Research Article |
Corresponding author: James Liebherr ( jkl5@cornell.edu ) Academic editor: Alexey Solodovnikov
© 2016 James Liebherr.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Liebherr JK (2016) Cyphocoleus Chaudoir (Coleoptera, Carabidae, Odacanthini): descriptive taxonomy, phylogenetic relationships, and the Cenozoic history of New Caledonia. Deutsche Entomologische Zeitschrift 63(2): 211-270. https://doi.org/10.3897/dez.63.10241
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The precinctive New Caledonian genus Cyphocoleus Chaudoir is revised with 22 species recognized, 12 newly described: C. lissus sp. n., C. prolixus sp. n., C. parovicollis sp. n., C. burwelli sp. n., C. angustatus sp. n., C. monteithi sp. n., C. fasciatus sp. n., C. lescheni Liebherr & Will, sp. n., C. cordatus sp. n., C. bourailensis sp. n., C. subulatus sp. n., and C. iledespinsensis sp. n. Atongolium Park & Will is found to be a junior synonym of Cyphocoleus, with its two species recombined as C. mirabilis comb. n. and C. moorei comb. n. Results of a survey of Harpalinae Bonelli place Cyphocoleus as a member of Odacanthini based on synapomorphies of the eighth abdominal tergite and the female spermathecal assembly. Cyphocoleus shares with five other generic-level taxa – Homethes Newman, Aeolodermus Andrewes, Stenocheila Laporte, Quammenis Erwin and Diplacanthogaster Liebke – a single-segmented maxillary galea that is appressed to the outer margin of the maxillary lacinia. These six generic-level taxa are newly classified as members of subtribe Homethinasubtrib. n. (type genus Homethes). Cladistic analysis including 79 taxa and utilizing 119 morphological characters supports division of Odacanthini into four monophyletic subtribes: 1, Actenonycina (Actenonyx White); 2, Homethina; 3, Pentagonicina (Pentagonica Dana, Parascopodes Darlington, Scopodes Erichson); and 4, Odacanthina (24 genera in this analysis monophyletically defined by Lasiocera Dejean and its adelphotaxon). These subtribes are phylogenetically arranged as: (Actenonycina (Homethina (Pentagonicina + Odacanthina). Area relationships defined within Homethina – (New Caledonia (Australia (South America + Central America))) – support the origin of New Caledonian Cyphocoleus prior to amphiantarctic vicariance between South America and Australia. Consistent with previous molecular dating of 100–105 Ma for the origin of Odacanthini, a general vicariance-based hypothesis proposes that New Zealandian Actenonyx and New Caledonian Cyphocoleus were emplaced on Zealandia prior to the completion of rifting between Zealandia and Australia during Late Cretaceous, and that both fragments of Zealandia remained subaerial throughout the Cenozoic. Alternatively, under a very specific time-constrained biogeographic hypothesis ladened with an added assumption of dispersal, the ancestor of Cyphocoleus could have colonized New Caledonia during a 2–5 Ma period after its proposed subaerial reemergence at 37 Ma. A clade within Cyphocoleus synapomorphously exhibits an environmental patina: a varnish-like coating to the dorsal body surface that is hypothesized to enable crypsis of the adult beetle. Several specializations of elytral setae are also synapomorphies of this clade, suggesting evolutionary association of the patina and the setal specializations.
