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T TA /Î-T Dissertation i J l Y l i Information Service
University Microfilms International A Bell & Howell Information Company 300 N. Z eeb Road, Ann Arbor, Micfiigan 48106
8618856
Stribling, James Bentley
WORLD GENERIC REViSION OF PTILODACTYLIDAE (COLEOPTERA;DRYOPOIDEA)
The Ohio State University Ph.D. 1986
University Microfilms I nternsti 0 nâl SOO N. zeeb Road, Ann Arbor, Ml 48106
WORLD GENERIC REVISION OF PTILODACTYLIDAE (COLEOPTERA:
DRYOPOIDEA)
DISSERTATION
Presented in Partial Fulfillment of the Requirements for
the Degree Doctor of Philosophy in the Graduate
School of The Ohio State University
By
James B. Stribling, B.S., M.S.
*****
The Ohio State University
1986
Dissertation Committee: Approved by
C. A. Triplehorn
N. F. Johnson Û- Adviser D. E. Johnston Department of Entomology
B. D. Valentine Copyright by James Bentley Stribling 1986 To My Parents
ii ACKNOWLEDGEMENTS
There are many people to whom I am very grateful for academic, moral, physical, and financial support during my graduate career at The Ohio State University: my adviser.
Dr. Charles A. Triplehorn, for sound advice for the duration and understanding when I needed it most; Dr.
Norman F. Johnson for numerous discussions, suggestions, and welcome advice on systematic theory and taxonomic procedure; my officemates, colleagues, and fellow graduate students, especially, Paul S. Cwikla, Michael A. Ivie,
Richard S. Miller, and John A. Shuey, for seemingly unending discussions on systematics, for companionship on collecting trips, and for tolerance during these last few months; John F. Lawrence for much advice and comment on this as well as smaller simultaneous projects; W. Calvin
Welbourn for many hours spent in setting up the AT&T 3B2 and QTREE and then tutoring me on their use; John F.
Lawrence, Charles W. O'Brien, and Arthur J. Gilbert for gifts of specimens; Roland L= Seymour (Department of Botany, The Ohio State University) for fungal identifica
iii tions; Ho-Yih Liu (also Department of Botany, 0. S. 0.) for translating some of Nakane's works from Japanese; all of the curators in the "Materials" section for giving me the
opportunity to study their specimens (even past expiration dates on loan forms); Niki Lowell and Adam Rubinstein for the habitus illustrations (Figures 3 (N. L«) and 1 and 2
(A. R.); and Helen W. Monk for typing part of the manuscript.
Partial funding of museum trips was provided by Sigma
Xi (Grant-in-Aid of Research), The Ohio State University
Graduate School (Graduate Student Alumni Research Award), and the Knull Fund, Department of Entomology. I am very
grateful to them.
I owe immense thanks to my wonderful parents. Dr. and
Mrs. Loutrelle, for their financial support, endless moral
support, encouragement, and for all of those times I needed
"a good talkin' to" and got it. Finally, I thank my wife,
Sally, for the hours of assistance she has given me in my
research, taking time out of her own to process specimens,
to type, or to take over household duties when I might be nearing a deadline. My greatest thanks to her are for her
unwavering love, encouragement, and companionship.
IV VITA
October 13, 1958...... Born - Jackson, Mississippi
1980...... B. S., The University of Mississippi, Oxford, Mississippi
1982 # ••••«•••••• # #M# S #, The Ohio State University, Columbus, Ohio
1980-present...... Graduate Research/Teaching Associate, Department of Entomology, The Ohio State University
PRESENTATIONS
1985. Phylogenetic Implications of Metathoracic Wing Venation in Dryopoidea (Coleoptera). Annual Meeting of Entomological Society of America, Hollywood, Florida. The Biology and Systematics of Anchvtarsus (Coleoptera: Ptilodactylidae). Annual Meeting of the North Central Branch, Entomological Society of America, Lexington, Kentucky.
The Biology, Phylogeny, and Zoogeography of the Wing-Toe Beetles (Ptilodactylidae). Departmental seminar. Department of Entomology, The Ohio State University, Columbus.
1984. The Faces of Systematic Biology. Invited seminar. Department of Biology, The University of Mississippi, Oxford. 1981. The New World Genera of the Ptilodactylidae (Coleoptera). Annual Meeting of the Entomological Society of America, San Diego, California.
PUBLICATIONS
1986. Revision of Anchvtarsus (Coleoptera: Dryopoidea) and a key to the New World genera of Ptilodactylidae. Ann. Entomol. Soc. Amer. 79: 219-234.
1984. Taxonomic and nomenclatorial notes on Caribbean Tropicus Pacheco (Coleoptera: Heteroceridae). Proc. Ent. Soc. Wash. 86 (4): 946-950. (with M. A. Ivie) Report on the LeConte symposium on the classification and phylogeny of the Coleoptera, Detroit and Columbus, 1983. Coleop. Bull. 38 (1): 98-102. (with M. A. Ivie)
FIELDS OF STUDY
Major Field: Systematic Entomology with specialization in Coleoptera biosystematics.
VI TABLE OF CONTENTS
ACKNOWLEDGEMENTS...... iii VITA...... V
LIST OF TABLES...... ix
LIST OF FIGURES...... x CHAPTER PAGE
I. INTRODUCTION...... 1
II. METHODS AND MATERIALS ...... 4
Methods Phylogenetic...... 4 General...... 8 Material...... 11
III. NATURAL HISTORY...... 16 IV. CHARACTERS, POLARIZATION, PHYLOGENETIC ANALYSIS, AND CLASSIFICATION...... 19 Monophyly of the Ptilodactylidae. . . . 19 Character discussion...... 19 H e a d ...... 20 Thorax...... 23 Abdomen...... 28 Character polarization...... 29 Discussion of phylogenetic analysis and resulting classification ..... 32
V. TAXONOMY...... 38 Key to World Genera of Ptilodactylidae (Adults)...... 38 The World Genera of Ptilodactylidae Anchvtarsus Guerin-Meneville . . . 42
vii Cladotoma Westwood ...... 44 Paraiichas White ...... 47 Hovactvla Fairmaire...... 50 Pseudocladotoma Pic...... 52 Octoqlossa Guérin-Méneville. . . . 54 Bradvtoma Guérin-Méneville .... 57 Aploqlossa Guérin-Méneville. . . . 60 Anchycteis Horn...... 63 Epilichas White...... 66 Bvrrocrvptus Broun ...... 69 Araeopidius Cockerell...... 72 Stenafricus NEW GENUS...... 74 Daemon de Laporte...... 77 Falsotherius P i c ...... 8 1 Pseudoepilichas Armstrong and Nakane ...... 83 Ectvphodactvla NEW GENUS ..... 85 Ptiloderes NEW G E N U S ...... 89 Pherocladus Fairmaire...... 91 Lachnodactvla Champion ...... 94 Ptilodactvla Illiger ...... 97 Stirophora Champion...... 118 Chelonariomorphus Pic...... 121 Epiptyqma NEW GENUS...... 123 Therius Guerin-Mènev i l i e ...... 126 Incertae sedis...... 129 Exclusions...... 129
LITERATURE CITED...... 130 APPEmiX
A. Figures ...... 144
Vlll LIST OF TABLES
TABLE PAGE
1. Fungal spores and hyphae identified from gut content of some species of Ptilodactlidae. . . 18
2. Hypothesized polarities of characters used in phylogenetic analysis of the genera of Ptilodactylidae...... 35
3. Phylogenetic hypothesis resulting from QTREE analysis of characters in Table 2. Reversals are indicated by dots at the upper right hand side of the character numbers; characters with multiple apomorphic appearances are 2, 3, 4, 5, 7, 9, 11, 12, 13, 14, 15, 16, 18, 20, 23, 25, 27, 29, 31, 35, and 3 8 ...... 36
4» Proposed supraspecific classification of the Ptilodactylidae based on phyletic sequencing • 37
ix LIST OF FIGURES
FIGURES PAGE 1. Dorsal habitus. Anchvcteis velutina, male...... 146
2. Dorsal habitus, Ptilodactvla militaris, male...... 148
3. Dorsal habitus. Ptiloderes sp., female. » . . 150
4. Serrate antenna; Bradvtoma lineata...... 152
5. Ramous antenna, nonarticulated rami; Paralichas q u e r i n i ...... 152
6. Ramous antenna, nonarticulated rami, Cladotoma maculicollis...... 152
7. Ramous antenna, articulated rami; Ptilodactvla sp...... 152
S. Tarsus, lateral view showing simple tarsomeres and large onychium; Eulichas sp...... 154
9. Tarsomere IV, dorsal view showing large onychium; Eul%cnas sp...... o. 154
10. Pectinate ungues; Paralichas querini. .... 154 11. Bifid ungues; Ptilodactvla sp. The mesal furcations are often more or less truncate. . 154
12. Modified tarsus showing reduction of tarsomere IV, ventral lobe of III, simple ungues, and reduced onychium; Cladotoma cucullatus...... 154
13. Simple tarsus: Araeopidius monachus...... 154
14. Simple tarsus showing minor lobes on tarsomeres II and III, and no reduction of IV; Pseudocladotoma maculata...... 154
15. Labium, ventral view, palp and setae omitted from
X right half; modified ligula, biacuminate with each half tapered to blunt apex; Ptiloderes sp.. . 156
16. Labium, ventral view, palp and setae omitted from right half; hexacuminate ligula; Bradvtoma a u r i t a ...... 156 17. Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate with each half tapered to blunt apex, apical palpomere with highlighted area of membranous region showing portion of setal clothing; Ectvphodactvla fiski...... 156 18. Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate; Falsotherius atricolor...... 156
19. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate with lateral apices reduced; Anchvcteis velutina . 158
20. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Bvrrocryptus sp...... 158
21. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Epilichas candezei...... 158
22. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Cladotoma sp...... 158
23. Labium, ventral view, palp and setae omitted from right half; simple ligula folded ventrally; Stirophora Ivciformis ...... 160
24. Labium, ventral view, palp and setae omitted from right half; modified ligula, multiacuminate; Paralichas trivittis...... 160
25. Labium, ventral view, palp and setae omitted from right half; modified ligula, hexacuminate; Octoqlossa sp...... 160
26. Left maxilla, ventral view; multiacuminate galea; Paralichas sp...... 162
27. Right maxilla, ventral view, lacinia forming spore
xi brush, apical palpomere elongate and largely membranous; Lachnodactvla parviscutum .... 162
28. Left maxilla, ventral view, biacuminate galea; Epilichas candezei...... 164
29. Left maxilla, ventral view, biacuminate galea with mesal branch absent; Anchvcteis velutina. . . 166
30. Left maxilla, ventral view, biacuminate galea with mesal branch reduced; Bvrrocrvptus sp. . . . 166
31. Left maxilla, ventral view, lacinia forming spore brush; Aploqlossa aureonotata ...... 166
32. Left maxilla, ventral view, galea and lacinia forming spore brush; Daemon gigas ...... 166
33. Left maxilla, ventral view, cylindrical galea with apical cluster of setae; Bradvtoma aurita . . 168
34. Left maxilla, ventral view; Ptiloderes sp.. . 168
35. Right maxilla, ventral view; Falsotherius at r1 color ...... 168
36. Right maxilla, ventral view, apical palpomere with highlighted area of membranous region showing portion of setal clothing; Ectvphodactvla fiski. 168 37. Left mandible, dorsal view; Aploqlossa aureonotata ...... 170
38. Left mandible, dorsal view, showing concave molar area; Daemon gigas...... 170
39. Left mandible, dorsal view; Paralichas querini ...... 170 40. Right mandible, dorsal view; Octoqlossa cvaneipennis...... 170
41. Left mandible, dorsal view; Lachnodactvla parviscutum...... 172
42. Left mandible, dorsal view; Anchvcteis velutina ...... 172
43. Left mandible, dorsal view; Epilichas candezei. .
xii ...... 172 44. Labrum, dorsal view; Falsotherius atricolor . 174
45. Labrum, dorsal view; Aploqlossa aureonotata . 174
46. Labrum, dorsal view; Cladotoma sp...... 174
47. Labrum, dorsal view; Ptilodactvla sp..... 174
48. Labrum, dorsal view; Anchvcteis velutina. . . 174
49. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal Insertions; Octoqlossa sp...... 176
50. Head and prothorax, frontal view, shaded areas - eyes, blackened areas = antennal insertions; Araeopidius monachus...... 176
51. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Cladotoma sp...... 176
52. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ptiloderes sp...... 176
53. Bead and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ectvphodactvla fiski...... 176
54. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ptilodactvla sp...... 176
55. Tergum and sternum of male visible abdominal segment VI, lateral view; Eoiptvcma rufa. . . 178
56. Male visible abdominal segment VI, dorsal view of Figure 55; Epiptyqma rufa ...... 178
57. Tergum and sternum of male visible abdominal segment VI, lateral view; Epiptyqma sp. ... 178
58. Male visible abdominal segment VI, dorsal view of Figure 57; Epiptyqma sp...... 178
59. Tergum and sternum of male visible abdominal segment VI, lateral view; Epiptyqma sp. ... 178
xiii 60. Male visible abdominal segment VI, dorsal view of Figure 59; Epiptvoma sp...... 178
61. Head and prothorax, lateral view, shaded area = eye; Pherocladus sp...... 178 62. Head and prothorax, lateral view, shaded area = eye; Araeopidius m o n a c h u s ...... 178
63. Head and prothorax, lateral view, shaded area = eye; Octoqlossa sp...... 180
64. Head and prothorax, lateral view, shaded area = eye; Paralichas q u e r i n i ...... 180
65. Head and prothorax, lateral view, shaded area = eye; Bradvtoma lineata...... 180
66. Head and prothorax, lateral view, shaded area = eye; Ectvphodactvla fiski ...... 180
67. Head and prothorax, lateral view, shaded area = eye; Chelonariom orphus sp...... 180
68. Head and prothorax, lateral view, shaded area = eye; Ptilodactvla sp...... 180
69. Anterior dorsum; Araeopidius monachus .... 182
70. Anterior dorsum; Chelonariomorphus subconvexus. . 182
71. Anterior dorsum; Ectvphodactvla fiski .... 182
72. Anterior dorsum; Paralichas querini ...... 182
73. Anterior dorsum; Ptiloderes sp...... 184
74. Pronotum, dorsal view; Ptiloderes sp...... 184 75. Pronotum, dorsal view; Ptiloderes sp...... 184
76. Pronotum, dorsal view; Ptiloderes sp. .... 184
77. Ovipositor, dorsal view; Aploqlossa sp. . . . 186
78. Ovipositor, dorsal view; Araeopidius monachus . . 186
XIV 79. Ovipositor, distal portion, dorsal view; Paralichas trivittis...... 188
80. Ovipositor, distal portion, dorsal view; Stirophora sulcipennis...... 188
81. Ovipositor, distal portion, dorsal view; Daemon sp...... 188 82. Ovipositor, distal portion, dorsal view; Falsotherius atricolor...... 188
83. Ovipositor, dorsal view; Cladotoma sp» . . . 190
84. Ovipositor, right lateral view of Figure 83; Cladotoma sp...... 190
85. Aedeagus, ventral view; Bradvtoma aurita. . . 192
86. Aedeagus, right lateral view of Figure 85; Bradvtoma aurita...... 192
87. Aedeagus, dorsal view; Aploqlossa aureonotata . , ..... 194
88. Aedeagus, right lateral view of Figure 87; Aploqlossa aureonotata...... 194
89. Aedeagus, dorsal view; Ectvphodactvla fiski . 196
90. Aedeagus, dorsal view; Chelonariomorphus sp.. 196
91. Aedeagus, dorsal view; Paralichas querini . . 196
92. Aedeagus, dorsal view; Stenafricus sp» ... 196
93. Aedeagus, dorsal view; Cladotoma sp.. . . . 198
94. Aedeagus, dorsal view; stirophora Ivciformis. 198
95. Aedeagus, dorsal view; Epilichas candezei . . 200 96. Aedeagus, dorsal view; Lachnodactvla monticola . . 200
XV CHAPTER I
INTRODUCTION
The Ptilodactylidae is a family of semiaquatic and terrestrial beetles which is nearly cosmopolitan in distribution (Stribling, 1986). Species are present in every major biogeographic region except the Palearctic and
Chile. The greatest species diversity is in tropical areas with roughly similar numbers in the Old and New Worlds.
Characteristics traditionally used to recognize ptilodactylids have been a heart-shaped (= cordiform) mesoscutellum, pseudotetramerous tarsi (with the third tarsomere ventrally lobed and the fourth reduced), and the pectinate antennae of the males. Exclusive use of these characters has allowed recognition of several genera as ptilodactylids, but has also led to confusion regarding phylogenetic relationships among these genera as well as between the Ptilodactylidae and other Dryopoidea (Crowson,
1967, 1978; Lawrence and Newton, 1982; Arnett 1968). An extension of this confusion is a misunderstanding of the generic composition of the family (Arnett, 1968; Borror, ^ al. 1978).
Boving and Craighead (1931) recognized affinities between the larvae of Anchvtarsus Guerin-Meneville and
Ptilodactvla Illiger and placed them together, elevating 2 the group to familial status. Later, fourteen more genera were transferred from the Dascillidae to the
Ptilodactylidae (Crowson, 1967). For a more complete review of the suprageneric taxonomic history see Stribling (1986).
There have been few regional studies of the
Ptilodactylidae in the Old World. Deleve (1972) investigated the fauna of the Philippines and the Bismarck
Archipelago, describing 20 species of Ptilodactvla, one of
Pherocladus Fairmaire, and the genus valoka Deleve. Many papers have treated various Oriental species (White, 1859;
Nakane, et 1963; Nakane, 1948, 1950, 1952, 1956, 1963,
1977; Sato, 1964, 1968, 1979, 1983; Chûjô and Sato, 1970;
Lewis, 1895) usually describing small numbers of Ptilodac tvla, Epilichas White, and Paralichas White. Armstrong and
Nakane (1956) described Pseudoepilichas from Japan.
Even fewer studies have dealt with the New World fauna.
Johnson and Freytag (1978, 1982) described species and reviewed the Ptilodactvla of the United States. Champion, in the "Biologia Centrali-Americana" (1897), recognized fifty new species of Ptilodactvla and four new genera from
Mexico and Central America. Between 1916 and 1958, Maurice Pic described 206 species of Ptilodactvla (102 in the New
World), and 7 monotypic genera.
This is the first comprehensive study of the Ptilodactylidae. Although I was unable to examine the 3 types of all described species, I was able to study
representatives of nearly all genera and of many species.
As a result, I present a phylogenetic hypothesis of the
major monophyletic lineages of the Ptilodactylidae of the world. This hypothesis will allow evaluation of phylogenetic relationships of subsequently studied or
discovered taxonomic units, and of the significance of new characters and character states. CHAPTER II
METHODS AND MATERIALS
Phylogenetic
Delineation of supraspecific taxonomic categories in classifications has long been highly subjective, often being based on overall similarity rather than on objective statements of phylogenetic relationship. Cladistic methodology (Hennig, 1966; Eldredge and Cracraft, 1980;
Wiley, 1981) has provided systematic biology with a method for recognition of groups of organisms sharing unique phylogenetic history. Construction of phylogenetic hypotheses through employment of these methods produces a hierarchically-arranged diagram of operational taxonomic units (OTü's) which may be translated into the hierarchy of the Linnean system of classification. Representation of
this common history in classificatory form has been discussed in regard to its information content (Farris,
1980) and application and necessity for non-systematic biological subdisciplines (Dobson, 1985).
In this study I use outgroup comparison analysis for
postulation of derived, or apomorphic, character states. Joint possession of apomorphic character states among OTü's
indicates probable common ancestry and monophyly. I
recognize monophyletic taxa (Hennig, 1966) as those 5 composed of: 1) the most recent common ancestor which is inferred to possess a particular apomorphy, and 2) all known descendants of that ancestor. Paraphyletic taxa
(Farris, 1974) are converted into monophyletic ones by synonymization and/or splitting of previously recognized taxa. In some cases this requires erection of new taxa.
Eldredge and Cracraft (1980: 39) point out that monophyletic higher taxa must consist of two or more species. Monotypic genera, therefore, cause non-monophyly
(specifically, paraphyly) in their sister-genera by allowing exclusion of one of the descendants of that ancestor. The only ancestor which a species has in common with a lineage is also ancestral to another part of that lineage. To avoid paraphyly, all descendants of that ancestor must be included in the taxon. If the sister-OTU of the species is of preexisting generic status, they are combined. In all cases this tactic produces a monophyletic genus. In other words, a monotypic genus may only be recognized when its species possesses no synapomorphy with a preexisting genus. The most important single component of phylogenetic methodology is the postulation of the relative polarity of character states. Although many criteria have been discussed as potential aids in this inference (Crisci and
Stuessy, 1980; de Jong, 1980), outgroup comparison is 6 regarded as the most reliable method (Watrous and Wheeler,
1981; Haddison et al., 1984; Donoghue and Cantino, 1984) and I have used this technique exclusively.
