallel rows (11)*; tibial spur formula 0-0-0/ 0-0-0 or 0-1-1 (16); scutellum with lateral grooves which receive elytra (26); alar ridge modified into elevated process (27); brachelytrous (28); brachial cell absent (30); wedge cell absent (31); 1A fused with 2A (34); la-2a crossvein absent (38); wings fold longitudinally only (39); furcal arms strongly anteriorly-oriented (40) (metendosternite character); spurious baculus present near coxital baculus (48); coxite leathery (at least distally) (49); lateral lobes greatly enlarged (56); tegminal plate bifid (59); tarsungulus pleuri setose (64)*; lateral lobes with 2 or 3 teeth on mesal surface (74). Genus Australymexylon (autapomorphies): head narrowed behind form ing neck (2)*; eyes emarginate (4)*; antennae serrate (6)*; maxillary palp-organ absent in male (75); male abdominal sternite VIII with large, setose pads (54); median lobe (sometimes) reduced in size (57); tegminal struts reduced in size (58)*; tarsungulus of larva bisetose, but with many small spines (65); anterior tendons of the metendostem- ite arise on the furcal arms (42). Genus Melittomma (autapomorphies): paratergites of abdominal segment IX long, appendiculate (55); tegminal plate short, small (60); 505. rim of larval abdominal segment IX with teeth (70). Genus Melittommopsis (autapomorphies): apical article of maxi llary palpus securiform (female)(76); apical palpomere of female labial palpus securiform (77); anterior area of mesostemum long and narrow (78); distal part of coxite membranous and narrow, basal band sclero tized (79); apical h of stylus membranous and setose, basal h s c le ro tized and glabrous (80). Genus Melanomelittomma (autapomorphies): anterior part of meso- sternum deeply emarginate (19); lateral lobes very long, large and with mesal (apical) row of spines (61); apex of larval abdominal segment IX crenulate (83); distal part of coxite of female genitalia highly-differentiated from basal part, heavily sclerotized (81). Subfamily Hylecoetinae (Genera included; Hylecoetus). As I discussed above, the Hylecoetinae are separated by a wide gap from all other lymexylids. The characters cited above under Hylecoetus apply h ere. Subfamily Lymexylinae, sensu stricto (Genera included; Lymexylon, Atractocerus). My use of Lymexylinae is new and restricted. Both Boving § Craighead and Crowson include genera of my "Melittoraminae" here as well. The following synapomorphies support the Lymexylinae as a monophyletic unit; maxillary palp-organ with tertiary branches (10); pronotum without sharp, lateral margins (12); tongue 8 groove locking mechanism of elytra absent (29); crossvein la-2a of hindwing incomplete (33); larval abdominal segment IX large, membranous, bulbous and "bustle-like" (67) ; larval abdominal segment IX with asperites (68). Subfamily Melittomminae (Genera included: Australymexylon, new genus; Melittomma, sensu stricto; Melittommopsis; and Me1anome1ittomma, new genus). Serrate antennae (6), although also present in Hylecoetus, seem synapomorphic for the subfamily; hypomeron of pronotum with supracoxal slot (13); procoxa with slot (14); anterior part of mesepi- sternum elevated (21) (this is apparently lost in Melittomma brasil iense ) ; mesepimeron with depressed lateral area which receives the elytral epipleural fold (23); larval abdominal segment IX round, truncate, heavily sclerotized (69). Melittomma-complex of genera CGenera included: Melittomma, Melanomelittomma, and Melittommopsis). The anterior area of the meso- sternum is elevated (18) (Melittomma brasiliense is an exception); lamina of metendosternite short and broad (43). What are the relationships among the genera in this complex? The elongate, appendiculate paratergites of male abdominal segment IX (55) and the small tegminal plate of the male genitalia (60) support the monophyly of Melittomma, sensu stricto, and the carinal margin of the mesepimeral depression (24) supports sister-group relationship for Melittomma and Melanomelittomma, as does the proximal position (at least near the middle) of the proctigeral baculi of the female geni talia (50). Melittommopsis, then, is indicated as sister group of Melittomma-Melanomelittomma. Classification of Lymexylidae My classification of the lymexylids is summarized in Table 18, and is expressed as a phyletic sequence scheme (see Wheeler, 1979a). This classification marks a refinement over its predecesors. I use three subfamilies (previous authors have used two since 1931), and propose two new genera to efficiently handle the observed diversity. In the phyletic sequence each taxon is the sister group of all taxa of equal or lesser rank listed below it. Thus, Hylecoetinae is sister to Lymexylinae + Melittomminae, Lymexylinae sister to Melittomminae, Australymexylon sister to the Melittomma-complex, and so forth. 508. Speculations on the Origin of Lymexyloidea What are the evolutionary origins of the lymexylid beetles? In b r ie f , I do not know. What I would lik e to do h ere, however, is to summarize the prevailing hypothesis regarding origins of lymexyids and to speculate cursorily based on my studies of these beetles. All of the evidence is not yet in (the largest gap being lack of detailed studies of the comparative structure of Cucujoidea families), but some educated guesses can serve to stimulate the process of gathering this d ata. Roy Crowson (1955, 1960, 1964) has made the most significant contributions towards a comprehensive understanding of the higher relationships of the Coleoptera, including the position of Lymexyloidea. In 1955, Crowson suggested that the Lymexyloidea might be sister-group to Cleroidea/Cucujoidea. His most recent scenario, however, pictures these superfamilies as an unresolved trichotomy (see my figure 292). Specifically, Crowson (1960) envisions a