Antarctica, biogeography, morphology, revisionary systematics, taxonomic revision
When Baron Maximilien de
Chaudoir’s first stated yet tentative placement of Cyphocoleus in tribe Platynini was summarily adopted by the only other species describer and reviser of the group (
This revision aims primarily to delineate and describe the species that comprise Cyphocoleus. Taxonomic placement of the genus is based on cladistic analysis using morphological characters of the adult beetles. Comprehensive character analysis of Cyphocoleus shows clearly that these species are members of the tribe Odacanthini. Moreover, within Odacanthini, Cyphocoleus and five other genera form a monophyletic group defined in part by a remarkably salient character: a one-segmented maxillary galea that lies appressed to the outer margin of the maxillary lacinia. Its evolution may be an evolutionary specialization associated with elongation of the mandibles in these beetles, as a similar maxillary configuration – a one-segmented galea that is held free apically from the lacinia – is observed in the long-jawed bembidiine genus Amerizus
Taxonomic material. The taxonomic revision of Cyphocoleus is based on 826 specimens borrowed from 19 institutions (see Acknowledgements). All type specimens were examined for 20 of the 22 species. For two species – Cyphocoleus mirabilis (Park & Will) and C. moorei (Park & Will) – a paratype specimen (
Laboratory techniques. Standard light microscopy, dissection and staining protocols used in this study are described in
Descriptive conventions. All ingroup Cyphocoleus specimens were determined to species through comprehensive examination, including both external characteristics and male genitalia. In order to assess the latter in a comprehensive manner, males of most collecting series, and in some instances numerous males, were dissected. The dissected males were labeled in succession as “male 1, male 2, ...”, with these working labels retained on the specimens along with the genitalia stored in polyethylene genitalia vials. Once specimens had been sorted to species, five specimens (or as many as possible for more rarely collected species) were chosen for mensural analysis. The five specimens included the largest and smallest beetle for the species, plus a representation of males and females. These specimens were labeled as “measured specimen 1 ...” and subjected to intensive examination of external characters. Based on the sample specimens, body size range, and variation among various body ratios were estimated. Also, qualitative characters were assessed among the five to ensure consistency among the specimens sorted to species. All data were entered into multiple Excel® spreadsheet pages, with these data serving as the basis for the dichotomous key, diagnoses, and descriptions.
Several measurements proved useful for diagnosing species. Eye development was quantified by the ocular ratio: MHW/mFW, or maximum head width across eyes divided by the minimum frons width between eyes. Dimensions of the pronotum were routinely used to diagnose several of the species, with four measurements made: 1, MPW, maximum pronotal (or prothoracic) width, measured either across the pronotum, or across the externally bulging proepisterna if visible in dorsal view; 2, APW, apical pronotal width, measured between the two most anterior points along the front of the pronotum; 3, BPW, basal pronotal width, measured across the base of the pronotum along the lateral marginal bead; and 4, PL, pronotal length, or the distance from the apex of the pronotum to the basal margin measured along the midline. These measurements were variously combined into the ratios APW/BPW, MPW/BPW, and MPW/PL, in order to describe prothoracic configuration. The ranges of these ratios are used for descriptive purposes only without any statistical connotations.
Three measurements of the elytra were also used to describe body shape: 1, MEW, maximal elytral width; 2, HuW, humeral width, or the distance between the anterior-most points along the basal elytral margin on both sides of the scutellum; and 3, EL, elytral length, or the distance from the base of the scutellum to the elytral apex, measured parallel to the fused suture, and to the farther of the two apices if they are separated by a median invagination at the suture. These measurements were combined into two ratios, MEW/EL, and MEW/HuW. The former describes the relative elytral breadth, whereas the latter ratio describes the relative width at the front of the elytra, i.e. the humeral development, versus the maximal elytral width near the elytral midlength.
Variation among individuals for counts and relative positions of the lateral elytral setae – i.e. those setae present in the broadened eight elytral interval – is presented using the convention of a(b) + c + d(e). In this sequence, “a” represents the modal number of setae in the anterior series of lateral elytral setae commencing just laterad the humerus, and “(b)” represents the number of setae in the minority of specimens, assuming there is variation observed. The value “c” is used if a seta is isolated between the anterior and posterior series of lateral setae. And “d” represents the modal number of setae in the posterior series of lateral elytral setae situated just anterad the elytral subapical sinuation, with “(e)” representing the number of setae in the minority of specimens for this setal series, assuming variation is observed.
Standardized body length used to describe body size is the sum of three measurements: 1, head length measured from the labral medioapical margin to the cervical ridge at the head-pronotal juncture; 2, pronotal length as measured above; 3, elytral length as measured above. As this sum of measurements ignores the apical portion of the elongate mandibles (always in variable positions specimen to specimen) and any distended portions of the abdomen (also variable dependent on specimen condition when prepared), the standardized body length measure will be smaller than the size perceived by eye.