The Ptilodactylidae are included in the Dryopoidea
(Crowson, 1955, 1960, 1978; Lawrence, 1982; Lawrence and
Newton, 1982) the latter having been hypothesized to consist of two main lineages (Lawrence and Newton, 1982):
1) Callirhipidae - Eulichadidae - Ptilodactylidae -
Chelonariidae; 2) Heteroceridae - Lutrochidae - Limnichidae
- Dryopidae - Elmidae - Psephenidae. The Eulichadidae have
also been hypothesized to be the sister-group of the Ptilodactylidae (Crowson, 1978).
Assuming monophyly of the former lineage, I used the
following taxa for outgroup comparison:
Eulichadidae
Eulichas spp.
Stenocolus scutellaris LeConte
Callirhipidae
Zenoa picea (Beauvois)
unidentified sp. from Vietnam
Psephenidae
Psephenus herricki (DeKay)
After demarcation of characters within Ptilodactylidae
into binary components, hypotheses of apomorphy through
the process of polarization were sought. This process entailed the following steps: 7
1) Check for occurrence of character state in
Eulichadidae. Absence indicates probable apomorphy.
2) Check for occurrence in Callirhipidae. Absence corroborates hypothesis of apomorphy.
3) Check for occurrence in Psephenidae. Absence, further corroboration. If the character state is found in Eulichadidae and either Callirhipidae or Psephenidae, it is hypothesized plesiomorphic and its alternative state, apomorphic.
In some cases, reference to outgroups did not provide evidence for the polarity of complex variation, such as that of the mouthparts and tarsi. Rather than attempt to align this variation into polarized transformation series I have chosen to follow what I consider to be a more conservative approach. With no evidence as to polarity or of the interrelationships of the character states, any attempt at coding them as a series would produce immediate subjective bias. I experimented with coding each of the states as independent apomorphies; however, this introduced much homoplasy and longer trees. Elimination of these characters from the analysis lowered homoplasy and consequently shortened trees. The phylogenetic computer program used for character matrix analysis was QTREE, written in C by Dr. L. E. 8 Watrous (FMNH); it was executed on an AT&T 3B2/300. The
program searches for the most parsimonious trees using the
mode of search specified by the options chosen. I used the following options:
-bO This specifies the number of pathways to be
explored in the search for the shortest tree. The
numerical value can be altered from 0 - 3, each increase
causing more trees to be examined.
-k2 This specifies that the computer randomly sort the
OTD*s; the numerical value, how many times the sort should
be repeated.
General
Relaxing. - Specimens were relaxed in an all
purpose ammonia and detergent household cleanser in a sonic
cleaner. They were then vibrated in tap water to remove
the ammonia and detergent. Specimens mounted on points or cards were removed before relaxing.
Dissecting. - All mouthparts and genitalia were removed
from relaxed specimens with jewelers' forceps (no. 3 or no.
5) and/or minuten pins mounted in wooden splints.
Dissected parts were dried of excess water and placed in
glycerine in small glass dishes in a wooden holder (DeLong
and Davidson, 1937) for serial observation. Clearing. - when necessary for viewing detail. 9 dissected structures were cleared in hot KOH and water solution for approximately 3 minutes. The specimen was then removed; rinsed in cold tap water, and examined. If degree of clearing was insufficient, it was placed back in the KOH and heated further.
Illustration. - Mouthparts or genitalia were placed on a shallow depression slide with a minimal amount of glycerine. The specimens were covered with a drop of K-Y
(registered trademark) Lubricating Jelly (Johnson &
Johnson), positioned, and allowed to stand for 2-3 minutes.
The specimen was then covered with a drop of glycerine and allowed to stand for another 2-3 minutes until the K-
Y/glycerin interface stabilized. Illustration then commenced. The K-Y gel is water soluble and, when allowed to stand uncovered, loses moisture. Thus, it becomes somewhat coagulated, preventing drifting of the specimens.
This technique (P. S. Cwikla, pers. comm.) is ideal for illustration purposes. Illustrations were made with a camera lucida on Wild M5A dissecting scope for all except the mouthparts and genitalia which were illustrated with a drawing tube on a Wild H12 compound microscope.
Storage of dissections. - Dissected structures are stored in polyethylene microvials with a drop of glycerine.
The vials are mounted on the same pins with the specimens.
Taxonomic lists. - The taxonomic lists following the 10 descriptions contain all names, specific, subspecific, and varietal, which have been published under the ptilodactylid genera. Asterisks (*) indicate those of which I have seen holotypes, type-compared specimens, or specimens which I feel were reliably determined. The specific epithets are listed in alphabetical order followed by author and date of description; acronyms in parentheses indicate the location of the holotype or type series as known by my examination of the specimen, personal communication with curators, or from the literature. At the end of almost all entries is a geographic locality indicating where the type specimen(s) was (were) collected; in some instances additional information was added from label data. Occasionally, no locality was published with the description and I have seen no determined specimens. 11 Materials
This study is based on approximately 25,000 specimens accumulated through loans from the following museum, university, and private collections. Acronyms listed are those used in the text for citation of specimens and for type repositories,
AMNH American Museum of Natural History, New York, NY;
L. H, Herman, Jr.
ANSP Academy of Natural Sciences of Philadelphia,
Philadelphia, PA; Donald Azuma.
BMNH British Museum (Natural History), London, England;
C, M. F. von Hayek, P. M, Hammond,
CASC California Academy of Sciences, San Francisco,
CA; D. H, Kavanaugh, Jr,
CBBC Cheryl B, Barr, private collection. Baton Rouge,
LA,
CISB California Insect Survey, University of California, Berkeley, CA; J. A, Chemsak,
CNCI Canadian National Collection of Insects,
Agriculture Canada, Ottawa, Ontario; J, M,
Campbell, L. LeSage, A. Smetana,
CSIR Commonwealth Scientific and Industrial Research Organization, Canberra, ACT; J, F, Lawrence, 12 CüCC Department of Entomology, Clemson University, Clemson, SC; J. C. Horse.
DKYC Daniel K. Young, personal collection, Madison, WI.
EEHC Enns Entomological Museum, University of Missouri, Columbia, MO; E. G. Riley.
EGRC Edward G. Riley, personal collection. Baton Rouge,
LA.
FMNH Field Museum of Natural History, Chicago,
IL; H. G. Nelson. FSCA Florida State Collection of Arthropods,
Gainsville, FL; M. C. Thomas, R. E. Woodruff.
GLPC Gary L. Peters, personal collection, Corvallis,
OR. HAHC Henry F. and Ann T. Howden, personal collection,
Ottawa, Ontario.
HUIC Museum fur Naturkunde der Humboldt-Universistat zu
Berlin; M. Uhlig.
ICCM Carnegie Museum of Natural History, Pittsburgh,
PA; G. Ekis.
INHS Illinois Natural History Survey, Urbana, IL; D. W.
Webb.
INPA Institute Nacional de Pesguisas da Amazonia,
Manaus, Brasil; V. Py-Daniel.
IZWP Institut Zoologique, Académie Polonaise des
Sciences, Warszawa, Poland; S. A. Slipinski. 13 JBSC James B. Stribling, personal collection, Columbus,
OH.
JEWC James E. Wappes, personal collection. Woodlands,
TX.
KJOC Karl Joplin, personal collection, Columbus, OH.
KÜBC Kagoshima University Biological Collections, Kagoshima-Shi, Japan; Takehiko Nakane.
LACM Los Angeles County Museum of Natural History, Los
Angeles, CA; J. P. Donoghue, R. R. Snelling.
LGBZ Larry G. Bezark, personal collection, Sacramento,
CA. LSUC Louisiana State University, Baton Rouge, LA; C. B.
Barr, J. B. Chapin.
MAIC Michael A. Ivie, personal collection, Bozeman, NT.
MCZC Museum of Comparative Zoology, Harvard University,
Cambridge, MA; A. F. Newton, Jr.
MNHP Museum National D'Histoire Naturelle, Paris, France; N. Berti, C. Girard.
MRAC Musée Royal de L'Afrique Centrale, Tervuren,
Belgium; J. Decelle.
MSUC Michigan State University, Lansing, MI; R. L.
Fisher.
HUIC Mississippi State University, Starkville, MS; R.
Brown. MZSP Museu de Zoologia da Universidade de Sao Paulo, 14 Sao Paulo, Brazil; Cleide Costa. NARZ Northern Arizona University, Flagstaff, AZ; C. D,
Johnson.
NCSÜ North Carolina State University, Raleigh, NC; C. Parron.
NDSU North Dakota State University, Fargo, ND; E. U.
Balsbaugh, Jr.
NMNH National Museum of Natural History, Smithsonian
Institution, Washington, D. C.; P. J. Spangler, J.
M. Kingsolver.
OSUC Ohio State University, Columbus, OH; C. A. Triplehorn.
OSUO Oregon State University, Corvallis, OR; G. L.
Peters.
RHTC Robert H. Turnbow, personal collection. Fort
Rucker, AL. RSMC Richard S. Miller, personal collection, Columbus,
OH. RSNB Institut Royal des Science Naturelles de Belgique,
Bruxelles, Belgium; P. Dessart.
RUIC Rutgers University, New Brunswick, NJ; G. W.
Wolfe.
SMSH Stovall Museum of Science and History, University
of Oklahoma, Norman, OK; H. P. Brown.
TAMT Texas A & M University, College Station, TX; S. Merritt.
ÜART University of Arizona, Tuscon, AZ; F. G. Werner.
UBCZ University of British Colombia, Vancouver, BC, Canada; S. G. Cannings.
UCDC University of California, Davis, CA; R. 0.
Schuster.
UICM University of Idaho, Moscow, ID; W. F. Barr, Jr.
UMCE Université de Montreal, Quebec, Canada; M. Coulloudon.
UMIS University of Mississippi, Oxford, MS; P. R. Lago.
UNTC Universidad Nacional de Tucuman, Argentina; A.
Willink. VPSU Virginia Polytechnic Institute and state
University, Blacksburg, VA; M. Kosztarab.
The following acronyms are used only in reference to type repositories.
ANMM National Museum of Victoria, Melbourn, Australia.
BANM Museu de Ciencias Naturales, Buenos Aires,
Argentina.
DEIE Institute of Plant Protection Research, Eberswalde,
Germany (formerly, Deutsches Entomologisches Institut).
HECM Hope Entomological Museum, Oxford University,
Cambridge, England. UZHD Universiteits Zoologiske Museum, Copenhagen. CHAPTER III
NATURAL HISTORY
In general, the biology of ptilodactylids is poorly known. The larvae of Anchvtarsus are aquatic and feed on decaying vegetable material in lotie situations (LeSage and
Harper, 1976; Spangler, 1966, 1981, 1982, 1983; Stribling,
1986). Though found in flowing water, individuals are not subjected to the force of the current due to specific location within leaf packs. Anchvtarsus bicolor has also been collected in large numbers from beaver dams in Ontario
(L. LeSage, pers. comm.).
Other larvae known to be aquatic, Anchvcteis velutina Horn and Araeopidius monachus (LeConte), have been reported to burrow in rocky substrates and to feed on the roots of emergent vegetation (Leech and Chandler, 1956; Brown, 1972),
They have also been listed as feeding on plant materials by shredding (Doyen and Ulrich, 1978) and as boring in soft wood (Dudley and Anderson, 1982). Specimens of Anchvcteis have been collected from the vegetation within a bog runoff; and Araeopidius from duff and leaves in creeks (H. P. Brown, pers. comm.).
On 3 July, 1982, in Oaxaca (near Vista Hermosa) I collected five species of Ptilodactvla (95 specimens) by
16 17 beating in an area of approximately 20 square meters. Some of the vegetation was spotted with sooty mold; gut content analysis of these specimens has yielded no structures which
I could identify as fungal spores or hyphae. However, fungal components were discovered in the guts of other specimens in the course of genitalic dissections (see Table
1). White (1859) described Paralichas querini from specimens which had emerged from structures he proposed were pupal cases of the beetles. When I examined these specimens in the BMNH I found an anonymous note attached to one of them saying that the structure is a tineid larval case (Lepidoptera). A noté on another specimen states that both the tineid larva and beetle have been removed.
The largest numbers of adults are collected nocturnally at lights. Diurnally, they are found on vegetation with the aquatic species usually near water. 18 Table 1. Fungal spores and hyphae identified from gut contents of some species of Ptilodactylidae (fungal identi fications by R. L. Seymour).
.Species. Contents Epjptygma rufa (Champion) Pteroconium sp. Botrvoderma sp.
Stirophora sulcipennis Champion mixed unknown hyphomycete and coelomycete spores
Ptilodactvla sp, mixed unknown hyphomycete and coelomycete spores
Aploalossa sp. Alternaria sp. Capnodendron sp. coelomycetespores Lachnodactvla monticola Champion Pestalotia sp. mixed unknown hyphomycete and coelomycete spores Epiptvqma sp. Coniosporium sp. Pestalotia sp.
Epiptvqma maculata (Champion) Alternaria sp.
Epiptvqma rufipennis (Pic) Pestalotia sp,
Daemon sp. Pestalotia sp. Scleroqraphium sp. possibly, Hvcoenterolobium sp. CHAPTER IV
CHARACTERS, POLARITY, PHYLOGENETIC HYPOTHESIS, AND
CLASSIFICATION
Honophvlv of the Ptilodactylidae
The lineage of the Dryopoidea consisting of the
Callirhipidae, Eulichadidae, Ptilodactylidae, and
Chelonariidae has been hypothesized as monophyletic
(Lawrence and Newton, 1982). Crowson (1978) indicated the probable sister-group relationship of the Eulichadidae to the Ptilodactylidae (Crowson, 1978). The Eulichadidae, as well as the Callirhipidae, possesses a large onychium
(Figs. 8, 9); in the Ptilodactylidae this structure is absent or very highly reduced (Figs. 12-14). I hypothesize this as evidence in support of monophyly of the Ptilodactylidae.
Character discussion
Suss and Puppin (1976) presented an investigation of the adult and larval anatomy of Ptilodactvla exotica
Chapin, Therefore, I will discuss and illustrate only those adult integumental features used in phylogenetic inference and descriptions.
The head and thorax have provided the majority of characters used in this study. Abdominal characters are limited to the form of male visible abdominal sterna V and
19 20 VI (including tergum of the latter), the ovipositor, and the aedeagus.
Head. The 11-articled antennae are often sexually dimorphic with the females usually more conservative. With the exception of some female Cladotoma. ramous antennae are found only in males and have the mesally-directed rami either articulated or nonarticulated. The articulated rami
(Figs. 2, 7) occur from antennomeres IV-X and have a narrow membranous base. The rigid nonarticulated rami (Figs. 1, 5,
6) occur on antennomeres III-X and arise proximally, medially, or distally on the antennomeres. This type of ramus is found in Cladotoma and is very broadly dorsoventrally flattened (Fig. 6). In some female
Cladotoma, rami are present but are much shorter and narrower than in conspecific males.
Serrate antennae vary from extremely weak (Fig. 3) to very strong serrations (Fig. 4). Almost all females possess this type of antennal structure and show the entire range of serrations. Serrate antennae in males are usually more strongly serrate than in conspecific females. The antennomeres are mesally serrate from III-IX.
The epistomal sulcus extends across the frons below and approximate to the antennal insertions. It is sometimes deeply impressed with the frons even with or raised above the clypeal level. The sulcus may be weakly impressed with 21 the frons and clypeus equal in level; it is also occasionally absent. Head shape is usually approximately quadrate (Pigs. 51, 52, 54). Octoqlossa. Ectvphodactvla
NEW GENUS, and Araeopidius depart from this configuration with a more or less elongate head capsule (Figs. 49, 50,
53).
The maxillary variation is large, with the major differences occurring in the galea and lacinia. In many genera, particularly within the Ptilodactylinae, the lacinia has a very well-developed spore brush similar to that described by Ashe (1984) for the subtribe Gyrophaenina
(Staphylinidae: Aleocharinae) in which the lacinial apex forms a discoid structure densely beset with short, stout spines (Figs. 27, 31). The spines and setae of the lacinia are sometimes thinner, more elongate, less dense, and not in a discoid structure or arrangement (Figs. 26, 28-30, 33-
36). I restrict my definition of spore brush to exclude these types.
In Daemon and Stenafricus NEW GENUS, the galea is closely associated with the spore brush, having similar short, very stout spines (Fig. 32). Likewise, the galea of
Ptilodactylinae is closely associated with the spore brush disc but possess longer and finer spines or setae (Fig. 27,
34). In this study, the galeae of Daemon and Stenafricus are considered to be part of the spore brush. 22 In other forms, the laclniae may be apically blunt
(Fig. 36) or acute, almost always heavily setose, with the galeae exhibiting the greatest variation, being either biacuminate or multiacuminate. Eoilichas and Anchvcteis are described as having biacuminate galeae (Figs. 28, 29); however, the mesad furcation of Anchvcteis is underdeveloped in relation to the laterad (Fig. 29).
Multiacuminate galeae are seen in most Cladotominae and possess three or more furcations, each with a number of short, apparently random subbranches (Fig. 26).
Mandibular variation includes 1-3 denticles, presence or absence of molar areas, dorsal ridge, and prostheca. In
Eoilichas (Fig. 43), the thin dorsomedian portion of the mandibles often possesses an emargination which I do not consider to be homologous to true multidenticulations seen, e. g., in Aploglossa (Fig. 37) and Daemon (Fig. 38).
Distinct molar areas (Figs. 37, 38, 41, 42) are often present with that of the left mandible usually concave
(molar socket) and the right, more or less convex. The prostheca is located mesally and distally to the molar areas; it is a lightly melanized lobe, with shorter or longer setae (Figs. 37, 38, 40-43).
The labrum varies in shape, condition of anterior margin, and presence or absence of anterolateral tufts of setae. In the Cladotominae, the labrum is strongly transverse (basal width approximately 3 1/2 times its greatest length. Fig. 46). All other forms have the basal width no more than approximately twice the median length
(Figs. 44, 45, 47, 48). The anterior margin of the labrum is usually slightly rounded or straight; in Falsotherius and Pherocladus it is concave (Fig. 44).
The ligula is variable in shape, being anteriorly simple, bi-, guadri-, hex-, or multiacuminate. These conditions consist of each half of the ligula forming one, two, three or more apices which are variable in their acuteness (Figs. 15-22, 24, 25). In the multiacuminate condition of Paralichas, each half has four or five branches similar in form to the multiacuminate galea, each branch with a number of apparently random subbranches (Fig.
24). In some forms the ligula is simple and is more or less ventrally and transversely folded (Fig. 23).
Thorax. The lateral margins of the prothorax are either obsolete or acute. The acute condition is characterized by a smooth carina-like ridge which begins at the posterior margin and extends variable distances anteriorly (Figs. 51, 52, 54, 61, 64, 67, 68).
In incompletely acute margins (Figs. 52, 54, 61, 67,
68), the ridge extends anywhere from 1/2 to nearly the entire lateral prothoracic length, the anterior end usually curving somewhat ventrally. Completely acute margins are 24 continuous to the anterior edge (Figs. 51, 64). Lateral margins are either in the same plane as that of the anterior margin (termed, equiplanar. Fig. 67), or approximately mediolateral on the prothorax. Some forms possessing incompletely acute margins appear to be equiplanar, e. g. Stirophora. some Pherocladus. and
Ptiloderes NEW GENUS (Fig. 52); however, the anterior end of the lateral ridge curves ventrally, or, in the case of
Pherocladus is actually dorsal to the anterior margin (Fig.
61), and are not considered equiplanar. Genera possessing equiplanar margins are Daemon and Chelonariomorohus, the former with the lateral and anterior margins continuous and the latter, continuous or very nearly so (Fig. 67).
In genera possessing obsolete lateral margination there
is no smooth carina. Occasionally, e.g. in Bradvtoma.
Aploglossa. and some Anchvtarsus. there may appear to be an acute lateroposterior margin; however, when closely examined, it is seen that the ridge is not smooth and is thus considered obsolete.
The pronotum is anteriorly produced dorsal to the anterior margin in Pherocladus (Fig. 67); Ectvphodactvla has the pronotum smoothly expanded dorsally, laterally, and anteriorly (Fig. 66), the head in repose nearly flush with the margins of anterior prothoracic opening.
The posterior pronotal margin is bisinuate and usually 25 more or less crenulate (Pigs. 69, 71, 72), Chelonariomor- phus and Ptiloderes are exceptions to this with a smooth basal margin (Figs. 70, 73-76). Of those genera possessing crenulate basal margins, Aploglossa, Pseudoepilichas,
Chaetodactvla, and some Bradvtoma and Anchvtarsus have these crenulations very weakly formed. The mediobasal lobe of the hind margin may be smooth or crenulate and usually has three produced denticles (Figs. 69, 73-76). Even with denticles, the mediobasal lobe is considered smooth when there is lack of one or more interdenticular crenulations.
The mediobasal lobe may also be crenulate without produced denticles; this condition is rare and is exhibited only by
Pseudocladotoma and some Paralichas (Fig. 72).
The notai projection is always present, usually long, ventrally acute or truncate (Figs. 61, 62, 64, 66, 67, 68).
In some genera this projection is shorter and usually not acute (Figs. 63, 65). Often present is a posterior and lateral expansion of the prosternum which has been termed
'cowling' (Hlavac, 1975). The cowling often meets the anterior edge of the notai projection causing concealment of the protrochantin. Both conditions, the visibility
(Fig. 62-65) and concealment (Fig. 61, 66-68) of the protrochantin, are widespread among the genera. In
Falsotherius, Ectvphodactvla, and Pherocladus the meeting of the cowling with the notai projection is loose (Pigs. 61, 66).