mid-Jurassic trichotomy. Lymexyloidea is wood-boring and mycophagous, Cleroidea is carnivorous, and Cucujoidea continues the plesiomorphic ecological relationship with dead wood/fungi (i.e., under bark, in soft-wood, etc., and asso ciated with mycelia and sporophores. Crowson points out an intrigueing parallel between this trichotomy (and tri-ecological radiation) and an earlier one in the evolution of beetles. This preceeding radiation 509. included Polyphaga (decaying wood-fungi), Myxophaga (wood-boring), and Adephaga (predaceous). In my estimation, the lymexylids are quite similar to the basic plan for cucujoid beetles (see Table 19), and sim ilarities to the cleroids are superficial and very likely resultant of convergence. I lack detailed information for the cucujoids, and can therefore not formulate an empirical argument for this view. I strongly urge detailed comparative morphological studies of Heteromera and Clavicornia. Lawrence (1977), Watt (1974), and others are already hard at work on the heteromeran problem, but the clavicorns remain largely ignored. Persistance in pursual of this line of inquiry will hopefull permit resolution of this fascinating question of superfamilial relationships in the foreseeable future. Table 16. Character state polarities. No. Apomorphic s ta te ______Plesiomorphic state 1 Soft-bodied (lightly sclerotized) Heavily sclerotized 2 Head narrowed behind eyes forming distinct Head broadly rounded behind eyes, not narrowed to "neck" (Figs. 218, 257) neck (Figs. 2, 118) 3 Eyes large, separated by less than single Eyes small, separated by single eye width (or more) eye width (Figs. 120, 218) (Figs. 2, 82, 179) 4 Eyes emarginate anteriorly (Figs.218,257) Eyes entire (Fig. 2) 5 Epicranial pit present (Figs. 2, 30) Epicranial pit absent 6 Antennae serrate (Figs. 180-183) Antennae filiform (Fig. 222) 7 Antennae short, pointed, spindle-shaped Antennae elongate, narrow (F ig. 123) 8 Maxillary palp-organ present in male, Maxillary palp-organ not present Arising from apex of palpomere III (F ig .7) 9 Maxillary palp-organ present in female, Maxillary palp-organ present in males only but smaller than in male (Fig. 124) 10 Maxillary palp-organ with tertiary Maxillary palp-organ with secondary branches branches (Figs. 97, 130) (males) only (Fig. 7) in Table 16. Character state polarities (cont'd.). No. Apomorphic s ta te ______Plesiomorphic state 11 Maxillary palp-organ with tertiary branches MPO without parallel tertiary branches arranged in paired, parallel rows (Fig. ) 12 Pronotum lacking lateral margin Pronotum with sharp lateral margin 13 Hypomeron o f pronotum w ith s lo t (Fig* 206) Hypomeron sim ple 14 Procoxae apically slotted (Fig. 273) Procoxae apically simple 15 Tibial spur formula 0-1-1/0-1-1 (male/female) TSF 2-2-212-2-2 16 Tibial spur formula 0-0-0/0-0-0 or 0-1-1 TSF 2-2-212-2-2 17 Tibial spur formula 0-2-2/1-2-2 TSF 2-2-212-2-2 18 Mesosternum elevated anteriorly •Mesosternum simple (Fig. 36) 19 Elevated, anterior part of mesosternum deeply Elevated area shallowly emarginate posteriorly emarginate posteriorly 20 Elevated, anterior part of mesosternum long, Elevated area broad, not long, narrow, pointed narrow, pointed posteriorly (Fig. 283) 21 Anterior part of mesepisternum elevated Mesepisternum not elevated anteriorly (Fig* 207) Table 16. Character state polarities (cont'd.). No. Apomorphic s ta te ______Plesiomorphic state ______ 22 Elevated, anterior part of mesepisternum Carina lacking separated from posterior part by carina 23 Mesepimeron with depressed, lateral area Mesepimeron Without depression which receives elytral epimeron 24 Mesepimeral depression separated by carina Carina lacking 25 Scutellum with median keel (Fig- 37) Scutellum simple, or with lateral areas raised 26 Scutellum with lateral grooves which Scutellum simple receive elytra 27 Alar ridge modified into elevated process Alar ridge simple (Fig. 134) (Fig. 135) 28 Brachelytrous (elytra not reaching metacoxae) Elytra long or slightly shortened 29 Tongue-and-groove absent along elytral suture Tongue-and-groove present (although it may end before apex) 30 Brachial cell absent (Fig. 156) Brachial cell present (Fig. 46) 31 Wedge cell absent (Fig. 156) Wedge cell present (Fig. 46) 32 Spurious base of M absent (Fig. 156) Spurious base of M present (Fig. 46) 512. 33 la-2a crossvein incomplete (Figs. 91,156) la-2a crossvein complete (Fig. 46) Table 16. Character state polarities (Cont'd.). No. Apomorphic s ta te ______Plesiomorphic state 34 1A fused with 2A (Fig. 156) 1A separate from 2A, arising spuriously 35 Spurious Rs absent Spurious Rs present 36 c u -la cro ssv ein absent cu-la crossvein present 37 Extra crossvein between 1A1-1A2 present Single 1A1-1A2 crossvein present (Fig. 91) 38 la-2a crossvein lost (Fig. 156) la-2a crossvein present 39 Wing folded longitudinally only (fan-like) Wing folded transversely (and longitudinally) 40 Furcal arms anteriorly directed (Figs, 176, Furcal arms antero-laterally directed (Figs. 47, 177, 178) 48, 49) 41 Anterior tendons separated (Figs, 262, 94) Anterior tendons proximal, nearly contiguous (Figs. 48, 49) 42 Anterior tendons arise on furcal arm (Fig.186) Anterior tendons arise between arms (Fig. 262} 43 Laminae short, wide (Figs, 236, 262) Laminae longer, narrower (Figs. 48, 186) 44 Abdominal segment VIII telescoped into VII Abdominal segment VIII exposed 45 Abdominal segment II absent (or nearly so) Abdominal segment II present 513 46 Male with one more exposed abdominal segment Male 8 female w ith same number o f segments Table 16. Character state polarities (ContM,), No. Apomorphic s ta te ______Plesiomorphic state ______ 47 Coxital fold present, dividing coxite into Coxital fold absent, coxite not divided basal part and distal part CFigs. 