Ingroup circumscription. The principal intents of this study are to revise the species-level taxa assigned to Cyphocoleus, and to deduce the phylogenetic relationships of those taxa. The latter goal requires robust definition of outgroup taxa to be included in the phylogenetic analysis of Cyphocoleus. This task was complicated by the variety of published proposals regarding taxa found in this study to be related to Cyphocoleus. The female reproductive tract of Cyphocoleus supports inclusion of the genus in Odacanthini, as the spermathecal assembly exhibits a unique configuration (Figs
Based on DNA molecular sequence data,
These starkly different sets of results suggest the need for reevaluation of the previously proposed relationship of Lachnophorini and Odacanthini based principally on spermathecal configuration. Whether to include lachnophorine taxa as outgroups for the phylogenetic analysis of Cyphocoleus formed the proximate question in this endeavor. More broadly, an anatomical survey of taxa placed in Lachnophorini and Odacanthini was undertaken to allow a robust circumscription of taxa to be assigned to Odacanthini. By necessity, such circumscription must precede cladistic analysis of the constituent taxa.
In a seminal morphological analysis of the abdominal terminalia of Carabidae,
Given this distinctive difference in abdominal configuration, abdominal segments VIII and IX and associated male genitalia were dissected from a variety of taxa previously assigned to Odacanthini and Lachnophorini (Figs
Male aedeagus with associated abdominal segments VIII and IX, dorsal view, for taxa assigned to Odacanthini (see Suppl. material
14–18 Male aedeagus with associated abdominal segments VIII and IX, dorsal view for taxa assigned to Odacanthini (see Suppl. material
Abbreviation | Structure |
---|---|
ae | male aedeagal median lobe |
bc | female bursa copulatrix |
cc | collecting canal of defensive gland |
co | female common oviduct |
dgd | defensive gland duct |
dgr | defensive gland reservoir |
ga | female gonocoxal apodeme |
gc | female gonocoxa |
hg | hindgut |
lp | left paramere of male aedeagus |
ltVIII | laterotergite of abdominal segment VIII |
lvVIII | lateroventrite of abdominal segment VIII |
mtIX | mediotergite of abdominal segment IX |
mxg | maxillary galea |
mxl | maxillary lacinia |
mxp | maxillary palp |
mxs | maxillary stipes |
sab | apical bulb of female spermatheca |
sbb | basal bulb of female spermatheca |
sbs | female spermathecal basal sclerite |
sg | female spermathecal gland |
sp | female spermatheca |
sph | spermatophore |
spi | spiracle |
Conversely, males of taxa consistently assigned to Lachnophorini do not exhibit a divided mediotergite VIII (Figs
Pertinent to the goal of this survey, Cyphocoleus males demonstrate the odacanthine abdominal segment VIII condition, with the mediotergite broadly divided into two laterotergites, and the spiracles incorporated into the lateral portion of the tergites (Figs
Right maxillary stipes, lacinia, galea, and palp, Cyphocoleus subulatus: 26, dorsal view; 27, ventral view. Single-segmented galea of Cyphocoleus and other Homethina lies laterad the lacinia, with a rabbeted groove on the dorsomedial surface of the galea accommodating the lateral margin of the lacinia, thereby supporting lacinia during alternating, rotary movements of the maxillae. For abbreviations see Table
28–32 Male aedeagus with associated abdominal segments VIII and IX, dorsal view, Cyphocoleus spp.: 28, C. heterogenus; 29, C. angustatus; 30, C. mirabilis; 31, C. cordatus; 32, C. subulatus. 33. Abdominal lateroventrites VIII (left) and laterotergites VIII (right), showing membranous division of laterotergites, male C. latipennis. Dorsal gland reservoir is globose without dorsal lobe. For abbreviations see Table
Based on these results, a reexamination of the homology of the bipartite spermatheca observed in lachnophorine and odacanthine taxa is in order. Within Odacanthini, the spermathecal assembly is based on a well-sclerotized spermathecal basal sclerite (sbs) which attaches the spermathecal assembly to the common oviduct (Fig.