The anterior margin of the mesoscutellum is associated with the mediobasal lobe of the pronotum in what has been called an interlocking mechanism or system (Hlavac, 1975;
Lawrence, 1982, and personal communication). It takes three forms: smooth (Fig. 70), crenulate (Fig. 72), and cordiform (Pig. 69, 71, 73). The interlocking mechanism is best developed in forms having cordiform mesoscutella in which the median denticle of the basal lobe corresponds with the scutellar notch. Elytra may be strongly or obsoletely striate; either form may possess deeply impressed sutural striae which are usually most evident in the basal one-fifth or one-quarter bounding the mesoscutellum (e. g., Fig. 69). The prosternai process extends posteriorly between the mesocoxae and is usually received in a groove formed by the mesad ridges of the mesocoxal cavities and is not a true excavation. Sometimes a mesosternal excavation is present, it is a depression of the mesosternum into which the prosternai process fits, and may be shallow or relatively deep. A mesosternal keel is best developed in Stirophora and
Chelonariomorphus. It is consistently present but weakly developed in Epiptvqma; rarely, a minor keel is present in Ptilodactvla. The metasternal transverse suture is visible 27 just anterior to the metacoxae, usually most distinct on either side of the discrimen; it less distinct laterally.
In some cases, e. g. Chelonariomorphus, Aploglossa. and
Epiptvqma, the suture is not visible. The metacoxal plates
form the ventral wall of the coxal grooves which serve for reception of the metafemora, and may be very weakly or
relatively well-developed.
The tarsi provide characters involving tarsomere IV,
the ventral surface of the tarsomeres, and the ungual form.
Tarsomere IV is often reduced, sometimes being
inconspicuous. When not reduced, IV may be simple and approximately the same size as III (Fig. 13, 14), or it may have a ventral lobe. Frequently, when tarsomere IV is reduced. III has an expanded ventral lobe (Fig. 12). All ventral tarsomere lobes in Ptilodactylidae are entire and
not bilobed as in some Dascillidae and Rhipiceridae. Tarsi
are termed simple if tarsomere IV is not reduced or lobed. Occasionally, as in Epilichas, Anchvcteis, and Therius, IV
may be lobed. In these and Falsotherius, the remaining
tarsomeres have ventral fleshy pads which occasionally are produced into weak lobes. In Pseudocladotoma there is no
reduction in IV and II - III have short, narrow lobes (Fig.
14). Ongues occur in three forms: simple (Fig. 12-14), bifid
(Fig. 11), and pectinate (Fig. 10). Bifid ungues have two 28 acute points (Fig. 11) or the inner furcation shortened and more or less truncate. In the pectinate ungues of
Paralichas (Fig. 10) each unguis has a mesally-directed row of fine teeth similar to those of alleculines
(Tenebrionidae).
Abdomen. The apex of male visible abdominal sternum V is variable in its shape and may be bisinuate, emarginate or simple. In male Epiptvqma NEW GENUS, visible tergum and sternum VI are sometimes very heavily sclerotized with variously shaped and ventrally-directed tergal flanges
(Figs. 55-60). This segment is very rarely extruded and is usually visible in caudal view.
The aedeagus is of the trilobed type and exhibits much variation. Consistent variation is found in the presence of subapical accessory projections of the median lobe
(Figs. 94). Parameres are movable and may have apical or subapical nonmelanized projections (Figs. 85, 86, 90, 92), and in some cases, e. g. Aploglossa, are very short in relation to the basal piece (Fig. 87, 88).
The ovipositor is usually very long with the bacula + valvulae 3 to 4x the length of the coxites. In some genera the ovipositor is shortened, the length of the bacula + valvulae less than twice that of the coxites (Figs. 77, 78). This condition is most pronounced in Araeopidius (Fig. 78). The coxites may (Figs. 77, 78, 81) or may not 29 (Fig. 19, 80, 82-84) be divided into proximal and distal
sections; styli may be present or absent, and, when present, apical (Fig. 78) or subapical (Figs. 77, 81). In
the Cladotominae, the coxites are completely or almost
completely fused with the bacula and valvulae (Fig. 79, 83,
84). The coxites in Cladotoma are apparently fused with
heavily sclerotized paraprocts, forming a very rigid structure (Fig. 83, 84).
Character polarization
In this section, the character numbers, including those
in parentheses, refer to Table 2.
Char. 1. - Antennal rami. Articulated rami are
postulated to be apomorphic due to unique occurrence within the Ptilodactylidae; antennal rami of the Callirhipidae (as
well as all others known to me) are nonarticulated.
Char. 2. - Epistomal sulcus. Reference to Eulichadidae
and Callirhipidae showing absence of this sulcus indicates
probable apomorphy in its presence.
Char. 3, - Head shape. Occurrence of the basically
quadrate condition in the examined outgroups requires postulation of its plesiomorphy; thus, elongation is
considered apomorphic.
Chars. 5. - Mandibles. Absence of prostheca (4) and
molar areas (5) are coded as apomorphic as a result of 30 presence of the former in Eulichadidae and Callirhipidae and the latter in the Eulichadidae.
Char. 37. - Ligula. In Eulichadidae, the ligula is simple; thus, modification of this structure from the simple condition is coded as apomorphic.
Chars. 6, 26. 27. - Labrum. The short, transverse configuration of the labrum (6) is postulated as apomorphic with the alternative state occurring in Eulichadidae and
Callirhipidae. Other labral modifications, anterior margin concave (26) and presence of anterolateral setal tufts (27) are likewise considered apomorphic due to their absence in the outgroup.
Chars. 23, 24. 31-33. 38. - Maxilla. Presence of distinct lacinial (23) and galeal (24) spore brushes is hypothesized as apomorphic with the absence of these structures in the outgroup. Coded as separate apomorphies are the biacumination of the galea (31) and the reduction of the mesad branch of this condition (33). A further modification is the apomorphic polyacuminate galea (32).
The apical palpomere being elongate and largely membranous
(48) is also considered apomorphic. None of these conditions are found in the Eulichadidae or Callirhipidae.
Chars. 7-10. 29. - Pronotum, Presence of incompletely acute lateral margination (8) is considered apomorphic due to its absence in the Eulichadidae and Psephenidae; thus. 31 the obsolete and completely acute conditions are hypothesized plesiomorphic. The equiplanar margins (9), shortened notai projections (29), concealment of the protrochantins (10), and the anterior pronotal production
(10), not found in any component of the outgroup, are postulated apomorphic» Chars. 11, 28, 34. - Thoracic sterna. Presence of the metasternal transverse suture (11) in eulichadids and callirhipids indicates probable plesiomorphy; therefore, the alternative absence is postulated apomorphic.
Reduction or absence of a mesosternal excavation (28) is considered apomorphic due to the broad and deep condition of that feature in the Eulichadidae. The mesosternal keel
(34) is not found in the outgroup, thus, its presence is apomorphic. Chars. 17-21, 36. - Tarsi. Eulichadidae and
Callirhipidae have simple tarsi and ungues. Any modification is thus hypothesized as apomorphic (17, 20).
The various tarsal modifications, tarsomere IV reduced (19) or Icbed (35), presence of ventral, fleshy pads on two or more tarsomeres (18), and ungues pectinate (21) and basal tooth/bifid (36), are considered apomorphic.
Char. 22. - Sixth visible abdominal segment, males.
The unique occurrence of this heavily sclerotized abdominal segment with a ventrally directed tergal flange and its 32 absence in the outgroup indicates an autapomorphy for
Epiptyqma.
Chars. 12. 25, 30. - Aedeagus. The subapical accessory projections (25) of the median lobe are postulated apomorphic with their absence in the outgroup; presence of nonmelanized parameral projections (12) and short parameres
(30) are also considered apomorphic due to absence of the projections and presence of long parameres in Eulichadidae and Callirhipidae.
Chars. 13-16. 39. - Ovipositor. In the Eulichadidae, the ovipositor is long, the coxites transversely divided and articulated with the bacula, and styli apical; therefore, a shortened ovipositor (16), coxites not divided
(13) or articulated with the bacula (fused; 39), and subapical (15) or absent (14) styli are hypothesized apomorphic.
Discussion of phvlooenetic analysis and resulting classification
Previous classifications of Ptilodactylidae (possibly only implied) (Hlavac, 1975; Lawrence and Newton, 1982) have recognized two or three subfamilies ([Anchytarsinae +
Ptilodactylinae], or [Araeopidiinae + Anchytarsinae +
PtilodactylinaeJ). Crowson (1967) suggested that several subfamilies exist within the Ptilodactylidae. The 33 phylogenetic hypothesis presented (Table 3) supports his statement. Basing groupings only on synapomorphy and by phyletic sequencing (Wiley, 1979), I propose recognition of six subfamilies (Table 4). The numbers in parentheses below refer to characters listed in Table 2 and the phylogenetic hypothesis (Table 3),
The proposed Ptilodactylinae is similar to that formerly recognized and with the addition of Daemonini and
Pseudoepilichini. The former is included on the basis of ungual modification (bifid) (20, 36) and the latter of ovipositor simplification (13, 14) and concealment of protrochantins (10).
The Epilichinae are hypothesized monophyletic on the synapomorphic biacuminate galea (31), a character appearing twice in the proposed phylogeny, also exhibited by the basal Anchytarsinae. The apomorphic shortening of parameres (with a corresponding lengthening of the basal piece) (30) suggests monophyly of Aploglossinae; the
Octoglossinae and Cladotominae are each hypothesized monophyletic on a number of synapomorphiies.
Two unresolved multichotomies occurred in the analysis.
Both consistently appeared in multiple runs of QTREE and could probably be resolved by addition of further characters, especially larval.
Some genera were not included in the phylogenetic 34 analysis due to lack of information, i. e., only having knowledge of one sex (in some cases, that of the type of the generic type species) or only a few specimens in poor condition. In constructing the key I have been able to use characters which I have never observed as sexually dimorphic. Therefore, I have placed those genera in the key and provided descriptions. I will discuss the probable subfamilial or tribal placement for two of them in the phylogenetic diagram, and thus, in the classification. The third Therius is placed incertae sedis due to lack of males and the poor condition of the examined material.
Hovactvla possesses a transverse labrum (6), no molar areas (5) or prostheca (4), and a polyacuroinate galea (32), and is placed in the Cladotominae. pseudocladotoma possesses a transverse labrum (6), no molar areas (5) or prostheca (4), and coxites not articulated with bacula
(39), and is also placed in the Cladotominae. 35
Table 2. Hypothesized polarities of characters used in phylogenetic analysis of the genera of Ptilodactylidae.
Characters States Plesiomorphic Apomorphic 1. Rami non-articulated articulated 2 . Epistomal sulcus absent present 3. Head shape quadrate elongate 4. Prostheca present absent 5. Molar area present absent 6. Labral length long short 7. Pronotum not ant.'ly produced ant.'ly produced 8. Pronotum: lat, marg. not incompl. acute 9. Acute margins not equiplanar equiplanar 10. Protrochantins visible concealed 11. Metasternal trans. suture present absent 12. Parameres w/o non-mel. projs. with 13. Ovipositor: coxites divided not divided 14. Ovipositor: styli present absent 15. Styli present apical subapical 16. Valvifers + baculae:length much greater than coxites less than 2x 17. Tarsi simple not simple 18. Tarsi: ventral aspect w/o fleshy pads w/pads 19. Tarsomere IV not reduced reduced 20. Ungues simple not simple 21. Ungues: not simple not pectinate pectinate 22. Abd. tergum VI (male) w/o vertical flange with flange 23. Lacinia not forming spore brush 24. Spore brush lacinia only lacinia + galea 25. Aedeagus:median lobe w/o access, proj. w/access. proj. 26. Labrum:ant. angles w/o setal tufts w/tufts 27. Labrum not anteriorly concave 28. Mesosternal excavation deep shallow 29. Notai projection long short 30. Parameres long short 31. Galea not biacuminate biacuminate 32. Galea not multiacuminate multiacuminate 33. Galea:biacuminate mesai furc, well-dev. not 34. Mesosternal keel absent present 35. Tarsomere 4:not reduced not lobed lobed 36. Ungues:not simple not bifid or toothed 37. Ligula simple not simple 38. Apical max. palpomere (male) not elong./memb. elong./memb. Coxite/baculum/valvifer fusion no yes 36
Table 3. Phylogenetic hypothesis resulting from QTREE analysis of characters in Table 2. Reversals are indicated by dots at the upper right hand side of the character numbers; characters with multiple apomorphic appearances are 2, 3, 4, 5, 7, 9, 11, 12, 13, 14, 15, 16, 18, 20, 23, 25, 27, 29, 31, 35, and 38.
' i r 38 22 25 27/31 27 38
/ 23 /2 5 34 18 I 33 23 / 27 35 24 28 37' 37" 2 0" 12 3 8" 16 2d 14 10 32 39 14 30 19*
20 36
26
.17 3 7J® 37 Table 4. Proposed supraspecific classification of the
Ptilodactylidae ranked by phyletic sequencing.
PTILODACTYLIDAE
Anchytarsinae ^ ^ Anchvtarsus Guerin-Meneville Cladotominae Cladotoma Westwood Paralichas White Pseudocladotoma Pic Hovactvla Pairmaire
Octoglossinae ^ ^ Octoqlossa Guerin-Meneville Aploglossinae Bradvtoma Guerin-Meneville Aploqlossa Gulrin-Meneville
Epilichinae Anchvcteis Horn Epilichas White Bvrrocrvptus Broun Araeopidius Cockerell
Ptilodactylinae
Daemonini Stenafricus NEW GENÜS Daemon de Laporte
Pseudoepilichini Falsotherius Pic Pseudoepilichas Armstrong and Nakane Ectvphodactvla NEW GENUS
Ptilodactylini Ptiloderes NEW GENUS Pherocladus Pairmaire Lachnodactvla Champion Ptilodactvla Illiger Stirophora Champion Chelonaricmorppus Pic Epiptvqma NEW GENUS CHAPTER V TAXONOMY
KEY TO WORLD GENERA OF PTILODACTYLIDAE (ADULTS)
1. Protrochantins exposed (Figs, 62-65) ...... 2
1*. Protrochantins concealed (Figs. 61, 6 6 -6 8 ) . c , .17
2. Tarsi simple (Fig. 13) ...... 3 2'. Tarsi not simple, tarsomere IV reduced (Fig. 12) or
lobed, II and III lobed (Fig. 14), or 1 or
more tarsomeres with ventral fleshy pads . 7
3. Labrum transverse (Fig. 46), basal width more than 3x
greatest length...... 4
3'. Labrum not transverse (Figs. 44, 45, 47, 48), basal
width approx. 2x greatest length . . . . . 5
4. Ungues pectinate (Fig. 10) ...... Paralichas
4*. Ungues simple (Figs. 12-14)...... Hovactvla
5. Mesoscutellum cordiform (Figs. 1-3, 69, 71, 73), . 6
5'. Mesoscutellum not cordiform, basally smooth (Fig. 70),
or crenulate (Fig. 7 2 ) ...... Anchvtarsus
6 . Lateral prothoracic margination incompletely acute
(Figs. 52, 54, 61, 67, 6 8 ) • . .Bvrrocrvptus
6 *. Lateral prothoracic margination obsolete (Figs. 49,
50, 53, 62, 63, 65, 6 6 )...... Anchvcteis
7. Tarsomere IV reduced (Fig. 12) ...... 8 7'. Tarsomere IV not reduced, usually lobed...... 13
38 39 8 . Mesoscutellum basally smooth (Fig. 70) or crenulate
(Fig. 72)...... 9
8 '. Mesoscutellum cordiform (Figs. 1-3, 69, 71, 73). .12
9. Tarsomeres II and III ventrally lobed...... 10
9'. Tarsomere II not lobed (Figs. 12, 13)...... 11
10. Mandibles bidentate; Oriental region ...... Pseudoepilichas
10'. Mandibles tridentate; Neotropical region...... Bradvtoma
11. Labrum transverse (Pig. 46), lateral prothoracic
margination complete (Figs. 51, 64)......
...... Cladotoma
11'. Labrum not transverse (Pigs. 44, 45, 47, 48),
lateral margination obsolete (Figs. 49, 50,
53, 63, 65, 66)...... Aploqlossa 12. Prothoracic margination equiplanar (Figs. 67),
complete; head quadrate (as in Figs. 52, 54);
Madagascar...... Daemon
12'. Prothoracic margination not equiplanar, obsolete (Fig. 63); head usually elongate (Fig. 49);
Neotropics...... Octoqlossa
13. Epistomal sulcus absent (Figs. 49, 51, 53) . . . .14 13'. Epistomal sulcus present (Figs. 50, 52, 54). , . .15
14. Tarsomere IV ventrally lobed ...... Epilichas 14'. Tarsomere IV not lobed (Fig. 14) . . Pseudocladotoma 40 15. Lateral prothoracic margination obsolete (as in Figs.
63r 65)...... Anchvcteis 15'. Lateral prothoracic margins acute...... 16
16. Ungues simple (Figs. 12-14); southern Africa...... Therius
16'. Ungues with basal teeth (as in Fig. 11 with mesal
furcations more or less truncate); central Africa...... Stenafricus NEW GENUS
17. Mesoscutellum basally smooth (Fig. 70) or crenulate
(Fig. 72)...... 18
17'. Mesoscutellum cordiform (Figs. 2, 3, 71, 73) . . .19
18. Prothoracic margination equiplanar (Fig. 67);
Neotropics...... Chelonariomorphus
18'. Prothoracic margination not equiplanar; Indonesia ...... Falsotherius
19. Prothorax bulbous, lateral margination obsolete, cowling loosely meeting notai projection
(Fig. 53, 6 6 ). . . .Ectvphodactvla NEW GENUS
19'. Prothorax not bulbous, lateral margination incompletely acute, cowling tightly fitting
against notai projection (Figs. 61, 6 8 ) . 20
20. With mesosternal keel...... 21
20'. Without mesosternal k e e l ...... 22 21. Dorsum usually glabrous, very shining, median mesoscutellary notch usually continuing as 41 distinct stride; males, tergum of visible
abdominal segment VI with vent rally-directed flange (Pigs. 55-60). . .Epiptvqma NEW GENUS
21*. Dorsum covered with dense, recumbant setae, or,
elytra very strongly striate; males, tergal
flange absent; aedeagus, median lobe with
subapical accessory projections (Fig. 94}
...... Stirophora 22. LateropeSterior pronotal angles broadly explanate
(Figs. 3, 52, 73-76); galeae not forming
spore brush (Fig. 34) .Ptiloderes NEW GENUS
22®. Lateroposterior pronotal angles not broadly
explanate (Pigs. 2, 54); galeae forming spore
brush (Fig. 27, and as in Fig. 31). . • .23
23. Pronotum with strong anterior production (Fig. 61);
Indonesia, Philippines . • • • .Pherocladus
23®. Pronotum without strong anterior production (Fig.
6 8 ); cosmopolitan ...... 24
24. Apical maxillary palpomere, males, enlarged,
elongate, and largely membranous (Fig. 27),
females, usually somewhat elongate and mostly
sclerotized ...... Lachnodactvla
24®. Apical maxillary palpomere simple, securiform (as in
Pig. 31)...... Ptilodactvla THE WORLD GENERA OF PTILODACTYLIDAE 42
Anchvtarsus
Anchvtarsus Guerin-Meneville, 1843a: 194 (type species, by
monotypy: Atopa bicolor Melsheimer, 1845: 221);
Blanchard, 1845: 56; Guerin-Meneville, 1849: 1;
Lacordaire, 1857: 264; LeConte, 1853: 229, 1861: 179,
1863-1866 (1865): 50; Horn, 1880: 8 6 , 1881: 87; LeConte
and Horn, 1883: 170; Champion, 1897: 593; Blatchley,
1910: 689; Bertrand, 1935: 138, 1939: 307, 1956: 275,
1966: 143, 1972: 391; Arnett, 1968: 443; Spangler,
1966: 397, 1981: 208, 1982: 386, 1983: 161; Brown,
1972: 24; 1975; 149; Hlavac, 1975: 182; Doyen and Ulrich, 1978: 229; White, 1980: 94; Stribling, 1986:
232. Tetraalossa Champion, 1897: 593 (type species, by monotypy: T, palpalis Champion, 1897: 594); Bertrand, 1956: 278,
1972: 393; Brown, 1975: 150; Stribling, 1986: 220.
I have recently redescribed this genus (Stribling,
1986) and will here give a synopsis of characters not pointed out in that paper as well as a diagnosis. Molar areas and dorsal ridge are present and well-developed. The apical maxillary palpomeres are simple and largely
sclerotized. The pronotal mediobasal lobe and mesoscutellum are either crenulate or smooth (as in Figs. 43 70 or 72). Sutural striae are deeply impressed and more evident than other striae near the mesoscutellum.
Diagnostic combination. This genus may be recognized by the following combination of characters: simple tarsi
(as in Fig. 13), protrochantins visible (as in Pig. 65), epistomal sulcus absent, lateral pronotal margination obsolete, male antennae serrate (as in Fig. 4), parameres with apical nonmelanized projections, and ovipositor with styli.