53-55) (on ventral surface only) 48 Spurious baculus present (Fig, 144,2) Spurious baculus absent 49 Coxite (distallyl thickened, leathery Coxite membranous 50 Proctigeral baculi proximal, at least near Proctigeral baculi separate and parallel middle 51 Proctigeral baculi fused along part of their Proctigeral baculi not fused, separate length (Figs. 274, 275) 52 Lateral lobes asymmetrical (Fig, 59) Lateral lobes symmetrical (Fig. 233) 53 Abdominal segment IX asymmetrical (Fig, 58) .Segment IX symmetrical 54 Abdominal sternite VIII with large, setose Abdominal sternite VIII without pads pads CFigs. 191, 192) 55 Paratergites of abdominal segment IX long, Paratergites of IX small, not appendiculate appendiculate (Figs, 235, 248) 56 Lateral lobes greatly enlarged (Figs, 145,165) Lateral lobes not greatly enlarged 514 57 Median lobe reduced in size (Fig, 1&7) Median lobe relative large (pig. 2QQ) Table 16. Character state polarities (Cont'd.). No. Apomorphic s ta te ______Plesiomorphic state 58 Tegminal struts reduced (Figs. 164, 197) Tegminal struts long (Figs. 57, 231) 59 Tegminal plate bifid (Fig. 167) Tegminal plate simple, not bifid 60 Tegminal plate small (Fig. 233) Tegminal plate large (Fig. 57) 61 Lateral lobes very long, stout, with Lateral lobes much smaller, simple, lobate apical (mesal) row of spines (Fig. 258) 62 Mesal margin of maxilla (of larva) membranous, Mesal margin not as described and bordered by long setae 63 Ocelli absent in late-instar larvae Ocelli present 64 Tarsungulus pleurisetose (Figs. 62, 147.2) Tarsungulus bisetose (Fig. 253) 65 Tarsungulus covered by short spines (Fig. Tarsungulus without spines (but bisetose) 203), and bisetose 66 Abdominal segment IX (larva) very long, Abdominal segment IX shorter, broader in form narrow, heavily sclerotized, with dorsal teeth, "sword-like" (Fig- 61) 67 Abdominal segment IX (larva) very large, Not as described bulbous, semi-membranous (Figs. 112, 148) 515 68 Apex o f IX w ith a s p e rite s (Fig* 148) Apex o f IX w ithout a s p e rite s Table 16. Character state polarities (Cont'd.). No. Apomorphic s ta te ______Plesiomorphic state 69 Apex of abdominal segment IX (larva) round, Not as described truncate and heavily sclerotized (Fig.278) 70 Rim of abdominal segment IX with teeth Rim of IX without teeth (Fig. 278) (Fig. 279) 71 Lateral lobes short, broad, truncate and Not as described widely separated (Fig. 99) 72 Apex o f abdominal segment IX (la rv a ) w ith Apex sim ple, w ith a s p e rite s but not to o th tooth-like process (Figs. 112, 113) 73 Labrum w ith th re e la te r a l se ta e (Fig. 114) Labrum with two setae (Fig. 65) 74 Lateral lobes with 2 or 3 large "teeth" Lateral lobes without large "teeth" mesally (Fig. 145) 75 Maxillary palp-organ lost (Fig* 210) Maxillary palp-organ present 76 Maxillary palpus of female securiform Maxillary palpus with apical segment parallel (Fig. 280) (apical segment) sided, or nearly so 77 Labial palpus of female with apical segment Labial palpus of female with apical segment securiform (Fig. 282) parallel-sided or nearly so 516 78 Anterior area of mesosternum long, narrow Anterior area broader (F ig. 283) Table 16. Character state polarities (Cont'd,). No. Apomorphic state ______Plesiomorphic state 79 Distal part of coxite membranous, except Distal part of coxite uniformly sclerotized for narrow, sclerotized basal area (Pig. (sometimes semi-membranous) 284) 80 Apical % of stylus membranous, setose and Stylus of uniform texture b asal h sclerotized, glabrous (Fig. 285) 81 Distal part of coxite very heavily sclero Distal part of coxite less distinctly separated tized, nearly separated from basal part by from basal part, not very heavily sclerotized coxital fold (Figs. 287, 288) 82 Apex o f male abdominal s te r n ite V III Apex o f s te r n ite V III sim ple, rounded produced (Fig, 286) 83 Apex o f la rv a l abdominal segment IX Apex not c ren u late crenulate (Fig. 278) 84 Lateral lobe with internal receptacle (Fig. Lateral lobe simple, without receptacle. 245) Table 17. My interpretations of plesiomorphic and apomorphic states of characters of the metendo- sternite of Coleoptera, based on data and discussion of Crowson, 1944: 305. Apomorphic state Plesiomorphic state - Stalk narrow* Anterior tendons separate Anterior tendons nearly contiguous* Anterior tendons widely separated Anterior tendons narrowly separated* - Median,projection present, bearing anterior tendons* - Complete absence of a posterior sclerotization** • Sternal suture present well before hind margin of metasternum (primitively connecting with ventral process of furca)+ ** Presence of furcal arms** - Furcal arms pointed antero-laterally Furcal arms pointed laterally* Presence of (well) differentiated lateral parts of the ventral process (in forms derived from the Hylecoetoid-type) *Hylecoetoid-type plesiomorphic for these characters; **apomorphic for these; see also my figs.47-49. 519. Table 18. Phyletic Sequence Classification of Lymexylidae. Subfamily HYLECOETINAE Genus Hylecoetus Latreille Subfamily LYMEXYLINAE Genus Lymexylon Fabricius Genus Atractocerus Palisot de Beauvois Subfamily MELITTOMMINAE Genus Australymexylon Wheeler, new genus Genus-Complex Melittomma Genus Melittommopsis Lane Genus Melittomma Murray, sensu stricto Genus Melanomelittomma Wheeler, new genus Fig. 291. Rate of description of Lymexylidae. Black dots- nominal species; open circles- nominal genera; triangles- species listed in most recent lists (including Zoological Record) some of which are known, but unresolved, synonyms. See text for discussion. < 9 0 , < 8 5 H80 sppP 0 7 5 a: 7 0 Ld CD 65 3 6 0 . Z 55 5 0 4 5 4 0 35. 30 2 5 2 0 15 1 0 -O' j O- YEAR:1740 1760 1780 1800 1820 1840 1860 1880 1900 1920 1940 1960 1980 Fig. 292. My concept (and illustration) of Crowson's scenario of the evolution of Lymexyloidea, Cleroidea, and Cucujoidea (Crowson, 1960). See text for discussion. Abbreviation: MYBP (millions of years before present.). 