Female gonocoxal ovipositor and female reproductive tract, ventral view for taxa assigned to Odacanthini (see Suppl. material
The spermatheca of Lachnophorini does not share this configuration because the spermathecal gland duct joins the spermatheca near the base of the basal bulb, not at the apex (
In conclusion, the outgroups considered appropriate to include with Cyphocoleus in a cladistic analysis include those assigned to Odacanthini and Pentagonicini (
Ingroup phylogenetic analysis. The 79 taxa included in the analysis (Suppl. material
Parsimony cladograms comprising exemplar taxa of Odacanthini used in phylogenetic analysis, rooted at the outgroup Celaenephes linearis (tribe Lebiini): 44, 852-step length strict consensus of 29 equally parsimonious trees of 835-step length (CI = 22, RI = 74) [A = subtribe Actenonycina; H = subtribe Homethina subtrib. n.; P = subtribe Pentagonicina; O = subtribe Odacanthina]; 45, majority-rule consensus of 29 equally parsimonious trees with all resolved nodes present in 100% of the multiple equally parsimonious trees; i.e. the majority-rule consensus is the result of collapsing 8 nodes. Abbreviations for geographical distributions include: Af, Africa; As, Asia; Eu, Europe; NC, New Caledonia; Nea, Nearctic; Neo, Neotropical; NZ, New Zealand; Oz, Australia.
Characters. Characters are numbered sequentially starting at the anterior of the head, posterad to the abdomen and then legs for external characters, then to internal anatomical, male genitalic, and female reproductive tract characters. The distribution of character states among taxa can be viewed under the “Diagnoser” toggle of WinClada (
Character 0. Labrum: truncate to slightly emarginate apically (0); broadly convex apically (1).
Character 1. Mandibular length: short, <= to ~1.5× distance antenna base-labral margin (0); moderately elongate, 1.5–1.75× distance (1); elongate, >1.8–2.25× distance (2) (Fig.
Character 2. Maxillary galea comprising: two subequal segments (0); a single elongate segment (1) (Figs
Character 3. Maxillary galea: free of the lacinia (0); appressed to the outer surface of lacinia (1) (Fig.
Character 4. Apical maxillary palpomeres: apparently glabrous (0); covered with short, sparsely distributed setae (1); covered with long, dense setae (2).
Character 5. Antennal scape: moderately long, length 2.0–2.3× breadth (0); elongate, length 2.4–3.2× breadth (1) (Fig.
Character 6. Antennal scape: gracile to moderately broad, anterior surface at most slightly bowed (0); bowed, swollen, anterior surface distinctly convex (1) (Figs
Character 7. Antennomeres 2–3: glabrous except for apical setae (ring setae on 3) (0); with short setae sparsely distributed along shaft (1); with short setae densely distributed along shaft (2).
Character 8. Antennomeres: concolorous or basal antennomeres paler (0); apical 3–4 antennomeres distinctly paler, testaceous (1); antennomeres 7–8 or 8–9 paler (2); antennomeres 7–10 paler (3).
Character 9. Frons: broadly convex (0) (Fig.
Character 10. Frons: smooth or irregularly upraised (0); distinctly punctate (1).
Character 11. Supraorbital setae: two, both anterior and posterior seta present (0); posterior seta present, anterior absent (1); neither anterior nor posterior seta present (2).
Character 12. Eyes: moderate large, greatly projected (0); hypertrophied, convexly projected (1).
Character 13. Eye size: moderate (0) (Fig.
Character 14. Number of ommatidia across horizontal diameter of eye: 10 or more (0); 8 (1).
Character 15. Eye convexity: moderately convex (0) (Fig.
Character 16. Eye convexity: moderately convex (0) (Fig.
Character 17. Neck: without any constriction behind vertex, sides not constricted (0) (Fig.
Character 18. Neck defined by: elongate gena, head stalked (0) (Fig.
Character 19. Submentum: with two setae each side (0); with single seta (inner) each side (1).
Character 20. Ventral surface of gena: glabrous (0); with seta each side (1).
Character 21. Mentum: without median tooth (0); with shallow bidentate tooth (1); with obtuse median tooth, apex rounded (2); with acute median tooth, apex pointed (3).
Character 22. Suture between mentum and gula/submentum straight and complete (0); reduced to an indistinct impression, straight or not (1).
Character 23. Suture between mentum and gula/submentum: complete across width, distinct or not (0); absent medially, mentum and gula fused (1).
Character 24. Pronotum: orbicular (0) (Fig.
Character 25. Pronotal length: subequal or less than width (0) (Fig.
Character 26. Pronotal length: less than to twice pronotal width (0); twice or more than twice pronotal width (1) (Fig.
Character 27. Front angles: rounded (0) (Fig.
Character 28. Pronotal hind angles: flat, not upraised (0) (Fig.
Character 29. Median base: bordered by evenly convex marginal bead (0) (Fig.