MATERIAL EXAMINED: 104 specimens (AMNH, MNHP, BMNH, CASC,
CNCI, EGRC, FMNH, ICCM, MSDC, MCZC, NMNH, NCSD, OSDC, SMSH,
JBSC, DMCE, OMIS).
Taxonomic list - Anchvtarsus
*bicolor (Melsheimer), 1845 (Atopa) (MCZC); eastern North
America *folliculipalpus Stribling, 1986 (CNCI); Central, S. America *instriatus Pic, 1931b (MNHP); Costa Rica
♦palpalis (Champion), 1897a (Tetraalossa) (BMNH); Mex.,
Cent., S. Amer.
♦substriatus Champion, 1897a (BMNH); Nicaragua Cladotoma 44
Cladotoma Westwood, 1837: 254 (type species: Ç. ovalis
Westwood 1837); Guerin-Meneville, 1843b: 194; 1843a: 1;
Blanchard, 1845: 56; Lacordaire, 1857: 276; Champion,
1897a: 626; Stribling, 1986.
Telon Champion, 1897a: 626 (type species: T. cucullatus
Champion 1897a: 626); Stribling, 1986: 232. NEW
SYNONYMY.
These two genera are synonymized due to joint possession of apomorphic pseudotetramery (Fig. 12) within the Cladotominae and the coxites of the ovipositor nonarticulated with the bacula (Figs. 83, 84).
Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margination complete (as in Fig. 64), labrum transverse (Fig. 46), tarsi (Fig. 12) tarsomere IV reduced,
III ventrally lobed, and simple ungues.
Description.
Body Size. Length 10.0-15.0 mm, width at humeral angles 3.5-5.0 mm.
Head. Quadrate; antennae, males with nonarticulated, broad, dorsoventrally flattened rami, antennomeres III-X
(Fig. 6 ), females usually strongly serrate, sometimes with much shorter rami formed as in males; epistomal sulcus 45 absent; labrum strongly transverse (Fig. 46); mandibles (as in Fig. 39) bidentate, without prostheca, molar area, or dorsal ridge; maxillae (as in Fig. 26), lacinia acute, galea polyacuminate; labium, ligula (Fig. 22) quadracuminate.
Thorax. Prothorax (as in Fig. 64) somewhat anteriorly produced, with complete, acute mediolateral margination
(Fig. 51, as in 64), strong basal crenulation, mediobasal lobe crenulate, with produced denticles, mesoscutellum crenulate (as in Fig. 72); notai projection very long, thin, trochantins visible (as in Fig. 64); mesosternal keel absent; elytra obsoletely striate, sutural striae not more deeply impressed; metasternal transverse suture visible near"discrimen, laterally obsolete; metacoxal plates well- developed; tarsi (Fig. 12), article IV reduced. III ventrally lobed, no fleshy pads, ungues simple.
Abdomen. Male sternum V modified, sinuate; aedeagus (Fig. 93), median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (Pigs. 83, 84) long, coxites not divided, not articulated with bacula, fused with heavily sclerotized paraprocts, styli absent.
Species excluded: russula Pic (Cerophytidae), vittata Pic
fPseudolichas: Dascillidae). 46 MATERIAL EXAMINED: 26 specimens (OSDC, EGRC, KJOC, CNCI,
MNHP, FMNH, INHS, BMNH).
Taxonomic list - Cladotoma
bruchii Pairmaire, 1904 (BANM); Argentina
*maculicollis Pairmaire, 1904 (MNHP); Argentina
*heptosa (NEW NAME FOR maculicollis Pic, 1911 [MNHP]);
Brazil (Rio Verde)
*marqinata Pic, 1933 (MNHP); Brazil
♦ovalis Westwood, 1837 (HEMC); Brazil
subvittata Guerin-Meneville, 1861; Brazil
thoracica Guerin-Meneville, 1843a; French Guiana Paralichas 4 7
Paralichas White, 1859; 287 (type species, by monotypy: P.
querini White 1859: 287); Nakane, 1948: 5, 1956: 52.
Eucteis Guerin-Meneville, 1861: 539 (type species, by
monotypy: E. bimaculata Guérin-Meneville, 1861: 539);
Pairmaire, 1886: 395; Lewis, 1895: 99.
Paralvchus Pairmaire, 1886: 395 (unjustified emendation).
Odontonvx Guérin-Meneville, 1843b: 194 (not Stephens, 1827
in Carabidae; type species, by monotypy: Atopa ornata
Melsheimer, 1845: 220 (transferred by Guérin-Meneville
(1843b) [ =Dasvtes trivittis Germar, 1824: 76,
Lacordaire (1857: 267)1; Blanchard, 1845: 56; Guérin-
Meneville, 1849: 1; LeConte, 1861: 179; Horn, 1880: 8 6 ; LeConte and Horn, 1883: 170; Blatchley, 1910; 689;
Arnett, 1968: 443; Hlavac, 1975: 182; Stribling, 1986:
232. NEW SYNONYMY.
This synonymy is based on joint possession of the apomorphic pectinate ungues (Pig. 10).
Diagnostic combination. This genus may be recognized
by the autapomorphic pectinate ungues.
Description.
Body Size. Length 7.5-11.0 mm, width at humeral angles
3o5-5o0 mm.
Head. Quadrate; antennae, males often with 48 nonarticulated rami, antennomeres III-X (Fig. 5), females, sometimes males, serrate; epistomal sulcus absent; labrum
(as in Fig. 46) strongly transverse, anteriorly truncate or slightly convex; mandibles (Fig. 39) bidentate, without prostheca, molar areas, or dorsal ridge; maxillae (Fig.
26), galea multiacuminate, apical palpomere mostly sclerotized and melanized; labium (Fig. 24), ligula multiacuminate.
Thorax. Prothorax (Fig. 64) somewhat anteriorly produced with complete, acute mediolateral margination, strong basal crenulation, mediobasal lobe crenulate, with or without produced denticles (Fig. 72); mesoscutellum crenulate (Fig, 72); notai projection very long, thin, trochantins visible (Fig. 64); mesosternal keel absent; elytra strongly striate, sutural striae not more deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well- developed; tarsi (as in Fig. 13) simple, without ventral fleshy pads; ungues pectinate (Fig. 10).
Abdomen. Male sternum V simple or modified, when modified, sinuate; aedeagus (Fig. 91), median lobe without mediolateral flanges and subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (Fig. 79) long, coxites not divided, not articulated with bacula; styli absent. 49 Species excluded: pjceiceps Pic (Pseudocladotoma\
MATERIAL EXAMINED: 136 specimens (ICCM, RHTC, MCZC, RÜIC, OSDC, KOBC, MNHP, NMNH, INHS, CASC).
There are two undescribed Neotropical species, one specimen each from Mexico and El Salvador.
Taxonomic list - Paralichas *bicolorpes Pic, 1911 (MNHP); Taiwan
*querini White, 1859 (BMNH); China Eucteis bimaculata Guerin-Meneville, 1861
Lewis, 1895 (synonymized with querini)
V . *disconiqer Pic, 1939 (MNHP); China
V . *diverselineatus Pic, 1939 (MNHP); China
V . *iunctus Pic, 1939 (MNHP); China
*qriseolineatus Pic, 1939 (MNHP); no locality
*hiqoniae Lewis, 1895 (MNHP); Japan ssp. qvotokui Nakane, 1963; Japan
niqer Pic, 1931a (MNHP); Madagascar *pectinata (Kiesenwetter), 1874 (Odontonvx) ; China, Japan
rufolimbata (Pairmaire), 1886 (Eucteis); Kiang-si, China
V . *apicornis Pic, 1939 (MNHP); China
*striolatus Pairmaire, 1897 (MNHP); Sze-tchouen
*subnitidus Pic, 1911 (MNHP); China *trivittus (Germar), 1824 (Dasvtes); eastern N. America
Atopa ornata Melsheimer, 1845 Guerin-Meneville, 1849 (transferred ornata to
Odontonvx) Hovactvla
Hovactvla Pairmaire, 1901: 174 (type species, by monotypy: H. dermestoides Pairmaire, 1901: 175).
Diagnostic combination. This genus may be recognized by the following combination of characters: labrum transverse (as in Pig. 46) and tarsi and ungues simple (as in Pig. 13).
Description.
Body size. Length 6.5-9.0 mm, width at humeral angles 2.5-3.0 mm. Head. Quadrate; antennae serrate (as in Pig. 4); epistomal sulcus absent, labrum transverse (as in Pig. 46), basal width approx. 4x greatest length, anteriorly convex; mandibles (as in Pig. 39) bidentate without prostheca or molar areas, with dorsal ridge; maxillae, lacinia acute not forming spore brush, galea multiacuminate (as in Pig. 26); apical maxillary palpomere mostly sclerotized; labium,
ligula quadracuminate.
Thorax. Pronotum somewhat anteriorly produced, with
mediolateral margins completely acute (as in Figs. 51, 64),
strong basal crenulation, mediobasal lobe crenulate, with produced denticles (as in Pig. 72), notai projection long,
protrochantins visible (as in Pig. 64); mesoscutellum
basally crenulate (as in Pig. 72), mesosternal excavation 51 deep, keel absent; elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete, coxal plates underdeveloped; tarsi simple (as in Fig. 13), without fleshy ventral pads, ungues simple.
Abdomen. Male sternum V slightly sinuate; aedeagus, median lobe without mediolateral spines or subapical accessory projections, parameres without nonmelanized projections. Females unknown.
MATERIAL EXAMINED: 2 specimens (MNHP, CNCI).
Specimens from the type locality and South Africa. The latter represents an undescribed species.
Species excluded: rufescens Pic (incertae sedis).
Taxonomic list - Hovactvla
*dermestoides Pairmaire, 1901 (MNHP); Madagascar 52
Pseudocladotoma
Pseudocladotoma Pic, 1918: 3 (type species, by monotypy: P. maculata Pic, 1918: 4).
Diagnostic combination. This genus may be recognized by the following combination of characters: labrum transverse, complete lateral margination, serrate antennae, simple ungues and, tarsomere IV not reduced.
Description.
Body size. Length 6.5-9.S mm, width at humeral angles
2.0-3.0 mm. Head. Quadrate; epistomal suture absent; labrum transverse (as in Fig. 46), basal width approx. 3x greatest length, anteriorly convex; mandibles monodentate (as in Fig. 40), without prostheca or molar areas (as in Fig. 39), with dorsal ridge; maxillae, lacinia acute, not forming spore brush, galea blunt; apical palpomere mostly sclerotized; labium, ligula biacuminate (as in Fig. 18).
Thorax. Lateral margins completely acute (as in Fig.
64), somewhat anteriorly produced; basally crenulate, mediobasal lobe crenulate without produced teeth; notai projection long, protrochantins visible (as in Fig. 64); mesosternal excavation, keel absent, mesoscutellum basally crenulate (as in Fig. 72); elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture 53 visible near discrimen, laterally obsolete; metacoxal plates underdeveloped; tarsi (Fig. 14} robust, article IV not reduced, II-III sometimes with very narrow ventral lobes, without ventral fleshy pads, ungues simple. Abdomen. Ovipositor long, coxites not divided, not articulated with bacula, styli absent.
Males unknown.
MATERIAL EXAMINED: 4 specimens (MNHP).
Taxonomic List - Pseudocladotoma
♦maculata Pic, 1918 (MNHP); Java
♦piceiceps (Pic), 1930a (Paralichas) NEW
COMBINATION (MNHP); China ♦rufonotatus (Pic), 1931b (Epilichas) NEW
COMBINATION (MNHP); Borneo ♦vitaticollis (Pic), 1930a (Epilichas) NEW
COMBINATION (MNHP); China 54 Octoqlossa
Octoqlossa Guerin-Meneville, 1843a: 1 (type species, by
monotypy: Q» femoralis Guerin-Meneville, 1843a: 2);
Guerin-Meneville, 1843b: 194; Lacordaire, 1857: 268;
Stribling, 1986: 232.
Astvloqlossa Pic, 1919: 6 (type species, by monotypy: A.
coerulipennis Pic, 1919: 7); Stribling, 1986. NEW
SYNONYMY.
These two genera are synonymized due to the synapomor phic elongation of the head, lack of molar areas, oviposi tors with subapical styli and, shortened notai projections.
Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margination obsolete, head elongate, trochantins visible (Figs. 49, 63), tarsomere IV reduced (as in Fig. 12 but somewhat distally expanded), II-III lobed ventrally, mesoscutellum cordiform. These species often have dorsal elytral patterns of metallic blue and green.
Description. Body size. Length 10.5-16.0 mm, width at humeral angles 2.5-4.5 mm.
Head. Elongate (Fig. 49; less so in 0. rufa): antennae serrate (as in Fig. 4), epistomal sulcus absent, labrum long, basal width less than 2x greatest length, anteriorly convex; mandibles (Fig. 40) long, monodentate, with prostheca, without dorsal ridge or molar areas; maxillae, galea acute, sometimes multiacuminate, lacinia acute, not forming spore brush; apical palpomere mostly sclerotized; labium, ligula hexacuminate (Fig. 25).
Thorax. Prothorax with obsolete lateral margins (Fig.
63); basal margins strongly crenulate, mediobasal lobe crenulate, with produced denticles (as in Fig. 71); notai projection short, protrochantins visible (Fig. 63); mesoscutellum cordiform; mesosternal excavation shallow, keel absent; elytra strongly striate, sutural striae not
deeply impressed; metasternal transverse suture visible near discrimen, laterlly obsolete, coxal plates underdeveloped; tarsi, article IV reduced (as in Fig. 12 but somewhat distally expanded), II-III lobed, without
ventral fleshy pads, ungues simple.
Abdomen. Male sternum V emarginate; aedeagus, median
lobe without mediolateral flanges or subapical accessory
projections; ovipositor (as in Fig. 77) short, bacula and
valvulae less than 2 x coxites, coxites divided, articulated
with bacula, styli subapical.
MATERIAL EXAMINED: 20 specimens (MCZC, BMNH, ANSP, MNHP,
NMNH, CNCI).
Specimens were examined from Panama, Costa Rica, Colombia, Ecuador, and Venezuela.
Taxonomie list - Octoqlossa
*coerulipennis Pic, 1919 (Astvloqlossa) NEW
COMBINATION (MNHP); Colombia
*cvaneipennis Pic, 1924a (MNHP); Costa Rica
femoralis Guérin-Meneville, 1843b; Panama, Colombia
*rufa Pic, 1924b (MNHP); Colombia Bradvtoma
Bradvtoma Guerin-Meneville, 1843a: 1 (type species: B.
aurita Guérin-Meneville, 1843a: 1, by monotypy);
Guerin-Meneville, 1843b: 194; Blanchard, 1845: 56;
Lacordaire, 1857: 277; Stribling, 1986: 232.
Brithvcera Erichson, 1847: 175 (type species: B. Ivciformis
Erichson, 1847: 175, by monotypy), 1848: 102;
Lacordaire, 1857: 276.
Diagnostic combination. This genus may be recognized by the following combination of characters: tarsomere IV reduced, II-III ventrally lobed, without ventral fleshy pads, labrum not anteriorly concave, lateral pronotal margination obsolete, male antennae serrate (Fig. 4), mesoscutellum basally smooth (as in Fig. 70) or crenulate
(as in Fig. 72), and ligula not simple, with variable acumination (Fig. 16).
Description.
Body size. Length 6.5-10.0 mm, width at humeral angles
2.0-3.5 mm.
Head. Quadrate; antennae serrate (Fig. 4); epistomal sulcus present or absent; labrum long, basal width 2 x or less greatest length (as in Fig. 47), anteriorly truncate or slightly convex; mandibles tridentate, with prostheca, 58 molar area, and dorsal ridge (as in Fig. 37); maxillae
(Fig. 33), lacinia blunt, not forming spore brush, galea cylindrical, with apical cluster of setae, not closely associated with lacinia; apical palpomere mostly sclero- tized; labium, ligula modified (Fig. 16), hexacuminate with varying degrees of reduction.
Thorax. Prothoracic margins (Pig. 65) obsolete, occasionally appearing posteriorly acute but margins without anterior, ventral curvature or smooth carina; basally crenulate, mediobasal lobe crenulate or not, with or without produced denticles; notai projection long, trochantins visible; mesosternal keel, excavation absent; mesoscutellum basally smooth, elytra strongly striate, sutural striae deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates weakly developed; tarsi, article IV reduced, II-III lobed, without ventral fleshy pads; ungues simple (as in Fig. 12).
Abdomen. Male sternum V simple or modified, when modified, sinuate; aedeagus (Figs. 85, 8 6 ), median lobe without mediolateral spines, parameres with nonmelanized projections, ovipositor (as in Fig. 81) long, coxites not divided, articulated with bacula, styli subapical.
MATERIAL EXAMINED: 12 specimens (MNHP, CNCI, RSN^ ). 55 Taxonomic list - Bradvtoma
*aurita Guerin-Meneville, 1843; Brasil
*lineata Pic, 1921 (MNHP); Brasil
Ivciformis (Erichson), 1847 (Brithvcera); Peru Aploalossa 60
Aploqlossa Guerin-Meneville, 1849; 1 (type species, here
designated: A. sallei Guerin-Meneville, 1849: 2
[ =Aploqlossa marqinata Guerin-Meneville, 1849: 2]);
Lacordaire, 1857: 276; Kasap and Crowson, 1975; 442;
Stribling 1986: 232.
Haploqlossa Gemminger and von Harold, 1869: 1623
(unjustified emendation); Champion, 1897: 623; Kasap
and Crowson, 1975: 442.
Diagnostic combination. This genus may be recognized
by the following combination of characters: tarsi (as in
Pig. 12), tarsomere IV reduced. III lobed ventrally,
epistomal sulcus present, protrochantins visible, obsolete
lateral margination (as in Fig. 65), male antennae serrate
(as in Fig. 4), mesoscutellum basally smooth (as in Fig.
70), and simple ligula.
Description.
Body size. Length 4.5-7.5 mm, width at humeral angles
1.0-2.5 mm.
Head. Quadrate; antennae (as in Fig. 4), serrate;
epistomal sulcus present; labrum (Fig. 45), not transverse,
anterior margin more or less concave, with tufts cf setae
at anterolateral angles; mandibles tridentate, with
prostheca, molar areas, and without dorsal ridge (Fig. 37); Cl maxillae (Fig. 31), lacinia forming spore brush, galea closely associated with spore brush, but, with longer, thinner, and less dense setae, apical palpomere mostly sclerotized; labium (as in Fig. 23), ligula simple.
Thorax. Prothorax (as in Fig. 65) with obsolete lateral margins, sometimes appearing posteriorly acute, but lacking smooth ridge, usually more or less angulate at posterior one-fifth; weak basal crenulation, mediobasal lobe crenulate, without produced denticles; mesoscutellum basally smooth (as in Fig. 70); notai projection short, trochantins visible (as in Fig. 65); mesosternal excavation shallow, keel absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture absent; metacoxal plates underdeveloped; tarsi (as in Fig.
12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues simple.
Abdomen. Male visible sternum V modified, emarginate; aedeagus, median lobe without mediolateral flanges, sometimes with subapical accessory projections (as in Fig.
94), parameres short, without nonmelanized apical projections (Pigs. 87, 88); ovipositor (Fig. 77) short, valvifers less than 2x coxites, coxites articulated with bacula; divided, with subapical styli.
MATERIAL EXAMINED: 145 specimens (NMNH, AMNH, CASC, BMNH,
CNCI, MAIC, FMNH, MNHP, JBSC, UNTC). 69 Taxonomie list - Aploqlossa
*anqustata Champion, 1897a (BMNE); Panama
*aureonotata Pic, 1913d (MNHP); Peru
V . qermaini Pic, 1927b (DEIE); Bolivia
*baeri Pic, 1913d (MNHP); Peru
*boliviensis Pic, 1913d (MNHP); Bolivia
*collaris Guérin-Meneville, 1849; Izabal, Guatemala
Champion, 1897a (redescription)
V . *reducta Pic, 1934e (MNHP); no locality
V . *testaceicollis Pic, 1913d (MNHP); no locality
*ferruqinea Champion, 1897a (BMNH); Guatemala
*laiovei Pic, 1913d (MNHP); Venezuela
sallei Guerin-Meneville, 1849; Caracas, Venezuela
marqinata Guérin-Meneville, 1849
*scutellaris Pic, 1913d (MNHP); Neotropics
V . *niqricollis Pic, 1913d (MNHP); Brazil
V . *obscuricolor Pic, 1913d (MNHP); Amazon
*suturalis Pic, 1934c (MNHP); Bolivia
V . *diversipes Pic, 1934c (MNHP); no locality
V , *humeralis Pic, 1934c (MNHP); no locality
V . *notaticollis Pic, 1934c (MNHP); no locality
*testaceipes Pic, 1934c (MNHP); Peru Anchvcteis
Anchvcteis Horn, 1880: 87 (type species, by monotypy: A.
velutina Horn, 1880: 87); LeConte and Horn, 1883: 170;
Leech and Chandler, 1956: 367; Arnett, 1968: 443;
Hlavac, 1975: 182; Doyen and Ulrich, 1978: 229;
Stribling, 1986: 232.