522. Period CLEROiratYMEXYlQDS CUCUJOIDS MYBP QUATERNARY TERTIARY 7 0 JURASSIC 1 8 0 TRIASSIC polyphagan stock Vi! dead-trce/mycophagous 2 2 5 v;i: habitat/feeding CROWSON’S EVOLUTIONARY SCENARD (1 9 6 0 ) K> Fig. 293. Cladogram of relationships among the genera of Lymexylidae. Numbers r e f e r to c h a ra c te rs (apomorphic s ta te s ) in ta b le 16. 524. 525. 11 55 17 60 40 22 70 42 86 81 35 82 77 36 84 46 66 43 13 23 47 Fig. 294. Cladogram and character matrix for Melittomminae. Numbers refer to apomorphic states in table 16. Abbreviations: AUSTV Austra- lymexylon; MELI- Melittomma; MELA- Melanomelittomma; MBRA- Melittomma brasiliense (rePresentative soft-bodied Melittomma); MSER- M. sericeum; MOPS- Melittommopsis. M E L I AUST MELA MBRA MSFR MOPF £ 0 K > 294 Table 19. Characters of ancestral Cucujoidea proposed by R.A. Crowson (1955). No. C haracter______Type of char. 1 Front coxae transverse, with exposed trochantins (e.g., Hypocoprus, Rhizophagus) Adults 2 Middle coxae with exposed trochantins Adults 3 Mesepimeron reaching middle coxal cavity " 4 5 free, visible sternites (abdominal) " 5 7 pairs of abdominal spiracles " 6 Antennae filiform or weakly clubbed, inserted laterally in front of eyes " 7 Mandibles with mola and prostheca " 8 Maxilla with 2 lobes (galea and lacinia) " 9 Gular sutures distinct and separate " 10 All tarsi 5-segmented (at least in female) " 11 Metendosternite of Hylecoetoid-type, anterior tendons close together " 12 Hindwings with complete Cantharoid venation (with radial.cell, anal cells, and " 5 anal veins in main group; possibly also, a spur on the r-m crossvein as in Chrysomelidae) 528 13 Elytra completely covering abdomen; more or less striate; with scutellary striole" Table 19. Characters of ancestral Cucujoidea proposed by R.A, Crowson (1955).(Cont'd.}. No. C haracter Type 14 Aedeagus of cucujoid type Adults 15 6 cryptonephridic Malpighian tubules II 16 Hind coxae strongly transverse, and nearly contiguous II 1 Head without median epicranial suture; with characteristically shaped frontal Larvae sutures (see Crowson, 1955, figure 122} 2 Distinct fronto-clypeal suture II 3 Labrum fre e If 4 Antennae 3-segmented It 5 Mandibles with mola and prostheca II 6 Maxilla with well-developed, articulating area, mala probably long and acute, II palpi 3-segmented (excluding palpiger) 7 Labial p a lp i 2-segmented, mentum 8 submentum c le a rly marked o ff It 8 Thoracic and abdominal segments with well-marked sclerotized tergites II U1 N> 9 Legs of normal Polyphagan type II tO Table 19. Characters of ancestral Cucujoidea proposed by R.A. Crowson (1955) (Cont'd.). No. C haracter Type 1 0 9 pairs of functional spiracles Larvae It 1 1 Postero-dorsal sclerotized out-growths (urogomphi, cerci) present on segment IX (abdominal tergite) 1 Loose bark and.dead trees (corresponding to earlier forms - i.e., upper Permian Proto- Habitat coleopterans and early Triassic Protopolyphagans, but with better drought-resistance. ZOOGEOGRAPHY In the absence of both species revisions and compilation of the known geographic localities involved in the individual species d istri butions, broad analysis of the extant pattern and in-depth historical scenarios are difficult to defend on empirical grounds. Nonetheless, my cladistic analysis of the genera provides insights into relationships among the genera, and correspondingly, relationships among the areas occupied by the genera. Not withstanding the absence of the species- level data, general distributions at the generic level can be surmized. It is from this cursory body of information that I develope my zoogeo- graphic presentation below. The Vicariant Model: An Introduction During the past decade a significant, new approach to biogeographic analysis has been taking form, emerging from the earlier writings of Leon Croizat (e.g., Croizat, 1958; his writings total more than 10,000 pages!), and being labelled "Vicariant Biogeography". Some especially important discussions of vicariance include Platnick 8 Nelson (1978), Rosen (1978), and Croizat et al (1974). A great deal of enthusiasm and energy has been unproductively spent in a heated argument over the comparative virtues of the dispersal 531. 532. model (the classical approach, encompassing variations of Darlington, 1958 and Pfennig, 1966; Brundin, 1966, 1972 advocates Hennig^s views) and the vicariant model. It should he obvious to most biogeographers that both dispersal and vicariant mechanisms have always been at work. Both models, in their usual form, are based on the allopatric speciation mode as advocated by Mayr CL969] and innumerable contemporary biologists. What then separates the dispersalist from the vicariance biogeographer? In brief, the difference lies in the temporal implemen tation of the isolating barrier, the barrier to "dispersal”. If a widespread population or species has its range divided into two or more smaller sub-units, the resultant, isolated populations (or, given enough time, species] have come about through vicariance. If, on the other hand, a barrier already' exists and some individuals of a population (or species) succeeds in crossing the barrier and founding a new population (and perhaps subsequently, species] on the other side then its origin has come about by dispersal in the classical sense. As discussed by Platnick § Nelson (1978], the vicariant model is based on the cladistic relationships of the areas (inferred from the cladistic analysis of the taxa). Once an initial pattern is established from the data in a revisional study, the pattern is compared to that seen in other organisms and a common, shared pattern or "generalized track" (Croizat et al, 1974) is searched