Amphicteis Bertrand, (1966: 144, 1972: 392; incorrect
subsequent spelling).
Diagnostic combination. This genus may be recognized by the following combination of characters: male antennae with nonarticulated rami (Pig. 1), epistomal sulcus present but shallow, lateral pronotal margins obsolete, protrochan tins visible (as in Fig. 65), tarsomeres I-IV with ventral fleshy pads, IV sometimes lobed, and molar areas present
(Fig. 42).
Description.
Body size. Length 9.5-11.0 mm, width at humeral angles
2.5-3.5 mm.
Head. Quadrate; antennae, males with non-articulated rami, antennomeres III-X (Figs. 1, 5), females serrate (as in Fig. 4); epistomal sulcus present, shallow; labrum (Fig.
48), not transverse, anteriorly convex; mandibles (Fig. 42) monodentate, with prostheca and molar areas, with or without dorsal ridge; maxillae (Fig, 29), lacinia blunt. 64 not forming spore brush, galea apically acute, biacuminate with mesal branch highly reduced or absent; apical palpomere mostly sclerotized and melanized, not exceptionally flattened or expanded; labium (Fig. 19), ligula quadracuminate with lateral branches highly reduced.
Thorax. Prothorax (as in Fig. 65) with lateral margination obsolete, basal margin strongly crenulate, mediobasal lobe crenulate, with produced denticles (as in
Fig. 71); mesoscutellum cordiform; notai projection long, trochantins visible (as in Fig. 65); mesosternal keel absent, excavation present, shallow; elytra strongly striate, sutural striae not more deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsomere IV not reduced, I-IV with ventral fleshy pads sometimes being drawn into small lobes; ungues simple.
Abdomen. Male visible sternum V modified, sinuate; aedeagus with mediolateral flanges, without subapical accessory projections; parameres long, without nonmelanized apical projections; ovipositor long, coxites articulated with bacula, divided or not, styli present or absent, when present, subapical.
MATERIAL EXAMINED: 56 specimens (UCDC, FMNH, OSUO, MCZC,
OSUC, UMCE, MNHP, BMNE, KÜBC, CASC, NMNH, CNCI, GLPC). 65 Taxonomie list - Anchvcteis.
*brunneicornis (Lewis), 1895 (Epilichas) NEW
COMBINATION (BMNH); Japan
ssp. usori (Nakane), 1958; Japan
*monticola (Nakane), 1952 (Epilichas) NEW
COMBINATION; Japan
*velutina Horn, 1880 (MCZC); northwestern ü. S. A, Epilichas
Epilichas White, 1859: 290 (type species, by monotypy: E.
candezei White, 1859; 289); Nakane, 1948: 5, 1956: 52;
Crowson, 1967: 50.
Diagnostic combination. This genus may be recognized by the following combination of characters: male antennae with nonarticulated rami (as in Figs. 1, 5), epistomal sulcus absent, lateral margination obsolete, protrochantin visible (as in Fig. 65), tarsomere IV lobed, I-III with ventral fleshy pads, and molar areas absent (Fig. 43).
Description.
Body size. Length 8.5-12.5 ram, width at humeral angles
2.5-3.5 mm.
Head. Quadrate; antennae, males with non-articulated rami on antennomeres III-X (as in Figs. 1, 5), females serrate (as in Fig. 4); epistomal sulcus absent; labrum (as in Fig. 48), not transverse, anteriorly truncate or convex; mandibles (Fig. 43) monodentate, with prostheca, without molar areas, with or without dorsal ridge; maxillae (Fig.
28), lacinia blunt, not forming spore brush, galea apically acute, biacuminate; apical palpomere mostly sclerotized, usually flattened and laterally expanded; labium (Fig. 21), ligula quadracuminate.
Thorax. Prothorax (as in Fig. 63) with lateral margination obsolete, basal margin strongly crenulate, mediobasal lobe crenulate, with produced teeth (as in Fig.
71); mesoscutellum cordiform; notai projection long, trochantins visible (as in Fig, 64); mesosternal keel absent, excavation shallow; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi, article IV lobed, articles I-III with ventral fleshy pads occasionally being drawn into small lobes; ungues simple.
Abdomen. Male visible sternum V modified, sinuate; aedeagus (Fig. 95), median lobe without mediolateral spines or subapical accessory projections; parameres, long, without apical flange or nonmelanized apical projections; ovipositor, long, coxites articulated with bacula, divided, with apical styli (as in Fig. 78 but bacula 4-5x coxital length).
Species excluded: brunneicornis Lewis (Anchvcteis), monticola Nakane (Anchvcteis), rufonotatus Pic
(Pseudocladotoma), varieqatus Carter (Bvrrocrvptus), vittaticollis Pic (Pseudocladotoma), apicicornis Pic
(Falsotherius).
MATERIAL EXAMINED: 46 specimens (MNHP, BMNH, NMNH,
CNCI, KUBC). 68 Taxonomie list - Epilichas
*atricolor Lewis, 1895 (BMNE); Japan
basipes Pic, 1927d (MNHP); Vietnam (Tonkin)
*candezei White, 1859 (BMNH); China
*flabellatus (Kiesenwetter), 1874 (Octoqlossa)r Japan
*testaceipes Pic, 1927a (MNHP)
Nakane, 1952 (synonymized with flabellatus)
V . amamianus Nakane, 1963; Japan
ssp. mutsuensis Nakane, 1958; Japan
ssp. vakushimensis Nakane, 1952; Japan
f, oshiïïianus Nakane, 1952; Japan
V . rubricollis Nakane, 1952; Japan
ssp. tamaii Sato, 1964; Iriomote-jima, Sakishima
mivatakei Nakane, 1952; Japan
niqrinis Carter, 1936; Australia
*niaripes Pic, 1914b; Vietnam (Tonkin)
V . testaceicollis Pic, 1914b; Vietnam
oblonqus (Carter), 1935 (Dascillus) (ANMM); Australia
obscurus (Pic), 1923a (Eulichas): Vietnam (Tonkin)
Deleve- 1972 (redescription)
rufescens Pic, 1927c; Vietnam (Tonkin)
serraticornis Carter, 1935; Australia Bvrrocrvptus
Bvrrocrvptus Broun, 1893: 1136 (type species, by monotypy:
Bvrrocrvptus urouharti Broun, 1893: 1137); Kasap and
Crowson, 1975: 442.
Diagnostic combination. This genus may be recognized by the following combination of characters: antennae serrate (as in Fig. 4), epistomal sulcus present, lateral pronotal margination incompletely acute, protrochantins visible (as in Pig. 65), mandibles bidentate (as in Fig.
41), and simple tarsi (as in Fig. 13).
Description.
Body size. Length 5.0-10.0 mm, width at humeral angles
1.5-3.5 mm.
Head. Quadrate; antennae serrate (as in Fig. 4); epistomal sulcus present; labrum (as in Fig. 47), basal width approx. 2x greatest length, anterior margin truncate or slightly'convex; mandibles (as in Fig. 41) bidentate, prostheca, molar areas, and dorsal ridge present; maxillae, lacinia blunt, not forming spore brush, galea (Fig. 30) biacuminate, mesal branch sometimes highly reduced, apical palpomere mostly sclerotized and melanized; labium (as in
Fig. 20), ligula quadracuminate.
Thorax. Prothorax with acute lateral margination, incomplete; basal margin strongly crenulate, mediobasal 70 lobe crenulate, with produced denticles (as in Fig. 71),
notai projection long, trochantins visible (as in Fig. 64);
mesoscutellum cordiform; sternal excavation shallow, keel
absent; elytra strongly striate, pronotum and elytra
sometimes with broad, irregular punctation, sutural striae
usually deeply impressed basally; metasternal transverse
suture visible near discrimen, laterally obsolete;
metacoxal plates well-developed; tarsi (as in Fig. 13)
simple without fleshy pads, ungues simple.
Abdomen. Male sternum V simple or modified, when
modified, sinuate; aedeagus, median lobe without
mediolateral flanges or subapical accessory projections;
parameres long, wtihout nonmelanized apical projections;
ovipositor, long, articulated with bacula, coxites divided,
with apical styli (as in Fig. 78 with length of bacula 4-5x
that of coxites).
MATERIAL EXAMINED: 136 specimens (CSIR, CNCI, BMNH,
JBSC).
Taxonomic list - Bvrrocrvptus
*urauharti Broun, 1893 (BMNH); New Zealand
*varieaatus (Carter), 1935 NEW COMBINATION (Epilichas)
(ANMM); N. Queensland
There are about 6 species with additional records from
Papua, New Guinea and New South Wales. It is probable that 71 the other species of Epilichas described by Carter (1935) belong here; however, examination of the types is necessary. Araeopidius 7 2
Araeopus LeConte, 1874: 56, not Spinola (1839) in Hemiptera;
Horn, 1880: 88; LeConte and Horn, 1883: 170.
Araeopidius Cockerell, 1906: 241 (type species: by
monotypy: Araeopus monachus LeConte, 1874: 56); Hlavac,
1975: 182; Lawrence and Newton, 1982: 278; Stribling,
1986: 232.
Diagnostic combination. This genus may be recognized by the following combination of characters; head somewhat elongate (Fig. 50), lateral pronotal margins obsolete, protrochantins visible (Fig. 62), mesoscutellum cordiform
(Fig. 69), tarsi simple (Fig. 13), and sutural striae deeply impressed (Fig. 69).
Description.
Body size. Length 9.0-12.5 mm, width at humeral angles
2.0-3.5 mm.
Head. Elongate (Fig. 50); antennae serrate (Fig. 4); epistomal sulcus shallow, labrum long, anteriorly truncate or slightly concave; mandibles (as in Pig. 42) monodentate, with prostheca, molar areas, and dorsal ridge; maxillae, lacinia acute (as in Fig. 29), not forming spore brush, galea biacuminate (as in Fig. 28); apical palpomere mostly sclerotized, somewhat elongate; labium, ligula quadracuminate (as in Fig. 21). 73 Thorax. Pronotum with obsolete lateral margins (Fig.
62)/ strong basal crenulation, mediobasal lobe crenulate, with produced denticles (Fig. 69); notai projection long, protrochantins visible (Fig. 62); mesoscutellum cordiform, mesosternal excavation deep, keel absent, elytra obsoletely striate, sutural striae deeply impressed (Fig. 69); metasternal transverse suture visible near discrimen, laterally obsolete, coxal plates well-developed; tarsi
(Fig. 13) simple, with reduced onychium, without ventral fleshy pads, ungues simple.
Abdomen. Male sternum V simple; aedeagus, median lobe without mediolateral flanges and subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (Fig. 78) short, bacula and valvulae slightly longer than coxites, coxites articulated with bacula, divided, styli apical.
MATERIAL EXAMINED: 175 specimens (GLPC, OSUO, COBC, INKS,
FMNH, MCZC, CASC, OSUC).
Taxonomic list - Araeopidius
♦monachus (LeConte), 1874 (Araeopus); northwestern North
America 7 4
Stenafricus NEW GENUS
Stenafricus NEW GENUS (type species, present designation:
Stenactvla auberti Pic, 1925b: 14).
Stenactvla Fairmaire, 1904: 346, in part.
Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margins acute, nearly complete, notai projection short (as in Fig. 63), protrochantins visible (as in Fig.
63), tarsomere IV lobed, no ventral fleshy pads, ungues with basal tooth, and lacinia and galea forming spore brush
(Fig. 32).
Description.
Body size. Length 4.0-7.0 mm, width at humeral angles
1.0-2.0 mm.
' Head. Quadrate; antennae serrate (as in Fig. 4); epistomal sulcus present; labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly convex; mandibles tridentate, with prostheca and molar areas (as in Fig. 37), without dorsal ridge; maxillae (Fig. 32), galea and lacinia forming spore brush, apical palpomere mostly sclerotized; labium, ligula simple, often somewhat ventrally, transversely (as in Fig. 23 though not as distinctly) folded.
Thorax. Prothorax with lateral margins acute, usually -s nearly complete, occasionally complete, margination more or
less mediolateral, not equiplanar; basal margin crenulate, mediobasal lobe crenulate, with produced teeth (as in Fig.
72), notai projection short, protrochantins visible (as in
Fig. 63); mesoscutellum cordiform or crenulate; mesosternal
excavation, keel absent; elytra obsoletely striate, sutural
striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete, metacoxal plates well-developed; tarsi, article IV lobed, without ventral fleshy pads, ungues with basal tooth.
Abdomen. Male sternum V more or less emarginate; median lobe with mediolateral flanges; parameres long, with nonmelanized subapical projections (Fig. 92); ovipositor
(as in Fig. 81} long, coxites divided, articulated with bacula, styli subapical.
MATERIAL EXAMINED: 37 specimens (MNHP, MRAC).
Seven or eight species from the type localities,
Nigeria, Angola, and Zaire. Further type studies are necessary to determine how many are described.
Taxonomic list - Stenafricus
*auberti (Pic), 1925b (Stenactvla) NEW COMBINATION
(MNHP); Gabon
*stranqulata (Pic) ,• 1928a (Stenactvla) NEW 76
COMBINATION (MNHP); French Congo
*nitidissimus (Pic), 1914a (Therius) NEW
COMBINATION (MNHP); west central Africa D a e m o n 7 7
Daemon de Laporte, 1836; 24 [type species, by monotypy:
Ptilodactvla gioas de Laporte, 1836: 23 (= Colobodera
ovatus Klug, 1838: 59)]; Lacordaire, 1857: 278; Kasap
and Crowson, 1975: 442,
Colobodera Klug, 1838: 67 (type species, present designa
tion: Ç. ovatus Klug, 1838: 67); Lacordaire, 1857: 278;
Fairmaire, 1896; 341.
Diagnostic combination. This genus may be recognized by the equiplanar prothoracic margination (as in Fig. 67 with lateral margins continuous with anterior) and visible protrochantins (as in Fig. 63).
Description.
Body size. Length 4.5-11.0 mm, width at humeral angles
1.5-4.0 mm.
Head. Quadrate; antennae very weakly serrate, not sexually dimorphic; epistomal sulcus present; labrum (as in
Fig. 47), not transverse, basal width approx. 2x basal width, anteriorly truncate or slightly convex; mandibles
(Fig. 38) bidentate, with prostheca, molar areas, and dorsal ridge, molar socket exaggerated; maxillae (as in
Fig. 32), lacinia forming spore brush, galea closely associated and integrated with spore brush, possessing similar spines; apical palpomere mostly sclerotized and 78 melanized; labium (as in Pig, 23), ligula simple, often somewhat ventrally, transversely folded.
Thorax, Prothorax with acute lateral margins, complete, equiplanar (as in Fig, 67), and continuous with anterior margin; strong basal crenulation, mediobasal lobe crenulate, with three produced denticles; mesoscutellum cordiform (as in Fig, 71); notai projection long, trochantins visible (as in Pig, 64); mesosternal excavation and keel absent; elytra obsoletely striate, sutural striae not deeply depressed; metasternal transverse sutrue visible near discrimen, laterally obsolete; metacoxal plates present, underdeveloped; tarsi (as in Fig, 12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues simple.
Abdomen, Male visible sternum V modified, emarginate; aedeagus, median lobe without mediolateral flanges, with or without subapical accessory projections; parameres without nonmelanized projections, often with long apical setae; ovipositor (Fig, 81), long, coxites divided, articulated with bacula, with subapicla styli.
Species excluded: subiunctus Pic (Ptilodactvla),
MATERIAL EXAMINED: 302 specimens (MNHP, MRAC, IZWP,
NMNH), Taxonomie list - Daemon 79
aegualis Fairmaire, 1896 (MNHP); Madagascar
*apicalis Pic, 1924b (MNHP); no locality
V . donckieri Pic, 1924b
atopoides Fairmaire, 1896 (MNHP); Madagascar
V . *notaticollis Pic, 1913b (MNHP); Madagascar
*atricornis Pic, 1924b (MNHP); Madagascar
*basalis Fairmaire, 1896 (MNHP); Madagascar
*berqrothi Pic, 1924b (MNHP); Madagascar
cebrionoides Fairmaire, 1896 (MNHP); Madagascar
colossus Fairmaire, 1896 (MNHP); Madagascar
distinquendus Fairmaire, 1896 (MNHP); Madagascar
*diversipes Pic, 1924b (MNHP); Madagascar
*elateroides Fairmaire, 1896 (MNHP); Madagascar
elonqatus (Klug), 1838 (Colobodera); Madagascar
Lacordaire, 1857 (transferred to Daemon)
Fairmaire, 1896 (redescription)
fasciolatus Fairmaire, 1896 (MNHP); Madagascar
*qeniculatus Pic, 1924b (MNHP); Madagascar
qiqas de Laporte, 1836 (MNHP); Madagascar
ovatus (Klug), 1838 (Colobodera)
Lacordaire, 1857 (synonymized with qiqas)
Fairmaire 1896 (redescription)
(Daemoniminus) grandis Pic, 1953b (MNHP); Madagascar
linearis de Laporte, 1836; Madagascar 80 Fairmaire, 1896 {redescription)
lineellus Fairmaire, 1896 (KNHP); Madagascar
*lonQulus Fairmaire, 1896 (MNHP); Madagascar
*minutus Fairmaire, 1896 (MNHP); Madagascar
mucronatus (Klug), 1838 (Colobodera) ; Madagascar
Lacordaire, 1857 (transferred to Daemon)
Fairmaire, 1896 (redescription)
*niqer Pic, 1953b (MNHP); Madagascar
*niariceps Fairmaire, 1896 (MNHP); Madagascar
*niqrithorax Pic, 1913b (MNHP); no locality
nitidus (Klug), 1838 (Colobodera); Madagascar
Lacordaire, 1857 (transferred to Daemon)
Fairmaire, 1896 (redescription)
notaticollis Pic, 1913b; no locality
*pallidipes Pic, 1924b (MNHP);.Madagascar
*quadrinotatus Pic, 1913b (MNHP); no locality
striatus (Klug), 1838 (Colobodera) ; Madagascar
Lacordaire, 1857 (transferred to Daemon)
Fairmaire, 1896 (redescription)
*subelonqatus Pic, 1913b (MNHP); no locality
*trifasciatus Fairmaire, 1896 (MNHP); Madagascar
*truncatulus Fairmaire, 1896 (MNHP); Madagascar
*vittatus Pic, 1949 (MNHP); Madagascar 81 Falsotherius
Falsotherius Pic, 1913a: 165 (type species, by subsequent
designation [Deleve, 1971: 355]): Falsotherius
atricolor Pic, 1913a: 165);
Diagnostic combination. This genus may be recognized
by the following combination of characters: antennae
slightly serrate, epistomal sulcus absent, protrochantins
loosely concealed, tarsomere IV reduced, I-III with
ventral, fleshy pads, II-III lobed, and males with apical
labial (Fig. 18) and maxillary (Fig. 35) palpomeres ovoid
and half-membranous.
Description.
Body size. Length 5.5-7.5 mm, width at humeral angles
1.5-2.5 mm.
Head, Quadrate; antennae slightly serrate, nearly
filiform; epistomal sulcus absent; labrum (Fig. 44) not
transverse, basal width 2x or less greatest length,
anterior margin concave, mandibles tridentate (as in Fig.
37), without prostheca, molar areas, or dorsal ridge;
maxillae (Fig. 18), lacinia blunt, not forming spore brush,
galea blunt with small production on lateral sub-apex;
labium (Fig. 35), ligula biacuminate; labial and maxillary
palpi shortened, in males, apical palpomeres ovoid, 1/2 membranous. 82 Thorax; Prothorax somewhat anteriorly produced, with acute mediolateral margins very short, strong basal crenulation, mediobasal lobe crenulate, with produced denticles (as in Fig. 71); mesoscutellum cordiform; notai projection long, cowling loosely meeting anterior edge, trochantins loosely concealed (as in Fig. 66); mesosternal keel, excavation absent; elytra strongly striate, sutural striae not more deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates underdeveloped; tarsi, article IV reduced
(as in Fig. 12), I-III with ventral, fleshy pads, II-III drawn into lobes; ungues simple.
Abdomen. Male sternum V modified, emarginate; aedeagus without mediolateral spines or flanges, or subapical accessory projections; parameres long, without nonmelanized apical projections; ovipositor (Fig. 82) long, coxites not divided, articulated with bacula, styli absent.
MATERIAL EXAMINED: 62 specimens (BMNH, NMNH, ÜCDC, MNHP,
IZWP).
Taxonomic list - Falsotherius
*apicicornis (Pic), 1934a (Eoilichas) NEW
COMBINATION (MNHP); Selangor (Malaysia)
*atricolor Pic 1913a (MNHP); Sumatra
*dohertvi Pic 1913a (MNHP); Malacca (Malaysia) Pseudoepilichas 83
Pseudoepilichas Armstrong and Nakane, 1956: 212 (type
species, by monotypy: Epilichas niponicus Lewis 1895: 101).
Diagnostic combination. This genus may be recognized by the following combinatin of characters: male antennae with nonarticulated rami (as in Fig. 5), epistomal sulcus present, protrochantins visible (as in Fig. 65), tarsomere
IV reduced (as in Fig. 12), II-III ventrally lobed, without fleshy pads, and apical palpomeres simple, largely sclerotized.