for. When a number of taxa support one pattern, we can feel more confidence in our picture of the relationships between these geographic areas. The underlying concept behind the vicariance model is that if a geographic area was subdivided by some barrier, then many taxa would be similarly isolated. When we examine the distriubtions of other taxa (with similar capacities for movement and similar ecological tolerances), they may corroborate our initial pattern (indicating a common history of isolation) or they may not. Because it is unlikely that a barrier sufficient enough to isolate our initial taxa would not affect any other species (or that all other affected species have become extinct) we can look upon unique patterns as the result of dispersal in the classical sense. If we fail, however, to entertain a vicariant explana tion first, we do not have a chance to test our explanation. Distribution Patterns for Lymexylidae: The Data and Some Comments Below, I present a brief discussion of the distribution pattern which is perceived for each genus of Lymexylidae: 1. HYLECOETUS (HOLARCTIC) - This genus occurs w idely in E urasia (Europe, Scandanavia, USSR, China, Japan), and in northeastern North America. One species in each major area. 2. LYMEXYLON (PALEARCTIC) - There is (I b e liev e) a sin g le species which has a pattern similar to the Old World species of Hyle- coetus: Europe north to scandanavian countries, USSR, China, and Japan. 3. ATRACTOCERUS (PANTROPICAL) - Many sp e c ie s. Mexico south to 534. Brazil, tropical Africa, Madagascar, India, Southeast Asia, Australia. 4. AUSTRALYMEXYLON (AUSTRALIA) - This genus is endemic in e a ste rn Australia, and contains only two species. 5. MELITTOMMOPSIS (SOUTH AMERICA) - This genus in clu d es th re e described species, all from Brazil. 6. MELANOMELITTOMMA (MADAGASCAR,SUMATRA) - This genus includes the coconut-pest M. insulare and an undescribed species (known only from female specimen, and tentatively placed) from Sumatra. M. insulare occurs in Madagascar and the Seychelles. Based on the discussion given by Brown (1956), it seems probable that the species is native to Madagascar and introduced to the Seychelles (it has no known native host in the Seychelles, and only occurs on a few islands although others within apparantly attainable distances and with suitable hosts are not colonized). Thus, for the purpose of bioggographic discussion I am treating insulare as a Madagascar species. 7. MELITTOMMA (PANTROPICAL, fo r th e m o stp a rt)- This i s a la rg e genus which is almost entirely Pantropical, occuring in the New World from Mexico south to Brazil, tropical Africa, Madagascar, India, South east Asia, Australia, and a single species occurs in the United States and southern Canada. Speciation Patterns: A Subjective Analysis I believe that an important part of any biogeographic analysis is an appraisal of the speciation pattern. Some general clues for differentiating islandic and continental patterns were formulated by Donald Whitehead (1972, 1976) and summarized by Wheeler (1979b). The lymexylids seem to be obvious examples of continental speciation Single species are incredibly vagile, and have remarkably wide ranges. Some examples include Atractocerus brasiliensis which ranges from northern Mexico south to Brazil (and which has been reported flying aboard ships as far as 50 miles from shore); A. brevicornis which is transcontinental in Africa (and also occurs in Madagascar); Melittomma javanicum which occurs from India all through Southeast Asia (a distribution which includes the Philippines, Malay Penninsula, Sumatra, Borneo, New Guinea, Java, etc.). On a global basis, the general image is one of very few species, each of which enjoys a very wide range. One of the implications of the continental pattern is a slow rate of speciation. Historical Zoogeography: A Preliminary Analysis The geographic distributions of the genera are summarized in figure 295 in a geographic-area-matrix. This pattern is presented in a simplified geological-cladogram in figure 296. The series of indicated ancestral ranges, following the branching sequence of the cladogram (Fig. 296) are shown in figure 297. 536. The pattern is complex, and the single thread which is wound throughout the pattern is one of Pantropical distribution. Melittomma species occur in South, Central § North Americas, Africa, Madagascar, India, S.E.Asia, and Australia; and Atractocerus shows a similarly wide ranging distribution pattern. All of this points to a very old age for the group, perhaps implying that the family pre-dated the break-up of Pangea (about 180 MYBP) (Hylecoetus/Melittomminae-Lymexy- linae split could conceivably line up with this event); and some genera, including Melittomma and Atractocerus were probably extant before the break-up of Gondwanaland. Fig. 295. Distribution matrix for genera of Lymexylidae. 537. 538 TAXON HYLECOE LYMEXYL ATRACT AUSTRAL M-OPSIS MELANOM MELITT iURASIA C.AMER S.AMER AFRICA MADIGAS INDIA 5.E.ASIA \USTRAL Figure 295 Fig. 296. Geocladogram showing relationships among geographic areas indicated by biocladogram of genera of Lymexylidae. Reduced from figure 295. HOLARCTIC PALEARCTIC PANTROPICAL § i AUSTRALIA N> KD O' SOUTH AMERICA MADAGASCAR PANTROPICAL 540 Table 20. Series of ancestral distributions indicated by cladogram (Fig. 296). HOLARCTIC/ PALEARCTIC + AUSTRALIA + S. AMERICA + AFRICA + MADAGASCAR + SOUTHEAST ASIA + N § C AMERICA PALEARCTIC + "PANTROPICAL" / "PANTROPICAL" PALEARCTIC/PANTROPICAL AUSTRALIA / AUSTRALIA + AFRICA + MADAGASCAR + S. AMERICA + N.$C. AMERICA + SE ASIA SOUTH AMER./ MADAGASCAR + AFRICA + S .,C .§ N. AMERICA + SE ASIA + AUSTRALIA MADAGASCAR/ AFRICA + MADAGASCAR + S .,C .§ N. AMERICA + SE ASIA + AUSTRALIA APPENDIX A. M is c e lla n e o u s O b s e r v a tio n s on L y m ex y lid a e During the course of this study a number of significant observa tions were made on Lymexylidae which, due to their fragmentary nature, were not entirely appropriate for the textual format. These notes add important, supplemental data to various portions of the text, and are referred to in appropriate places. Diverse elements of this appendix include justifications for certain nomenclatural and taxo nomic changes, annotations about genera and species-complexes, biolo gical notes, comments on type specimens, and some interpretive, historical notes. The order of inclusion of these separate items is totally arbitrary; each entry is cited in the text (.see Appendix A: Item 1; etc.). (1) Australymexylon fuscipennis (.Lea), NEW COMBINATION . There are 2 specimens in the type-series of Hylecoetus fuscipennis (1 male,l fe male) and both are mounted on a single card. The male is small (about 7.5 mm, with the head strongly deflexed), of typ ica l coloration Ci.e., with a distinct pale area at the bases of the elytra: see description in text); the antennae are missing, although a piece of one antenna is glued to the card; the antennomeres are greatly expanded (corro borating this species identification). The female is about 12.9 mm long, the head is slightly deflexed; the ovipositor is extended, and is about 7.8mm long. 542. 543. Lea (1911, p.466) stated that "The ovipositor of the type is extended..." (my italics). This qualifies the female specimen as the holotype by original designation, according to article 73(b) of the International Code of Zoological Nomenclature as cited by Blackwelder, 1967 (p. 593). Accordingly, I have placed labels on the pin of the specimens indicating that the female is holotype, and the male is paratype. This is in contradiction to a handwritten "TY" label under the male specimen, presumably inscribed by Lea himself. The following labels are on the pin of the specimens: "5332, Hylecoetus fuscipennis Lea, N.S. Wales, TYPE"/"S.A.Museum". The type material was borrowed from the South Australian Museum for stu d y . (2) Hylecoetus lin e a r is Lea, 1894 ; p . 603 = Australymexylon australe, NEW SYNONYMY. Lea's description was based on a single specimen from Tamworth, New South Wales. I have borrowed and examined the type specimen (South Australian Museum) and make the following observations. The subserrate antennae indicate that it is a male specimen of au strale. The specimen, unfortunately, is in poor condition: all legs except one front leg, the entire abdomen, the apical antennomeres (4 on one side, 5 on the other), and the left elytron are all missing, and the speci men was pinned with a hefty sized minuten which has badly corroded. The head, pronotum, and antennae are reddish-brown, the elytron and ventral surface paler and yellowish colored. I am confident that 544. the specimen belongs to au strale. Labels on the type read : "linearis Lea, TYPE, Tamworth"/"5331 Hylecoetus linearis Lea, N.S. Wales, TYPE"/ "S.A. Museum Specimen". (3) Lymexylon adelaidae Blackburn, 1898, p . 34, Trans. Royal Soc. South Australia, = Australymexylon australe, n. comb., NEW SYNONYMY. The type specmen was borrowe from the South Australian Museum for study. The specimen is unfortunately in poor condition: the abdomen, antennae, one hind leg, one front tarsus, and one hind tarsus are missing. A piece of one antenna is glued to the top of the head and includes five apical segments, which indicate that the specimen is a female. The type is labelled : "Lymexylon, adelaidae Blackb., TYPE"/ (this label is handwritten and difficult to make out) "Thiro C., 4.3. 99, Gamia (+ another word I cannot read)"/"(a letter, perhaps 'J' with the numbers '6736'), Lymexylon, adelaidae Bl, S. Australia, Syn. of, A. australis Er."/"S.A. Museum Specimen". (4) Hylecoetus vigilans Lea, 1911, p. 467, Proc. Linn. Soc. New South Wales, see also Lea's fig. 8, on plate xvii. I have seen the single, female type (borrowed from the South Australian Museum) and am confident that it is conspecific with Melittomma pervagus O llif f , NEW COMBINATION; there is nothing out of the ordinary about the specimen. Labels: "vigilans, Lea TYPE, L ittle Mulgrave R."/"15040, Hylecoetus, vigilans Lea, Queensland, syn. of pervagus O il., TYPE"/ "S.A.Museum Specimen". 545. (5) Atractocerus procerus Schenkling, 1914, p. 319. LECTOTYPE: here designated, female specimen (DEMB, borrowed) with following labels: "Brasil, Corumba, Alto Praguay, H, Richter"/ female symbol/ "Schenk ling det."/"TYPUS"/"Atractocerus procerus Schklg."/"Dtsch. Entomol. Institut Berlin". Schenkling (19141 did not designate a holotype in his original description. The eye facets are relatively large, coarse, and separated into distinct rows. The antenna is widest near the middle. The Base of the elytra have small, somewhat imbricate tubercles (see SEM photos in chapter on Atractocerus ). Another South American species C A tracto cerus valdivianus ?} has finely faceted eyes, and lackes the tubercles on the elytra bases. (6} Atractocerus ater Kraatz, 1895, p. 163. This species was consider ed to be synonymous with the common African species brevicornis, but has recently been recognized as a valid species by V illiers (1969), who provides several characters, including the male genitalia, by which these two species may be separated. I have seen the type specimen of a ter (DEMB, borrowed) which bears the following labels: "Togo Conradt"/"Coll. Kraatz"/female symbol/"Atractoc. ater Kraatz"/ "Schenkling det."/"A. brevicornis L."/"Dtsch. Entomol. Institut Berlin" /"Dtsch. Entomol. Institut Eberswalde"/"HoloType"/Atractocerus ater Kraatz, A. Villiers 1968". As noted by V illiers, the male holotype has the abdominal apex missing. 