Description.
Body size. Length 5.5-6.0 mm, width at humeral angles
1.5-2.0 mm.
Head. Quadrate; antennae, males with nonarticulated rami, antennomeres III-X (as in Fig. 5), females serrate; epistomal sulcus present; labrum, not transverse, basal width approx. 2x greatest length, anterior margin truncate or slightly convex (as in Fig. 47); mandibles bidentate, with prostheca, dorsal ridge (as in Fig. 41), without molar areas; maxillae, lacinia blunt, not forming spore brush, galea biacuminate (as in Fig. 28); labium, ligula quadracuminate (as in Fig. 21); apical palpomeres simple, mostly sclerotized. Q ^ Thorax. Prothorax with acute lateral margination, incomplete, anterior end of margin not curving ventrally; basal margin weakly crenulate, mediobasal lobe smooth, with three produced denticles; notai projection long, trochantin visible (as in Pig. 64); mesoscutellum basally smooth (as in Fig. 70); mesosternal excavation, keel, absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi, article IV reduced (as in Fig. 12), II-III lobed, without ventral fleshy pads; ungues simple.
Abdomen. Male sternum V simple; aedeagus, median lobe with mediolateral flanges, without subapical accessory processes; parameres long, without nonmelanized apical projections; ovipositor (as in Fig, 80) long, coxites not divided, articulated with bacula, styli absent.
MATERIAL EXAMINED; 8 specimens (KUBC, BMNH).
Taxonomic - Pseudoepilichas
*niponicus (Lewis), 1895 (Epilichas) (BMNH); Japan
robustior Nakane, 1963; Japan 85 EctvPhodactvla NEW GENUS
Ectvphodactvla NEW GENUS (type species, by monotypy: E.
fiski NEW SPECIES).
Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margination obsolete, protrochantins loosely concealed (Fig. 66), tarsomere IV reduced. III ventrally lobed, simple ungues (as in Fig. 12), head somewhat elongate, clypeus mostly covering labrum (Fig. 53), and males with apical maxillary and labial palpomeres elongate and mostly membranous (Figs. 17, 36).
Description.
Body size. Length 7.5-12.0 mm, width at humeral angles
3-4 mm.
Head. Rather elongate; antennae serrate; epistomal sulcus absent, labrum (Fig. 53) long, basal width 2x or less greatest length, anteriorly convex; mandibles tridentate, with prostheca and dorsal ridge (as in Fig.
37), without molar areas (as in Fig. 39); maxillae (Fig.
36), lacinia blunt, not forming spore brush, galea blunt; in males, apical maxillary and labial palpomeres elongate and largely membranous, membranous areas covered with short, evenly-spaced sensilla (Figs. 17, 36), in females, elongate and mostly sclerotized; labium (Fig. 17), ligula 86 biacuminate, each half tapered to blunt apex.
Thorax. Pronotum (Figs. 53, 66, 71) strongly dorsoanteriorly rounded, with obsolete lateral margination
(Fig. 66), strong basal crenulation, mediobasal lobe crenulate, with produced denticles (Fig. 71); notai projection long, protrochantins concealed, cowlings loosely meeting anterior edge of notai projection (Pig. 66); mesoscutellum cordiform (Fig. 71); mesosternal excavation, keel absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; coxal plates well- developed; tarsi (as in Fig. 12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues simple.
Abdomen, Male sternum V emarginate; aedeagus (Fig.
89), median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized accessory projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula, styli absent.
The following species description does not include characters discussed in the generic description.
Ectvphodactvla fiski NEW SPECIES
Type material. Holotype male, label data: COSTA RICA: 87 Heredia, Prov,, Pto. Viejo, Finca la Silva, II-1-1974,
FWFisk (OSÜC). Paratype males, 8 specimens: Mexico, V. C.
Volcan San Martin SE. slope, B & B Valentine/ beating at camp, el. 4000', 21-25-VII-59 (OSDC); Coyame, Lake
Catemaco, Veracruz, MEXICO, 10-18.VII.63-Black, Its. D. R.
Whitehead (NMNH); COSTA RICA, Turrialba, VII-15-19-65, P.
J. Spangler (2 spec., NMNH); COSTA RICA, Turrialba,
Cartage, III-11-67, W.R. Enns, coll./U Mo, C (EEMC); Costa
Rica: Heredia, pr. LaSelva nr. Puerto Viejo 50m 17-
20.11,80, RS Anderson (CNCI); COSTA RICA Cart. Pr., 20 Km.
E Turrialba, June 9, 11 1980, J.E. Wappes (OSUC); Honduras,
1940/ W. von Hagen (AMNH). Paratype females, 4 specimens:
MEX, Yerba Buena, 20 mi. N. Bochil, Chis. VI-10-1969, D.E.
Bright (CNCI); MEX, Veracruz, El Bastanal, nr., Coyame, ca.
2500', San Andres Mts., IX,19.65/ George E. Ball, D. R.
Whitehead, collectors (NMNH); Mexico, V. C. Volcan San
Martin SE. slope, B & B Valentine/ beating el,, 4-5000',
23-VII-59 (OSÜC); COSTA RICA: Puntarenas, Monteverde area,
6 June-14 June 1973, 1400-1700 meters/ Erwin & Hevel,
Central American Expedition, 1973 (NMNH).
Description. Antennae very weakly serrate; clypeus lengthened, nearly covering labrum (Fig. 53); dorsal mandibular ridge prominent; anterolateral angles of ligula more produced than inner (Fig. 17); prominent groove dorsomesally to each eye; pronotum and elytra minutely 88 punctate, sparsely covered with fine setae; aedeagus (Fig.
89), median lobe very thin, parameres dorsoventrally compressed, broad, blade-like, apex and subapex covered with short setae.
Larvae unknown.
Etymology. Ectvphodactvla is from a combination of the greek words "ektyphos" (puffed up, or swollen) and
"dactylos" (finger). "Ektyphos" is in reference to the bulbous pronotum; "dactylos" is taken from the family name,
Ptilodactylidae; and the specific epithet "fiski" is in honor of Dr. Frank W. Fisk, Emeritus Professor, Department of Entomology, The Ohio State University, collector of the holotype specimen.
MATERIAL EXAMINED: 13 specimens (NMNH, OSUC, CNCI, AMNH).
Specimens are from Mexico (Chiapas, Veracruz), Costa
Rica, and Honduras.
Taxonomic list - Ectvphodactvla
*fiski NEW SPECIES (OSUC); Mexico, Central America Ptiloderes NEW GENUS
Ptiloderes NEW GENUS (type species, by monotypy: P.
humeralifer Pic, 1928c: 8).
Diagnostic combination. This genus may be recognized by the following combination of characters: protrochantins concealed, lateral pronotal margination incompletely acute
(as in Fig. 68), posterolateral pronotal angles broadly explanate (Pigs. 3, 52, 73-76), tarsi (as in Fig. 12), tarsomere IV reduced. III ventrally lobed, mesoscutellum cordiform (Figs. 73-76), and lacinia not forming spore brush (Fig. 34).
Body size. Length 3.5-7.5 mm, width at humeral angles
1.5-3.5 mm.
Head. Quadrate; male antennae with articulated rami
(as in Fig. 7), female serrate; epistomal sulcus present, labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly concave; mandibles bidentate, with prostheca and dorsal ridge (as in
Fig. 41), without molar areas (as in Fig. 39); maxillae
(Fig. 34), lacinia blunt, not forming spore brush, galea acute, apical palpomere mostly sclerotized; labium (Fig.
15), ligula with each half tapered to blunt apex.
Thorax. Pronotum with incompletely acute lateral margins (Fig. 52, as in 58), hind angles broadly explanate 90 (Fig. 3, 52, 73-76), basal margin weakly crenulate,
mediobasal lobe smooth, with produced denticles (Figs. 73-
76); notai projection long, protrochantins concealed (as in
Fig. 68); mesoscutellum cordiform (Fig. 73); mesosternal
excavation, keel absent; elytra obsoletely striate, sutural
striae deeply impressed; metasternal transverse suture
visible near discrimen, laterally obsolete; coxal plates
well-developed; tarsi (as in Fig. 12), article IV reduced,
III lobed, without ventral fleshy pads, ungues bifid.
Abdomen. Male sternum V emarginate; aedeagus, median
lobe without mediolateral flanges or subapical accessory
projections, parameres long, witout nonmelanized apical
projections; ovipositor (as in Fig. 80) long, coxites not
divided, articulated with bacula, styli absent.
Etymology. Ptiloderes is a combination of "ptilon"
(Gr., feather, wing, leaf) and "dere" (Gr., fneck,
throat) and is in reference to the broadly explanate
posterolateral angles of the pronotum.
MATERIAL EXAMINED: 45 specimens (IZWP, INPA, CNCI, EGRC,
BMNH, LACM, MZSP, MNHB, NMNH, MNHP, CASC).
There are 6 or 7 species in this genus from Panama,
Ecuador, and Brazil. Further type studies are necessary to
determine which species are described.
Taxonomic list - Ptiloderes
*humeralifer (Pic), 1928c (Ptilodactvla) NEW
COMBINATION (MNHP); Brazil 91 Pherocladus
Pherocladus•Fairmaire 1881: 372 (type species, by monotypy:
P. dermestoides Fairmaire, 1881: 372); Deleve, 1972:
264.
Diagnostic combination. This genus may be recognized
by the following combination of characters: lateral
prothoracic margination incompletely acute, protrochantins
concealed and extreme anterior pronotal production (Fig.
61).
Description.
Body size. Length 3.0-6.0 mm, width at humeral angles
1.0-2.0 mm.
Head, Quadrate; male antennae (as in Figs. 2, 7) with articulated rami, antennomeres IV-X, females serrate (as in
Fig. 4); epistomal sulcus present; labrum (as in Fig. 47) not transverse, basal width 2x or more greatest length; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig. 31), lacinia forming spore brush, galea acute, closely associated with
spore brush and with longer, finer setae; apical maxillary palpomere mostly sclerotized; labium (as in Fig. 23),
ligula simple, ventrally, transversely folded.
Thorax. Prothoracic margins incompletely acute,
extreme anterior pronotal production, dorsal to anterior 92 margin (Fig. 61); basal margin strongly crenulate, mediobasal lobe crenulate, with three produced denticles
(as in Fig. 71); mesoscutellum cordiform; notai projection long, protrochantins concealed (Fig. 61); mesosternal keel, excavation absent; elytra obsoletely striate, sutural striae variable, may or may not be deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi
(as in Fig. 12), article IV reduced. III lobed, without ventral fleshy pads; ungues with basal teeth.
Abdomen. Male sternum V emarginate; aedeagus, median lobe without mediolateral spines or subapical accessory projections, parameres long, without apical nonmelanized projections; ovipositor (as in Pig. 80) long, coxites not divided, articulated with bacula, without styli.
Species excluded; andrewesi Bourgeois (Ptilodactvla)
MATERIAL EXAMINED: 9 specimens (MNHP, NMNH, BMNH, MCZC,
IZWP). 93 Taxonomie list - Pherocladus
atriceps Pic, 1923c; Sumatra
*capucina Pic, 1923c (MNHP); Sumatra
*castanescens Pic, 1923c (MNHP); Borneo
dermestoides Fairmaire, 1881; Insel Viti (Fiji Isl.)
*inlatefalis Pic, 1923c (MNHP); Sumatra
palawanus Deleve, 1972 (ÜZMD); Palawan, Philippines
*plicata Pic, 1923c (MNHP); Borneo
*sinqularicollis Pic, 1935 (BMNH); Borneo
*testaceomaculata Pic, 1923c (MNHP); Sumatra 94 Lachnodactvla
Lachnodactvla Champion, 1897a; 658 (type species, here
designated: L. monticola Champion, 1897a: 659);
Schaeffer, 1906: 115; Hlavac, 1975: 182; Stribling,
1986; 232.
Diagnostic combination. This genus may be recognized by the following combination of characters: protrochantins concealed (as in Fig. 68), lateral pronotal margination incompletely acute (as in Fig. 68), tarsomere IV reduced,
III ventrally lobed (as in Fig. 12), mesoscutellum usually crenulate (as in Fig. 72), males with apical maxillary palpomeres thickened, elongate, and largely membranous
(Fig. 27), and females usually with apical maxillary palpomere slender, elongate, and mostly sclerotized.
Description.
Body size. Length 3.5-9.0 mm, width at humeral angles
1.0-2.5 mm.
Head. Quadrate; males, antennae with articulated rami
(as in Figs. 2 and 7), females serrate (as in Pig. 4, usually more weakly serrate); epistomal sulcus present, labrum (as in Fig. 47) long, basal width 2x or less greatest length; mandibles (Fig. 41) bidentate, with prostheca, molar areas, and acute dorsal ridge; maxillae
(Fig. 27), lacinia forming spore brush, galea acute. 95 closely associated with spore brush, with longer, thinner, and less dense setae; in males, apical palpomere elongate, mesal 1/2 of more membranous, covered with evenly spaced, very short sensilla, in dried specimens often appearing as longitudinal furrow, in females usually somewhat elongate
(less so in L. arizonica Schaeffer), mostly sclerotized.
Thorax, Pronotum (as in Fig, 68) with incompletely acute lateral margins, strong basal crenulation, mediobasal lobe crenulate, with produced denticles; notai projection long, protrochantins concealed; mesoscutellum cordiform or crenulate; mesosternal excavation, keel absent; elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete or absent; coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, without ventral fleshy pads; ungues with basal tooth.
Abdomen, Male sternum V emarginate; aedeagus (Fig, 96) without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (as in Fig, 80) long, coxites not divided, articulated with bacula, styli absent,
MATERIAL EXAMINED: 303 specimens (CNCI, HUIC, FMNH, CISB,
FSCA, BMNH, UCDC, OSUC, AMNH, MCZC, COBC, JBSC, UCRC, RSNB,
NMNH, EGRC, MSUC, TAMT). 96
Specimens are from Arizona, Texas, Florida, Mexico, C.
America, and Colombia; those from Florida and Colombia
represent 2 undescribed species.
Taxonomic list - Lachnodactvla
*arizonica Schaeffer, 1906 (NMNH); southern Arizona
*monticola Champion, 1897 (BMNH); Mexico, Guatemala
*parviscutum Champion, 1897 (BMNH); Mexico, Guatemala,
Costa Rica
*texana Schaeffer, 1906 (NMNH); Texas, Mexico 97
Ptilodactvla
Ptilodactvla llliger, 1807: 342 (type species, by monotypy:
Ptilodactvla elaterina Illiger, 1807: 342 [= Pvrochroa
nitida DeGeer, 1775: 27]; Latreille, 1929: 461; de
Laporte, 1836: 21, 1840; 258; Blanchard, 1845; 57;
Lacordaire, 1857: 279; LeConte, 1861: 179; Chenu, 1870:
16; Kirsch, 1874: 371; Horn, 1880: 90; LeConte and
Horn, 1883: 171; Champion, 1897: 627; Blatchley, 1910:
690; Legros, 1947: 96; Nakane, 1956: 51; Arnett, 1968:
461; Deleve, 1972: 264; Stribling, 1986: 232.
Hvpselothorax Kirsch, 1866: 188 (I have never seen this
publication, thus I cannot here record the type
species).
Ptvlodactvla de Laporte, 1836: 21 (incorrect subsequent
spelling),
Stenactvla Fairmaire, 1896: 346 (type species, by monotypy:
Stenactvla pallioes Fairmaire, 1896: 346). NEW
SYNONYMY.
Falsodaemon Pic, 1913c: 4 (type species, by monotypy:
Falsodaemon madecassus Pic, 1913c: 4). NEW SYNONYMY.
Theriomorphus Pic, 1913c: 1 (type species, by monotypy:
Theriomorphus niasensis Pic, 1913c: 1). NEW SYNONYMY.
Due to the large number of species included in
Ptilodactvla it is premature to propose hypothetical synapomorphy. Therefore, since the type species of 98
Stenactvla, Falsodaemon, and Theriomorphus cannot be distinguished from Ptilodactvla, the genera are synonymized.
Diagnostic combination. This genus may be recognized by the following combination of characters; lateral pronotal margination incompletely acute, trochantins concealed (Fig. 68), tarsi (as in Fig. 12) tarsomere IV reduced. III lobed ventrally, ungues with basal tooth, mesoscutellum usually cordiform, apical palpomeres mostly sclerotized and securiform, and male antennae with articulated rami (Fig. 2, 7).
Description.
Body size. Length 2.5-11.0 mm, width at humeral angles
< 1.0-3.5 mm.
Head. Quadrate; male antennae with articulated rami
(Figs. 2, 7), female serrate; epistomal sulcus present, labrum long (Fig. 47), basal width 2x or less greatest length, anteriorly truncate or slightly convex; mandibles
(as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig. 31), lacinia forming spore brush, galea acute, closely associated with spore brush, with longer, thinner, and less dense setae, apical palpomere mostly sclerotized; labium (as in Fig. 23), . 99 ligula simple, usually somewhat ventrally, transversely folded.
Thorax. Pronotum with lateral margins incompletely acute (Fig. 68), basal margin strongly crenulate, mediobasal lobe crenulate, with produced denticles (as in
Fig. 71); notai projection long, protrochantins concealed
(Fig. 68); mesoscutellum cordiform, rarely, crenulate; mesosternal excavation absent, keel, rarely, slightly developed; elytra strongly or obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture usually not visible, coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, without ventral fleshy pads, ungues bifid or with basal tooth.
Abdomen, Male sternum V usually emarginate; aedeagus, median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula, styli absent.
SPECIES EXCLUDED: humeralifer Pic (Ptiloderes); all following species are transferred to Epiotvoma: rufa
Champion, rufipennis Pic, corvina Champion, ebenina
Champion, maculate Champion, nigricornis Champion.
More than 20,000 specimens are available for examination from the repositories in the "Material" section. Taxonomie list - Ptilodactvla 100
*acuta Johnson and Freytag, 1978 (NMNH); Georgia
*aequinoctialis Champion, 1897a (BMNH); Panama, Costa Rica
africana Pic, 1942 (MNHP); Cameroon
amamioshimana Nakane, 1963; Japan
amazonica Pic, 1947b; Amazon (Brazil)
*andrewesi (Bourgeois), 1896 (Pherocladus) NEW COMBINATION;
India
anqusta Kirsch, 1874; Peru
*anqustata Horn, 1880 (MCZC); eastern North America
anqustatipennis Pic, 1916b (MNHP); India
Sato, 1979 (diagnosis)
annulata Pic, 1924 (1923c) (MNHP); South America
annulicornis Chevrolat, 1870 (MHCC); Cuba
*antennalis Champion, 1897a (BMNH); Honduras
*antillarum Champion, 1897b (BMNH); Antilles
*apicalis (Pic), 1929 (Stenactvla) NEW COMBINATION (MNHP) ;
Africa
V . *robustior Pic, 1929 (MNHP); Africa
apiceniqra Pic, 1947b; Brazil
apicicornis Pic, 1916a (MNHP); Ceylon
ardua Sato, 1979 (NMBE); India
atra de Laporte, 1836; Cayenne (French Guiana)
atricollis Pic, 1917a; Vietnam
*atricolor Pic, 1947 (MNHP); Peru ICI *atricornis Pic, 1924 (1923c) (MNHP); Malay Archipelago
atrosuturalis Pic, 1916a (MNHP); America meridionalis
*australis Bourgeois, 1884 (RSNB); New Caledonia, New Hebrides
baeri Pic, 1916a (MNHP); Peru
*bakeri Pic, 1923c (MNHP); Borneo, Mindanao
Deleve, 1972 (redescription)
V. *curtithorax Pic, 1923c (MNHP)
V . *sexualis Pic, 1923c (MNHP)
V. *mindanaosa Pic, 1923c (MNHP)
*banatama Deleve, 1972 (üZMD); Lavongai, New Guinea
baroniurbanii Sato, 1979 (NMBE); Nepal
benhana Pic, 1929; Vietnam (Tonkin)
bhutanica Sato, 1979 (NMBE); Bhutan
bicolor Deieve, 1972 (ÜZMD); Palawan, Philippines
bicoloriceps Pic, 1923a (NMNH); Trinidad
boQorensis Pic, 1916a (MNHP); Malacca (Indonesia)
boootensis Pic, 1916a (MNHP); America meridionalis
borneensis Pic, 1916a (MNHP); New Guinea
boucardi Pic, 1923c; Sumatra
brancuccii Sato, 1979 (NMBE); Nepal
ssp. darieelinaiana Sato, 1979 (NMBE); India
bremensis Pic, 1947b; Brazil
breveimpressa Pic, 1923a (MNHP); Brazil
brevenotata Pic, 1947b; no locality
V . innotata Pic, 1947b; no locality 209 brevepubens Pic, 1947b; Colombia
*brevicollis Champion, 1897 (BMNH); Mexico
brevipennis Pic, 1916a (MNHP); Sumatra
*brevipubescens Deleve, 1972 (UZMD); Yalom, New Britian,
New Guinea
*breviramus Pic, 1923c (MNHP); Borneo, Philippines
Deleve, 1972 (redescription)
*breviscutum Champion, 1897a (BMNH); Guatemala
*bruneiensis Pic, 1924 (1923c) (MNHP); Borneo
brunneipennis Pic, 1923c; Borneo, Philippines
*brunneomarqinata Pic, 1924 (1923c) (MNHP); South America
brunneonotata Pic, 1916a (MNHP); Mexico
burqeoni Pic, 1934d (MRAC); Belgian Congo (Zaire)
calcuttana Pic, 1924 (1923c) (MNHP); India
calwelli Pic, 1934b; New Guinea
*canaliculata Champion 1897a (BMNH); Mexico, Guatemala
caracaensis Pic, 1947b; Venezuela
carbonaria Chevrolat, 1870 (MHCC); CUba
*carinata Johnson and Freytag, 1978 (NMNH); Kentucky
castanea de Laporte, 1836 ; Cayenne (French Guiana)
castaneonotata Pic, 1924 (1923c) (MNHP); South America
catharinae Pic, 1947b; Brazil
*caudata Champion, 1897a (BMNH), Central America
celebensis Pic, 1916a (MNHP); Malacca (Indonesia)
chiriquensis Champion, 1897a (BMNH); Panama cisteloides Kirsch, 1874; Peru 103
compressa Pic, 1947c (MNHP); Peru
*confinis Champion, 1897a (BMNH); Central America
confusa Deleve, 1972 (ÜZMD); Yalom, New Britian, New
Guinea
conradti Pic, 1947b; Guatemala
convexa Champion, 1897a (BMNH); Mexico, Nicaragua
convexicollis Champion, 1897a (BMNH); Central America
*corporaali Pic, 1923b (MNHP); Sumatra
costaricana Pic, 1947b (MNHP); Costa Rica
*costaricensis Champion, 1897a (BMNH); Costa Rica
Costata Pic, 1923c (MNHP); South America
crenuatostriata Redtenbacher, 1868; Brazil
*cruciata Kirsch, 1866; Colombia
cucullata Champion, 1897a (BMNH); Central America
curta Pic, 1923c; Sumatra
*curtipennis Pic, 1923c; Borneo; Philippines
curtiramous Pic, 1929; Bolivia
daemoniformis Pic, 1917b (MNHP); Banguey
dascilliformis Pic, 1923c; Sumatra
*debilis Champion, 1897a (BMNH); Mexico, Guatemala
decumana Erichson, 1847; Peru
*deleta Champion, 1897a (BMNH); Central America
delevi (NEW NAME for reducta Deleve, 1972) (UZMD); Tawi
Tawi, Philippines 104 densepunctata Pic, 1923c (MNHP); South America
*dentaticornis Pic, 1924 (1923c) (MNHP); South America
*denticollis Champion, 1897a (BMNH); Costa Rica
*denticulata Champion, 1897a (BMNH); Central America
deplanata Champion, 1897a (BMNH); Panama
deplanaticeps Pic, 1929; Bolivia
dermestoides Pic, 1923c (MNHP); Celebes (Sulawesi)
dilatata Pic, 1929 (MNHP); Bolivia
dilaticollis Champion, 1897a (BMNH); Panama
dilatithorax Pic, 1947b; Brazil
*discedens Deleve, 1972 (UZMD); Lorengau, Manus, N.Guinea
discicollis Pic, 1930a; Bolivia
discoidalis Pic, 1924 (1923c) (MNHP); South America
diversa Pic, 1923c (MNHP); Borneo; Philippines
Deleve 1972 (redescription)
diversecostata Pic, 1935; Borneo
diversepunctata Pic, 1924 (1923c) (MNHP); China
diversipubens Pic, 1924 (1923c) (MNHP); South America
*dohertvi Pic, 1916a (MNHP); Jobi
donckieri (see note)
Note; I do not know if this name is published as a
full species or is merely a manuscript name. I did not
find this type in the Pic collection; he (1952, 1955) published the two varieties under it. 105
V . ruandana Pic, 1955 (MRAC); Ruanda
V . flandriana Pic, 1952 (MRAC); Belgian Congo (Zaire)
elonqata de Laporte, 1836; Colombia
emarqinata Chevrolat, 1870 (MHCC); Cuba
*equilobata Chapin, 1927 (NMNH); United States
*exotica Chapin, 1927 (NMNH); United States
Suss and Puppin, 1977 (redescription)
favrei Pic, 1924 (1923c) (MNHP); Travancore, India
ferruqinea Boheman, 1858; Brazil
*forcioata Champion, 1897a (BMNH); Mexico
forteimpressa Pic, 1947c (MNHP); Brazil
*forticornis Champion, 1897a; Mexico
fruhstorferi Pic, 1947c; Brazil
frvi Pic, 1924 (1923c) (MNHP); South America
furcata Pic, 1954; China
fuscicornis Champion, 1897a (BMNH); Guatemala
qermaini Pic, 1916a (MNHP); Neotropics
*qermana Champion, 1897a (BMNH) ; Panama
*qibbicollis Champion, 1897a (BMNH); Panama
*qlabrata Champion, 1897a (BMNH); Costa Rica
qodavarensis Sato, 1979 (NMBE); Nepal
qorhami Pic, 1924 (1923c) (MNHP); South America
qounelli Pic, 1947b; Brazil
*qracilicornis Deleve, 1972 (UZMD); Yalom, New Britian, New Guinea -0^
*qracilis Champion, 1897a (BMNH); Guatemala
qranulata Pic, 1917b (MNHP); Borneo
*qranulicollis Champion, 1897a (BMNH); Mexico
qriseosuturalis Pic, 1930b; Argentina
V , rufescens Pic, 1930b.
qriseoundulata Pic, 1928d; Brazil
quadalupensis Legros, 1947 (MNHP); Guadeloupe
quatemalensis Champion, 1897a; Guatemala
hektosa (NEW NAME for rufithorax Pic, 1923c); Sumatra
*heterophva Kirsch, 1873; Peru
hickeri Pic, 1947b; Brazil
hirsuta Pic, 1923c; Borneo, Philippines
humeralis Motschulsky, 1863; Ceylon
V , picea Motschulsky, 1863; Ceylon
humeridens Pic, 1947c; Colombia
humerosa Champion, 1897a (BMNH); Mexico, Antilles
hYperqlotta Johnson and Freytag, 1982 (NMNH); Texas,
0. S. A.
impressa Pic, 1924 (1923c) (MNHP); Ganee, Malay
Archipelago
impressipennis Pic, 1916a (MNHP); Neotropics
incisa Pic, 1924 (1923c) (MNHP); South America
inhursuta Pic, 1923c; Sumatra
inlateralis Pic, 1924 (1923c) (MNHP); Sumatra 107 instriata Pic, 1916a (MNHP); Malacca (Indonesia)
*inteara Champion, 1897a (BMNH); Mexico, Guatemala
irregularis Pic, 1928e; Buenos Aires, Argentina
ishiqakiana Sato, 1968; Ryukyu Archipelago
isoloba Johnson and Freytag, 1982 (NMNH); Kentucky, Ü.S.A.
iaponensis (=macrophthalma Nakane)
iiriensis Sato, 1979 (NMBE); Nepal
*iokoensis (Pic), 1931b NEW COMBINATION (Stenactvla) (MNHP);
Cameroon
klapperichi Pic, 1954 (DEIE, MNHP); China
lacorderei de Laporte, 1836 ; Cayenne (French Guiana)
laiovei Pic, 1917b (MNHP); Ho Chi Minh City, Vietnam
lamellifera Kirsch, 1873; Peru
laoensis Pic, 1926a; Laos
lata Pic, 1926b (MNHP); Bolivia
lateralis Pic, 1923c; Sumatra
laticollis Pic, 1923c; Borneo, Philippines
latior Pic, 1923a (MNHP); Trinidad
latithorax Pic, 1924 (1923c) (MNHP); Malay Archipelago
Delève, 1972 (redescription)
leopoldi Pic, 1952 (MRAC); Belgian Congo (Zaire)
lerovi Pic, 1952 (MRAC); Belgian Congo (Zaire)
loiaensis Pic, 1947c; Ecuador
lonqicornis Pic, 1923c (MNHP); Borneo, Philippines
Delève, 1972 (redescription) lonqiramus Pic, 1923c (MNHP); Borneo, Philippines
Delève, 1972 (redescription)
lovasatoi Pic, 1947b; Uruguay
luteipes Pic, 1923c (MNHP); Sumatra
Delève, 1972 (redescription)
luteitarsus Pic, 1947a; Peru
*lutescens Champion, 1897a (BMNH); Mexico; Guatemala
luzonica Pic, 1923c (MNHP); Borneo; Philippines
Delève 1972 (redescription)
macrophtalma Legros, 1947 (MNHP); Guadeloupe
macrophthalma Nakane, 1963; Japan
iaponensis Nakane 1977 UNJUSTIFIED NAME CHANGE
madecassus (Pic), 1913c NEW COMBINATION (Palsodaemon)
(MNHP); Madagascar
madurensis Pic, 1917b (MNHP); India
maindroni Pic, 1924 (1923c) (MNHP); India
malaccana Pic, 1923a (MNHP); Malacca (Idonesia)
malabarensis Pic, 1938b (DEIE); Malabar (SW India)
manuanesis Delève, 1972 (UZMD); Manuan, Duke of York, New
Guinea
*marcida Champion, 1897a (BMNH); Guatemala
*marqinata Champion, 1897a (BMNH); Mexico, Guatemala
martapurana Pic, 1916a (MNHP); Malacca (Indonesia)
maxima Pic, 1923c (MNHP); Bogor (Java)
medanensis Pic, 1917b (MNHP); Sumatra 109 *mexicana Champion, 1897a (BMNH); Mexico
*militaris Chevrolet, 1870 (MHCC); Cuba
minacensis Pic, 1947b; Brazil
minima Pic, 1924 (1923c) (MNHP); New Guinea
minor Pic, 1923c (MNHP); Borneo, Philippines
V. diversicolor Pic, 1923c (MNHP); Borneo, Philippines
Delève, 1972 (redescription)
miniscula Pic, 1923c (MNHP); Malacca (Indonesia)
*minuta Kirsch, 1873; Peru
miscella Sato, 1979 (NMBE); Bhutan
modesta Pic, 1924 (1923c) (MNHP); South America
modiqlianii Pic, 1924 (1923c) (MNHP); Malay Archipelago
*montana Champion, 1897a (BMNH); Panama
mrazai Pic, 1928d (Prague); Brazil
multicostata Pic, 1947c; Brazil
multifaria Sato, 1979 (NMBE); Nepal
multiimoressa Pic, 1947c; Brazil
mussauiana Delève, 1972 (ÜZMD); Talumalaus, Mussau,
New Guinea
mutata Pic; Bolivia (=scutellaris Pic 1928b, not Kirsch)
nanoderma Johnson and Freytag, 1982 (NMNH); Indiana,
U.S.A.
nigra Kirsch, 1873; Peru
nigronotata Pic, 1916a (MNHP); America meridionalis
nitens de Laporte, 1836 ; Cayenne (French Guiana) 110
nitida (Degeer), 1775 (Pvrochroa) ; Cayenne (French Guiana)
elaterina Illiger 1807
Guerin-Meneville, 1829-44 (illustrated
elaterina)
nitidissima Pic, 1927b; Formosa
nitidor Pic, 1916a (MNHP); Sumatra
nodieri Pic, 1947b; Peru
notata Pic, 1929; Bolivia
notaticollis. Pic 1923b; Sumatra
novabritannica Delève, 1972 (UZMD); Yalom, New
Britian, NewGuinea
*novahibernica Delève, 1972 (UZMD); Danu, New
Ireland, NewGuinea
obesa Erichson, 1847; Peru
obiiauefasciata Pic, 1947b; Bolivia
*obovata Champion, 1897a (BMNH); Mexico
obscura Kirsch, 1873; Peru
obscuriceps Pic, 1923c; Sumatra
V . separata Pic, 1923c; Sumatra
obscuricollis Pic, 1923c; Borneo, Philippines
ocularis Pic, 1916a (MNHP); New Guinea
omospila Kirsch, 1873; Peru
*opacicollis Pic, 1924 (1923c) (MNHP); South America
opacithorax Pic, 1923c (MNHP); Colombia
*opima Champion, 1897a (BMNH); Guatemala, Panama *ornata de Laporte, 1836; Brazil 111
*ovalis Delève, 1972 (UZMD); Palawan, Philippines
pallescens Kirsch, 1873; Peru
pallida Steinheil, 1874; Argentina
pallidicolor Pic, 1923c; Sumatra
pallidior Pic, 1929; Bolivia
pallidipes Pic, 1916a (MNHP); Vietnam (Tonkin)
pallidomarqinata Pic, 1924 (1923c) (MNHP); Java
*pallipes (Pairmaire), 1896 NEW COMBINATION (Stenactvla)
(MNHP); Madagascar
*parallela Champion, 1897a (BMNH); Costa Rica, Panama
paranana Pic, 1928c; Argentina
particularicornis Pic, 1924 (1923c)- (MNHP); Mexico
pauli Pic, 1924 (1923c) (MNHP); Brazil
V . iuneta Pic, 1924 (1923c); South America
V , elonqatior Pic, 1928c; Brazil
pendleburvi Pic, 1934a; Malaysia
penta (NEW NAME for minuta Pic, 1947b); Brazil
peruviana Pic, 1923a (MNHP); Peru
peruviensis Pic, 1947b; Peru
*pici (NEW COMBINATION, NEW NAME for Stenactvla anqustata
Pic, 1929 [MNHP]; Africa)
piniqisana Delève, 1972 (UMZD); Palawan, Philippines
planatipennis Delève, 1972 (UMZD); Yalom, New Britian, New
Guinea 11 ') praecellens Kirsch, 1873; Peru
prescutellaris Pic, 1952 (MRAC); Belgian Congo (Zaire)
prima (NEW NAME for atricollis Pic, 1947b); Colombia
probanda Kirsch, 1873; Peru
*pruinosa Champion, 1897a (BMNH); Mexico
puberula Pic, 1928c; Brazil
pubescens Pic, 1923c; Borneo, Philippines
*punctatissima Champion, 1897a (BMNH); Mexico, Guatemala
punctatostriata Murray, 1868 (BMNH); Old Calabar
punosa Pic, 1947b; Peru
*ramea Lewis, 1895 (BMNH); Japan
ramicornis Chevrolat, 1870 (MHCC); Cuba
rectiramus Pic, 1929; Bolivia
V . notaticollis Pic, 1929; Bolivia
reducta Pic, 1924 (1923c) (MNHP); Argentina
reitteri Pic, 1924 (1923c) (MNHP); South America
robusta Pic, 1917b (MNHP); Sumatra
roonensis Pic, 1924 (1923c) (MNHP); Malay Archipelago
*rotundicollis Champion, 1897a (BMNH); Guatemala
rouveri Pic, 1917b (MNHP), Java
V . notaticollis Pic, 1923b; Sumatra
V , brunneosuturalis Pic, 1938a; Pahang (Malaysia)
rubricollis Pic, 1952 (MRAC); Belgian Congo (Zaire)
rufescens Pic, 1947b; Colombia
ruficolor Pic, 1916a (MNHP); American meridionalis 113 rufipes Pic, 1924 (1923c) (MNHP); South America
rufithorax Pic, 1917a; Bolivia
rufocincta Pic, 1916a; America meridionalis
rufohumeralis Pic, 1924 (1923c) (MNHP); Palawan,
Philippines
rufosuturalis Pic, 1924 (1923c) (MNHP); Bolivia
*rufonotata Pic, 1923a (MNHP); Brazil
rufotestacea Champion, 1897a (BMNH); Guatemala
rufula Pic, 1947b; Bolivia
*ruqulosa Champion, 1897a (BMNH); Mexico
saiqonensis Pic, 1929; Ho Chi Minh City, Vietnam
sallei Pic, 1924 (1923c) (MNHP); Mexico
salti Pic, 1953a; Kilimanjaro (Tanzania)
*sanctisvincentis Champion, 1897b (BMNH); Antilles
sanqirensis Pic, 1924 (1923c) (MNHP); Malay Archipelago
sapitensis Pic, 1924 (1923c) (MNHP); Lombok (Indonesia)
satoi (NEW NAME for P. bicolor Sato, 1979 [NMBE]); Bhutan
*scabrosa Champion, 1923 (BMNH); Seychelles
*scapularis Champion, 1897a (BMNH); Panama
scrutata Kirsch, 1866; Colombia
*scutata Champion, 1897a (BMNH); Panama
scutellaris Kirsch, 1873; Peru
scutellaris Pic ( =mutata Pic)
secedens Kirsch, 1873; Peru
secunda (NEW COMBINATION, NEW NAME for Stenactvla bicolor 114 Pic, 1925b [MNHP]); Madagascar
V . testaceicornis Pic, 1925b (MNHP); Madagascar
V . testaceipennis Pic, 1925b (MNHP); Madagascar
semiobscura Pic, 1923c; Borneo, Philippines
sericea de Laporte, 1836 ; Cayenne (French Guiana)
*serrata Champion, 1897a (BMNH); Costa Rica, Panama
*serricollis (Say), 1823 (Ptilinus) ; ü. S. A.
fusca Melsheimer, 1845
Johnson and Freytag, 1982 (neotype designation)
*sexualis Delève, 1972 (ÜMZD); Palawan, Philippines
simplex Chevrolat, 1870 (MHCC); Cuba
simulans Kirsch, 1873; Peru
sinensis Pic, 1923a; China
V . sauteri Pic, 1927b; Formosa
solidicornis Delève, 1972 (üZMD); Balabac, Philippines
sparsepunctata Pic, 1916a (MNHP); Borneo
strangulate Pic, 1924c; Sumatra
striatella Delève, 1972 (ÜZMD); Palawan, Philippines
striatipennis Pic, 1923c (MNHP); Borneo, Phillipines
Delève, 1972 (redescription)
*strictifrons Delève, 1972 (ÜZMD); Banatam, Lavongai,
NewGuinea
suapeinsis Pic, 1947b; Bolivia
subacuminata Pic, 1916a (MNHP); Java
subcastanea Motschulsky, 1863; Java subconvexa Pic, 1923c; Sumatra
subdepressa Pic, 1924 (1923c) (MNHP); Malacca (Malaysia)
subelonqata Pic, 1924 (1923c) (MNHP); Malacca (Malaysia)
subiunctus Pic, 1953b NEW COMBINATION (Daemon) (MNHP);
Madagascar
*submaculata Champion, 1897a (BMNH); Honduras, Nicaragua
subovata Pic, 1924 (1923c) (MNHP); South America
*subparallela Champion, 1897a (BMNH); Mexico
*substriata Champion, 1897a (BMNH); Costa Rica
substriatipennis Pic, 1928c; Brazil
subvittata Pic, 1924 (1923c) (MNHP); Malay Archipelago
♦sulcata Champion, 1897a (BMNH); Mexico
suturalifer Pic, 1929; Bolivia
suturalis Pic, 1916a (MNHP); America meridionalis
♦tabascoana Champion, 1897a (BMNH); Mexico
takahashii Sato, 1968; Ryukyu Archipelago
♦tanqana (Pic), 1914 NEW COMBINATION (Stenactvla) (MNHP);
no locality
♦tenanarivana (Pic), 1925 NEW COMBINATION (Stenactvla)
(MNHP); Madagascar
tenuipunctata Delève 1972 (ÜZMD); Yalom, New Britian,
New Guinea
♦tenuis Champion, 1897a (BMNH); Guatemala
testaceicollis Pic, 1916a (MNHP); America meridionalis
V . inhumeralis Pic, 1947b; Bolivia testaceicornis Pic, 1924 (1923c) (MNHP); Malay
Archipelago
testaceimembris Pic, 1924 (1923c) (MNHP); Madura (Java)
testaceipes Pic, 1924 (1923c) (MNHP); South America
*testaceohumeralis Pic, 1952 (MRAC); Belgian Congo (Zaire)
testaceonotata Pic, 1924 (1923c) (MNHP); Borneo,
Philippines
Delève 1972 (redescription)
*tetra (NEW COMBINATION, NEW NAME for Stenactvla donkieri
Pic, 1913c [MNHP]); Congo
theresae Pic, 1947a; Brazil
thoracica de Laporte, 1840 ; Cayenne (French Guiana)
thri (NEW NAME for curta Pic, 1925); Borneo
toliana Pic, 1924 (1923c) (MNHP); Celebes (Sulawesi)
tonkinea Pic, 1916a (MNHP); Vietnam (Tonkin)
transversicollis Pic, 1923a (MNHP); Brazil
tricoloricornis Pic, 1923a (MNHP); Peru
triimpressa Pic, 1924 (1923c) (MNHP); South America
trinotata Lacordaire, 1857; Brazil
*tropicalis Champion, 1897a; Central America
truncata Pic, 1928c; Brazil
truncaticollis Pic, 1947a; Peru
tucumana Pic, 1928c; Argentina
turrialbana Pic, 1947b; Costa Rica
ueleensis Pic, 1952 (MRAC); Belgian Congo (Zaire) undulata Pic, 1923c (MNHP); Borneo, Philippines
Deleve 1972 (redescription)
V . obscurior Pic, 1923c; Borneo, Philippines
V . subreqularis Pic, 1923c; Borneo, Philippines
V , proxima Pic, 1923c; Borneo, Philippines
ssp. iavanica Delève, 1972 (UZMD); Java
*varicornis Champion, 1897a (BMNH); Panama
varieqata Kirsch, 1889; Colombia
*venusta Delève 1972 (UZMD); Talumalaus, Mussau,
New Guinea
vicina Pic, 1923b; Sumatra
vilis Kirsch, 1873; Peru
*vittata-Pic, 1916a (MNHP); America meridionalis
V . invittata Pic, 1926b (MNHP); Brazil
vuilleti Pic, 1917a; Vietnam (Tonkin)
waqneri Pic, 1924 (1923c) (MNHP); South America
waterstradti Pic, 1924 (1923c) (MNHP); Malay
Archipelago
wittei Pic, 1950; Belgian Congo (Zaire)
wittmeri Sato, 1979 (NMBE); Bhutan Stirophora 115
Stirophora Champion, 1897a; 660 (type species, by monotypy:
S. sulcipennis Champion, 1897a: 660); Stribling, 1986:
232.