546. (7) Melittommopsis nigra Lane, 1955. I have examined a female paratype for this species (BMNH, borrowed), with the following labels: "Para-type"/"S.Paulo, Camp, de Jordao, 1-5.1.1948, F.Lane col."/female symbol/"Paratipo"/"Brit.Mus."/"Brit.M us.,1955-221"/"Melittommopsis nigra sp.n., paratipo, F.Lane det.1955". (8) Hylecoetus m atsushitai Kono, 1938, p. 199 = H. derm estoides, NEW SYNONYMY. The female holotype, which Kono falsely regarded as a male, is of typical color and form (pale with apex of elytra black, and head of typical pale color), and has the following labels: "Saghalien, H.Kono, Haga, Shimizu"/"Hylecoetus matsushitai Kono, type", male symbol/"Picea jezuensis". A second specimen (female paratype) was seen from Shari, Kitawi, Hokkaido, 14/vi/28, det. Konishi. These specimens were borrowed from the Hakkaido University, Sapporo, Japan courtesy of Dr. S. Takagi. (9) I have seen the monotypic holotype of Mordella barbatus S c h a lle r, 1783, p. 322, a long-recognized synonym of Hylecoetus dermestoides (ZMHB, borrowed). (10) I examined 2 female syntypes of Lytta francofurthanus H erbst, 1784, p. 145, and confirm the long-standing synonymy with Hylecoetus dermestoides (ZMHB, borrowed). (11) Lymexylon a u s tra lis Erichson, 1842, p. 147. I have studied the holotype for this species (ZMHB, borrowed), which is in poor condition, missing its head and 4 legs. The locality label could not be read. The specimen is female, and darkly colored. This species is designated as type-species for my new genus Australymexylon (see text). (12) Lymexylon flabellicornis Schneider, 1791, p. 109. I have seen the type-series for this species, consisting of seven syntypes (ZMHB, borrowed), and a male LECTOTYPE is hereby designated. (All of the specimens are male, and there is no label data). (13) Lymexylon proboscideum Fabricius. Three specimens were borrowed from the Kiel collection in Copenhagen. A LECTOTYPE is hereby designated; the specimen has no label and is a typical male Hylecoetus dermestoides (This synonymy has been recognized for many years). One of the paralectotypes is a typical male dermestoides and the other a specimen of the fla b e llic o r n is morph. (14) Lymexylon abbreviatum Fabricius (= Atractocerus brevicornis) . The type was borrowed from Copenhagen, and simply bears the label : "L. abbreviatum Pflueg.". C151 Lymexylon r u fic o lle Kurosawaf 1949-f p. 27, I have seen a female paratype of this species courtesy of Mr, Winkler (borrowedi from "Yunchana, Fukushima Pref,, 17-VI-1947, Y. Kurosawa, Lgt.rr. Aside from color differences, there is nothing to separate this form from "typical" Lymexylon navale females. The basic color of ru fic o lle is black; the pronotum is read, with a darker area along the posterior margin, and a slightly darkened area on the disc postero-medially; the humeral angle of each elytron is reddish colored; the ventral surface of the beetle is black. Kurosawa C1956, p .82} l a t e r d escrib ed a male fo r th is sp e c ie s. I have not been able to borrow a specimen of the male for study, and therefore hesitate to synonymize the species with navale. Based on the color differences alone, I doubt that this species is valid. C161. Hylecoetus cossis Lewis. I synonymize this Japanese species w ith Hylecoetus dermestoides (see textL. Female specimens of ',c o ssis,, have the head and apex of the elytra black. I have seen females with only the elytral apices black, and some with only the head black. 549. (17) Notes on the nomenclature of Atractocerus, with Synonymy of Atractorus, a lapsus calami by Palisot de Beauvois (1801) . P a lis o t de Beauvois described Atractocerus in an obscure French journal, and there had been some question as to the correct page citation for the original description (see Palisot de Beauvois, 1801, La Decade Philosophique, Literataire, et Politique, vol. 9, no. 4, pp. 513-518, with 1 plate) . Although Atractocerus appeared first in the paper (as part of the caption for a plate, which, incidentally, had no page number!), it did not appear a second time. Rather, the alterna tive spelling Atractourus appears throughout the text, a total of ten separate times, including the formal description of the genus on page 516. Atractourus was the only spelling to appear in the text; it appeared 10 times; and the only place Atractocerus appeared was in th e cap tio n fo r th e p la te . What was to be done? After describing the problem to Dr. Donald Borrer (Department of Entomology, The Ohio State University), we discussed the alternatives and concluded that Atractocerus could be justifiably retained under the following rationale: (1) Atractocerus is the correct derivation from the greek for "spindle-horn"; the name which Palisot (1801) said he had chosen; (2) Atractocerus appears first in the text (as the caption to an in serted p la te , opposite page 515); and (3) reten tion of this name would maintain nomenclatural stability, avoiding the disruptions caused by a change to Atractourus. Based on these considerations, I concluded that Atractourus was a lapsus calami which should be emended under ICZN, A rt. 32(a)(ii). 