Chaetodactvla Champion, 1897a: 660 (type species, by
monotypy: Ç. Ivciformis Champion, 1897a: 661); Hlavac,
1975: 182; Stribling, 1986: 232. NEW SYNONYMY.
This synonymy is on the basis of joint possession of the apomorphic mesosternal keel and the subapical accessory projections of the median aedeagal lobe (Fig. 94).
Diagnostic combination. This genus may be recognized by the following combination of characters: tarsomere IV reduced. III ventrally lobed (as in Fig. 12), protrochan- tins concealed (as in Fig. 68), epistomal sulcus present, lateral pronotal margins incompletely acute (as in Fig.
68), male antennae with articulated rami (as in Fig. 2, 7), mesosternal keel present, usually very prominent, and median aedeagal lobe with subapical accessory projections
(Fig. 94).
Description.
Body size. Length 5.5-7.0 mm; width at humeral angles
1.5-2.0 mm.
Head. Quadrate; antennae, males with articulated rami
(as in Fig. 2, 7), females serrate (as in Fig. 4); labrum 119 (as in Fig. 47), not transverse, basal width approx. 2x
greatest length, anteriorly truncate or slightly convex;
epistomal sulcus present; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and distinct dorsal
ridge; maxillae (as in Fig. 31), lacinia forming well- developed spore brush, galea closely associated with spore brush, with longer, less dense, and thinner setae, apical palpomere mostly sclerotized and melanized; labium (Fig.
23), ligula simple, usually ventrally, transversely, folded.
Thorax. Prothorax with acute, incomplete lateral margins, anteriorly curved toward venter (as in Fig. 68); weak basal crenulation, mediobasal lobe smooth, with three produced denticles; mesoscutellum cordiform; notai projection long, met anteriorly by cowling, protrochantin concealed (as in Fig. 68); mesosternal excavation absent, keel usually present; elytra strongly striate, striae sometimes obscured by setation, sutural stria not more evident; metasternal transverse suture present near discrimen, laterally obsolete; coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, ungues with basal tooth (as in Fig. 11 with mesal furcation truncate).
Abdomen. Male visible sternum V modified, emarginate; aedeagus (Fig. 94), median lobe with subapical accessory 120
projections, without mediolateral flanges, parameres long,
without nonmelanized apical projections; ovipositor (Fig.
80), coxites not divided, without styli, articulated with
bacula.
MATERIAL EXAMINED: 104 specimens (CNCI, HAHC, NDSÜ,
EEMC, EGRC, PMNH, NMNH, BMNH, ÜICM, AMNH).
Taxonomic list - Stirophora
*lvciformis (Champion), 1897a (Chaetodactyla) NEW COMBINATION
(BMNH); Panama, Costa Rica, Nicaragua, Colombia
*sulcipennis Champion, 1897a (BMNH); Panama
There are four species; the two undescribed are from
Costa Rica. 121 Chelonar iomorphus
Chelonariomorphus Pic, 1916a; 2 (type species: Ç.
subconvexus Pic, 1916a: 2); Stribling, 1986: 232.
Diagnostic combination. This genus may be recognized by the following combination of characters: prothoracic margins equiplanar and protrochantins concealed (Fig. 67).
Description.
Body size. Length 4.5-5.5 mm, width at humeral angles
1.5-2.5 mm.
Head. Quadrate; antennae (as in Figs. 2, 7), males with articulated rami on articles IV-X, females serrate; epistomal sulcus present; labrum (as in Fig. 47) not transverse, basal width approx, 2x greatest length, anterior margin truncate or slightly concave; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig, 31), lacinia forming spore brush, galea acute, closely associated with spore brush and with longer, finer setae; apical palpomere mostly sclerotized; ligula (as in Fig. 23) simple, ventrally, transversely folded.
Thorax, Prothoracic margins equiplanar, lateral margins continuous, or very nearly so, with anterior (Fig.
67); basal margin smooth, mediobasal lobe smooth, without produced denticles (Fig. 70); mesoscutellum basally smooth 122 (Fig. 70); notai projection long, trochantins concealed
(Fig.-67); mesosternal keel present, excavation shallow; elytra moderately or obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture absent; metacoxal plates well-developed; tarsi, artilce IV reduced,
III ventrally lobed (as in Fig. 12), without fleshy pads; ungues with basal teeth.
Abdomen. Male sternum V emarginate; aedeagus with mediolateral flanges, without accessory projections, parameres long, without apical nonmelanized projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula; styli absent.
MATERIAL EXAMINED; 18 specimens (BMNH, MNHP, INPA, LACM,
CNCI, FMNH, NMNH, MZSP).
Taxonomic list - Chelonariomorphus
*subconvexus Pic, 1916 (MNHP); Bolivia
I have examined three species (two undescribed) with specimens from Suriname, British Guiana, Peru, and Ecuador. 123 Epjptyqma NEW GENUS
Epjptygma NEW GENUS (type species, present designation:
Ptilodactyla rufa Champion, 1897a: 630).
Diagnostic combination. This genus may be recognized by the following combination of characters: protrochantins concealed, lateral pronotal margination incompletely acute
(as in Fig. 68), tarsi (as in Fig. 12), tarsomere IV reduced. III yentrally lobed, mesosternal keel present, mesoscutellum cordiform with median notch usually continuing as striole to or nearly to apex, and lacinia forming spore brush (as in Fig. 31), and male visible abdominal tergum VI with yentrally-directed flange (Figs.
55-60).
Description.
Body size. Length 5.0-7.0 mm, width at humeral angles
1.5-3.5 mm.
Head. Quadrate; male antennae with articulated rami
(as in Fig. 7), female serrate (as in Fig. 4); epistomal suture present, labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly conyex; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in
Fig. 31), lacinia forming spore brush, with longer, thinner, and less dense setae; apical palpomere mostly 124 sclerotized; labium (as in Pig. 23), ligula simple, often
somewhat ventrally, transversely folded.
Thorax, Pronotum with incompletely acute lateral
margins (as in Fig, 68), strong basal crenulations,
occasionally smooth on either side of mediobasal lobe,
mediobasal lobe crenulate, with produced denticles (as in
Fig 71); notai projection long, protrochantins concealed
(as in Fig, 68); mesoscutellum cordiform (as in Fig, 71);
mesosternal excavation absent, keel present; elytra
obsoletely striate, sutural striae usually deeply
impressed; metasternal transverse suture absent; coxal
plates well-developed; tarsi (as in Fig, 12), article IV
reduced. III ventrally lobed, without ventral fleshy pads,
ungues with basal tooth.
Abdomen, Male sternum V emarginate; segment VI usually heavily sclerotized and with ventrally-directed, posterior
tergal flange (Figs, 55-60); aedeagus, median lobe without
mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (as in Fig, 80) long, coxites not divided, articulated with bacula, styli absent.
Etymology, Epiptvcma is greek for "over-fold" and
"flap" and is in reference to the modification of the visible abdominal tergum VI in males of this genus, that of 125 the ventrally-directed flanges (Figs. 55-60).
MATERIAL EXAMINED: 317 specimens (PMNH, MCZC, CASC, EGRC,
KJOC, HAHC, IZWP, AMNH, CNCI, JBSC, MZSP, EEMC, NMNH, OSUC,
NDSÜ, BMNH, MNHP).
There are approximately 20 species from Suriname,
Bolivia, Panama, Brazil, Costa Rica, Mexico, Belize,
Venezuela, Ecuador, Peru, Nicaragua, and Guatemala.
Further type studies are necessary to determine which of these species are described.
Taxonomic list - Epi-ptvoma
*corvina (Champion), 1897a (Ptilodactvla)
NEW COMBINATION (BMNH); Mexico
*ebenina (Champion), 1897a (Ptilodactyla)
NEW COMBINATION (BMNH); Central America
*maculata (Champion), 1897a (Ptilodactyla)
NEW COMBINATION (BMNH); Central America
*niqricornis (Champion), 1897a (Ptilodactyla)
NEW COMBINATION (BMNH); Central America
*rufa (Champion), 1897a (Ptilodactyla)
NEW COMBINATION (BMNH); Central America
*rufjpennis (Pic), 1916a (Ptilodactyla)
NEW COMBINATION (MNHP); Neotropics Therius
Therius Guerin-Meneville, 1849: 1 (type species, here
designated: T. suturalis Guerin-Meneville, 1849: 2);
Lacordaire, 1857: 270; Kasap and Crowson, 1975: 442;
Hlavac, 1975: 182.
Thervus Pic, 1932: 20 (incorrect subsequent spelling).
Diagnostic combination. This genus may be recognized by the following combination of characters: tarsomere IV
lobed, II-III with ventral fleshy pads, lateral pronotal marginations acute, very nearly reaching anterior margins, protrochantins visible (as in Fig. 64), and mandibles tridentate (as in Fig. 37).
Description.
Body size. Length 5.5-6.0 mm, width at humeral angles
1.5-2.0 mm.
Head. Quadrate; female antennae serrate (as in Fig.
4), epistomal sulcus present; labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly convex; mandibles (as in Fig. 37) tridentate, with prostheca, molar areas, and dorsal ridge; maxillae,
lacinia blunt, not forming spore brush;'galea blunt; apical palpomere mostly sclerotized; labium, ligula with each half bluntly apexed (as in Fig. 15).
Thorax. Pronotal margins incompletely acute. 127 margination very nearly meeting anterior; basal margin
crenulate, mediobasal lobe crenulate, with produced
denticles (as in Fig. 72); notai projection short, protrochantins visible (as in Fig. 63); mesosternal excavation deep, keel absent; mesoscutellum cordiform;
elytra strongly striate, sutural striae not deeply
impressed; metasternal transverse suture visible near
discrimen, laterally obsolete; metacoxal plates well- developed; tarsi, article IV not reduced, lobed, with ventral fleshy pads; ungues simple.
Abdomen. Ovipositor (as in Fig. 81) long, coxites divided, articulated with bacula, styli subapical.
Males unknown.
MATERIAL EXAMINED: 10 specimens (MNHP), all in poor condition.
Species excluded: nitidissimus Pic (Stenafricus)
Taxonomic list - Therius
fulvipes Guerin-Meneville, 1849; S. Africa
iaspideus Fairmaire, 1878; China
*luridipennis Guerin-Meneville, 1849; S. Africa
perrieri Pic, 1931a; Madagascar
V . bicolor Pic, 1931a; Madagascar
*suturalis Guerin-Meneville, 1849; S. Africa
*uniformis Pic, 1913a (MNHP); no locality 128
COMBINATION (MNHP); French Congo
*nitidissimus (Pic)/ 1914a (Therius) NEW
COMBINATION (MNHP); west central Africa .29
INCERTAE SEDIS
Falsoptilodactvla Pic 1958
Valoka Deleve 1972
Podabrocephalus Pic 1913
Brounia Sharp 1878
Hovactvla rufescens Pic 1913
Stenactvla basicornis Fairmaire 1901
Stenactvla lutea Pic 1946
Stenactvla ruficeps 1952
Stenactvla striata Pic 1930
EXCLUSIONS
Cladotoma russula Fairmaire 1904a Cerophytidae
Cladotoma vittata Pic 1914 Pseudolichas
Drupeus Lewis 1895 Psephenidae;
Eubriinae
Pseudolichas Fairmaire 1878 Dascillidae
Pseudodactvlus Hampe 1866 Eurypogonidae
Singularodaemon Pic 1953b (?) Dascilloidea LITERATURE CITED
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FIGURES
144 145
Fig. 1. Dorsal habitus. Anchvcteis velutina, male. 146 147
Fig. 2. Dorsal habitus. Ptilodactvla militaris, male. 148 149
Fig. 3. Dorsal habitus. Ptiloderes sp., female. 150 151
Fig. 4. Serrate antenna; Bradvtoma lineata.
Fig. 5. Famous antenna, nonarticulated rami; Paralichas querini.
Fig. 6. Famous antenna, nonarticulated rami, Cladotoma maculicollis.
Fig. 7, Famous antenna, articulated rami; Ptilodactvla sp. 152 153
Fig. 8. Tarsus, lateral view showing simple tarsomeres and large onychium; Eulichas sp.
Fig. 9. Tarsomere IV, dorsal view showing large onychium; Eulichas sp.
Fig. 10. Pectinate ungues; Paralichas querini.
Fig. 11. Bifid ungues; Ptilodactvla sp. The mesal furcations are often more or less truncate.
Fig. 12, Modified tarsus showing reduction of tarsomere IV, ventral lobe of III, simple ungues, and reduced onychium; Cladotoma cucullatus.
Fig. 13, Simple tarsus; Araeopidius monachus.
Fig. 14. Simple tarsus showing minor lobes on tarsomeres II and III, and no reduction of IV; Pseudocladotoma maculata. 154
i ' / 155
Fig. 15. Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate with each half tapered to blunt apex; Ptiloderes sp.
Fig. 16. Labium, ventral view, palp and setae omitted from right half; hexacuminate ligula; Bradvtoma aurita.
Fig. 17. Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate with each half tapered to blunt apex, apical palpomere with highlighted area of membranous region showing portion of setal clothing; Ectvphodactvla fiski.
Fig. 18. Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate; Falsotherius atricolor. 156 157
Fig, 19. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate with lateral apices reduced; Anchvcteis velutina.
Fig. 20. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Bvrrocrvptus sp.
Fig. 21. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Epilichas candezei.
Fig. 22. Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Cladotoma sp. 00 m 159
Fig. 23. Labium, ventral view, palp and setae omitted from right half; simple ligula folded ventrally; Stirophora Ivciformis.
Fig. 24. Labium, ventral view, palp and setae omitted from right half; modified ligula, multiacuminate; Paralichas trivittis.
Fig. 25. Labium, ventral view, palp and setae omitted from right half; modified ligula, hexacuminate; Octoqlossa sp. .60 161
Fig. 26. Left maxilla, ventral view; multiacuminate galea; Paralichas sp.
Fig. 27. Right maxilla, ventral view, lacinia forming spore brush, apical palpomere elongate and largely membranous; Lachnodactvla parviscutum. 162
L ^ 3 & .62
Fig. 28. Left maxilla, ventral view, biacuminate galea; Epilichas candezei. 164 165
Fig. 29. Left maxilla, ventral view, biacuminate galea with mesal branch absent; Anchvcteis velutina.
Pig. 30. Left maxilla, ventral view, biacuminate galea with mesal branch reduced; Bvrrocrvptus sp.
Fig. 31. Left maxilla, ventral view, lacinia forming spore brush; Aploalossa aureonotata.
Fig. 32. Left maxilla, ventral view, galea and lacinia forming spore brush; Daemon gigas. 166 167
Fig. 33. Left maxilla, ventral view, cylindrical galea with apical cluster of setae; Bradvtoma aurita.
Fig. 34. Left maxilla, ventral view; Ptiloderes sp.
Fig. 35. Right maxilla, ventral view; Falsotherius atricolor.
Fig. 36. Right maxilla, ventral view, apical palpomere with highlighted area of membranous region showing portion of setal clothing; Ectvphodactvla fiski. 168
o 2 'KM 169
Fig. 37. Left mandible, dorsal view; Aploglossa aureonotata.
Fig. 38. Left mandible, dorsal view, showing concave molar area; Daemon gigas.
Fig. 39. Left mandible, dorsal view; Paralichas guerini.
Fig. 40. Right mandible, dorsal view; Octoglossa cvaneipennis. o
OO co 171
Fig. 41. Left mandible, dorsal view; Lachnodactvla parviscutum.
Fig. 42. Left mandible, dorsal view; Anchvcteis velutina.
Fig. 43. Left mandible, dorsal view; Epilichas candezei. 172
#
42 41 173
Pig. 44. Labrum, dorsal view; Falsotherius atricolor.
Fig. 45. Labrum, dorsal view; Aploalossa aureonotata.
Fig. 46. Labrum, dorsal view; Cladotoma sp.
Fig. 47. Labrum, dorsal view; Ptilodactvla sp.
Fig. 48. Labrum, dorsal view; Anchvcteis velutina. u
OO
cjm
4 ^ 175
Fig. 49. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Octoglossa sp.
Fig. 50. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Araeopidius monachus.
Fig, 51. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Cladotoma sp.
Fig. 52. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ptiloderes sp.
Fig. 53. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ectvphodactvla fiski.
Fig. 54. Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ptilodactvla sp. 176
SI 177
Fig. 55. Tergum and sternum of male visible abdominal segment VI, lateral view; Eoiptvama rufa.
Fig. 56. Male visible abdominal segment VI, dorsal view of Figure 55; Eoiotvoma rufa.
Fig. 57. Tergum and sternum of male visible abdominal segment VI, lateral view; Epiptvoma sp.
Fig. 58. Male visible abdominal segment VI, dorsal view of Figure 57; Epiptygma sp.
Fig. 59. Tergum and sternum of male visible abdominal segment VI, lateral view; Epiptygma sp.
Fig. 60. Male visible abdominal segment VI, dorsal view of Figure 59; Epiptygma sp.
Fig. 61. Head and prothorax, lateral view, shaded area = eye; Pherocladus sp.
Fig. 62. Head and prothorax, lateral view, shaded area = eye; Araeopidius monachus. e n en en
en « o
e n en OO en e n
M '-J 03 179
Fig. 63. Head and prothorax, lateral view, shaded area = eye; Octoqlossa sp.
Fig. 64. Head and prothorax, lateral view, shaded area = eye; Paralichas guerini.
Fig. 65. Head and prothorax, lateral view, shaded area = eye; Bradvtoma lineata.
Fig. 6 6 . Head and prothorax, lateral view, shaded area = eye; Ectvphodactvla fiski.
Fig. 67. Head and prothorax, lateral view, shaded area = eye; Chelonariomorphus sp.
Fig. 68. Head and prothorax, lateral view, shaded area = eye; Ptilodactyla sp. o 00 t-1
OO CD CjO c o
1^™ u n CD CD 181
Fig. 69. Anterior dorsum; Araeopidius monachus.
Fig. 70. Anterior dorsum; Chelonariomorphus subconvexus.
Fig. 71, Anterior dorsum; Ectvphodactvla fiski.
Fig. 72. Anterior dorsum; Paralichas querini. 182
71 183
Fig. 73. Anterior dorsum; Ptiloderes sp.
Fig. 74. Pronotum, dorsal view; Ptiloderes sp.
Fig. 75. Pronotum, dorsal view; Ptiloderes sp.
Fig. 76, Pronotum, dorsal view; Ptiloderes sp. 134
74
75
76 18!
Fig, 77, Ovipositor, dorsal view; Aoloalossa sp.
Fig, 78, Ovipositor, dorsal view; Araeopidius monachus 186 187
Fig. 79. Ovipositor, distal portion, dorsal view; Paralichas trivittis.
Fig. 80. Ovipositor, distal portion, dorsal view; Stirophora sulcipennis.
Fig. 81, Ovipositor, distal portion, dorsal view; Daemon sp.
Fig. 82. Ovipositor, distal portion, dorsal view; Falsotherius atricolor. 00 CO r- (
oo oo
o> oo 189
Fig, 83. Ovipositor, dorsal view; Cladotoma sp.
Fig, 84, Ovipositor, right lateral view of Figure 83; Cladotoma sp. VAM
06T 191
Fig. 85. Aedeagus, ventral view; Bradvtoma aurita.
Fig. 8 6 , Aedsagus, right lateral view of Figure 85; Bradvtoma aurita. 192
86 193
Fig. 87. Aedeagus, dorsal view; Aploglossa aureonotata
Fig. 8 8 . Aedeagus, right lateral view of Figure 87; Aploglossa aureonotata. 194
88 195
Fig. 89. Aedeagus, dorsal view; Ectvphodactvla fiski.
Fig. 90. Aedeagus, dorsal view; Chelonariomorphus sp.
Fig. 91. Aedeagus, dorsal view; Paralichas querini.
Fig. 92. Aedeagus, dorsal view; Stenafricus sp. 196
ii 197
Fig. 93, Aedeagus, dorsal view; Cladotoma sp.
Fig. 94. Aedeagus, dorsal view; Stirophora Ivciformis. 198
V.' 199
Fig. 95. Aedeagus/ dorsal view; Epilichas candezei.
Fig. 96. Aedeagus, dorsal view; Lachnodactvla monticola. 200
V.:'
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