550. I am particularly grateful to Donald J. Borrer and Charles A. Triplehorn for their advice in resolving this nomenclatural delima. (18) Justification for the synonymy of Hylecoetus flabellicornis with H. dermestoides. Because the species flabellicornis was recognized before Linneaus published his Systema naturae, and because it has been recognized as a valid species for more than two centuries since Schneid er validated the epithet in 1761 I was hesitant to synonymize it with H. dermestoides. This notion, however, is not mine alone. Pfeil (1859) hinted that flabellicornis might only be a form of the male of derm esto id es. The aedeagus of these two male forms are identical in form, and no females can be separated into two corresponding morphs. Further, I have seen specimens of both morphs from the same locality. I have not, however, thoroughly analyzed the geographic distributions of these morphs (or the other morphs of dermestoides), for which reason I discuss these forms in the text. (19) Synonymy o f genus NEOHYLECOETUS Kurosawaf 1956, with MELITTOMMA. I was not able to study the type-specimens of either Neohylecoetus javanus or Neohylecoetus philippensis. The descriptions,, and the very good drawings, given by Kurosawa, however, are sufficient to confidently place these species in Melittomma. I propose synonymy with Melittomma here. When the types can be studied, they should be compared with M. javanicum and related species. APPENDIX B: SUPPLEMENTARY ILLUSTRATIONS The two illustrations given in Appendix B were prepared by Larry Watrous and me for oral presentations on cladistics, and are included as a supplementary reference to my discussion on philosophical methods near the introduction of this paper. The text should be consulted for explanation. 551. Fig. 297. Flow-chart of systematics. Developed by L. E. Watrous and author. See text for discussion. 553. BORROW COLLECT OTHER TYPE SPECIMENS SPECIMENS AVAILABLE PURCHASE SPECIES alphi CONCEPT CHARACTER MATRIX CLASSIFICATION PHENETIC CLADISTIC ANALYSIS ANALYSIS INTERPRETATION biogaographica1 acological evolutionary F ig. 297 Fig. 298. Flow-chart of cladistic methodology. Developed by L.E.Watrous and the author. See text for discussion. INTERPRETATION: BIOGEOGRAPHY T CAUSAL EXPLANATIONS SELECTION OF BEST n ______CLADOGRAM: FALSE CHARACTER PARSIMONY & EXPLANATION: CHARACTER WEIGHTING PARALLELISM/CONVERGENCE REVERSALS I :..... n . : i . i ------HYPOTHESES OF *i ______MONOPHYLETIC RE-EVALUATION OF GROUPS CONFLICTING CHARACTERS POLARITY & HOMOLOGY CHARACTER POLARITY I DETERMINATION SEARCH FOR CHARACTERS 298 cn tn cn APPENDIX C: Another Response to E. King A quarter of a century ago, King (1955) argued that Atractocerus beetles are more primitive than generally appreciated, and that the sim ilarities to polyphagous beetle families are resultant of convergent evolution. His taxonomic solution to this situation was to elevate Atractocerus to the status of suborder, and recognize it as sister group to Adephaga + Polyphaga. This unorthodox position was quickly rebutted by Forbes (1956) and Selander (1959) on grounds of wing-venation characters, and has not been advocated since to my knowledge. Nonetheless, the other morphological arguments set forth by King in defence of his hypothesis have never been refuted. In the interest of thoroughness (and not in flailing a dead hypothesis), I believe it would be desirable to address his other points. This is done below, following a brief synopsis of the other arguments against King already in print. Forbes (1956) quickly pointed out that the hindwing venation was not in conflict with the inclusion of the genus in Polyphaga. Selander (1959) expanded upon this point, presenting an elegant scenario showing one possible explanation for the bizarre wing venation seen in A tracto cerus. Selander*s scheme, beginning with a relatively complete wing (,Melittomma ) and ending with this highly-modified version was simple, logical, and carefully ordered. He did not present it as the 556. 557. explanation, but wanted to show that if this explanation was consistant with known data, we could comprehend this type of wing without undoing the higher classification of the Coleoptera. Selander raised examples of other beetles with similarly shortened elytra, aside from the familiar Staphylinidae. He included Cantharids, Phengodids, Telegeusids, Cerambycids (Necydalines), Rhipiphorids (Rhipi- phrines + Rhipidiines), and Stylopids. Selander noted that Rhipiphorids, Telegeusids, Stylopids, Phengodids, and Atractocerus all share an in crease in the surface area of either the antennae or the palps. The usual explanation involves secretive habits and loss of flight in the females; but this is not true in Atractocerus. Selander suggested that low population density or an unusual factor in mating ethology or distribution has favored rapid flight in Atractocerus. Because the arguments which King made on morphological grounds, aside from characters of the hindwings (refuted by Forbes and Selander), are unsubstantiated I will briefly discuss each below: 1. FEMALE GENITALIA. The "proctiger" discussed by King (the "median sclerite" in my terminology) is present throughout the family Lymexylidae, though variously sclerotized. A partial division of the coxites, noted by Tanner (1927) - but not by King - is apparently a synapomorphy which unites all genera of Lymexylidae ( including A tracto cerus) . I do not believe that King's assertion that Tanner implied evolution of the polyphagan type female genitalia from tedephagan one is substantiated from Tanner's publication. 558. 2. MALE GENITALIA. The absence of "significant sim ilarities" to other male genitalia reported by King is simply false. I find no problem whatsoever in homologizing all parts of the genitalia of all genera of Lymexylidae with those of families in Cleroidea and Cucu- jo id e a . 3. THE MAXILLAE. The "thumb-like process" described by King in other genera of Lymexylidae is the fourth palpomere, and I have shown in my chapter on Atractocerus that it is actually present here as well (many authors preceeding King have also thought that palpomere IV was missing). The only really unusual thing about the maxillae in this genus is that females as well as males have a maxillary palp-organ. « This was not mentioned by King, and is, of course, consistant with other data for the family. 4. THE LARVAE. King suggested that the larva of Atractocerus was unlike that of other lymexylids. 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