Character 30. Pronotal basal margin: straight between pronotal hind angles (0); with median lobe extended posteriorly (1).
Character 31. Pronotal disc: smooth or punctate laterally (0) (Fig.
Character 32. Pronotal disc: smooth or transversely wrinkled overall (0); punctate overall (1).
Character 33. Pronotal disc: without parasagittal impressions (0) (Fig.
Character 34. Prothoracic episternum: not expanded nor visible in dorsal view (0) (Fig.
Character 35. Proepisternum: smooth, impunctate (0); punctured in posterior half (1); distinctly punctured or wrinkled over entire surface (2).
Character 36 (unordered). Pronotal lateral seta: present (0); absent (1); more than one seta present (2).
Character 37 (unordered). Pronotal basal seta: present (0); absent (1); more than one seta present each side (2).
Character 38. Pronotal base: smooth, wrinkled or punctate as anterior portion (0); punctate (1).
Character 39. Pronotal lateral margin: straight to convexly curved (0); angulate at lateral seta (1).
Character 40. Pronotal lateral margin: ridge-like, lateral depression present (0); absent, lateral seta if present not in groove (1).
Character 41. Elytral humerus: sinuously rounded laterad parascutellar striole (0) (Fig.
Character 42. Elytral humerus: broadly extended laterally (0) (Fig.
Character 43. Elytral basal margin: straight between humeri across scutellar base (0) (Fig.
Character 44. Elytral basal groove: of uniform depth from scutellum to humerus (0); with deep pit at base of stria 3 (2).
Character 45. Elytral lateral marginal depression: narrow outside anterior series of lateral elytral setae (0) (Fig.
Character 46. Elytral lateral margin: convex laterad anterior series of lateral elytral setae (0) (Fig.
Character 47. Parascutellar seta: present, doubled or not (0); absent (1).
Character 48. Parascutellar seta: in articulatory socket nearly to coplanar with interval 1 (0); in papillate articulatory socket elevated above interval 1 (1).
Character 49. Parascutellar seta: single each elytron (0); double each elytron (1).
Character 50. Dorsal elytral setae: 3–4 or more (0); 2 (anterior and medial setae present) (1); 1 (medial only) (2); 0 (3).
Character 51. Dorsal elytral setae: 3–4 or less (0); more than 3–4, present along length of interval 3 (1).
Character 52. Dorsal elytral setae: articulatory socket coplanar with elytral disc (0); set in foveate articulatory socket (1).
Character 53. Dorsal elytral setae: not surrounded by differentially colored cuticular spot (0); surrounded by differentially colored cuticular spot (1).
Character 54. Elytral macrosetae: present only on interval 3 (0); also present on interval 5 (1); also present on intervals 5 and 7 (2); also present in intervals 1, 5 and 7 (3); present on all intervals 1–7 (4).
Character 55. Elytra: evenly convex from base to apex (0); with shallow lateral depression at basal 1/3 of length (1); with deep, distinct lateral depression at basal 1/3 of length (2); with deep lateral depressions in basal and apical halves (3).
Character 56. Discal elytral intervals: flat to slightly convex (0); moderately convex (1); very convex (2) (Figs
Character 57. Elytral striae: present, shallow to deep (0); present basally and laterally, absent in apical half (1); absent (2).
Character 58. Elytral intervals 3, 5, and 7: of equal convexity (or nearly equal) to others on disc (0) (Fig.
Character 59. Elytral striae: impunctate to minutely punctate (0) (Fig.
Character 60. Lateral elytral setae: with articulatory sockets not greatly raised (0); with papillate articulatory sockets (1).
Character 61. Elytral apices: meeting at suture, margins evenly curved (0) (Figs
Character 62. Elytral subapical spine at apex of stria 3: absent (0); present (1).
Character 63. Subapical sinuation: nearly obsolete, elytral apices obliquely truncate (0); evident (1); very distinct, angularly concave (2); very distinct, with subapical spine laterally (3).
Character 64. Apical elytral setae: subapical and apical present (0); only subapical present (1).
Character 65. Mesepisternal fovea: absent (0); present (1).
Character 66. Metathoracic flight wings: present, functional with folded apex (1); vestigial (2).
Character 67. Metepisternal length to width ratio: 2.5–4.0 (0); 2.0–2.4 (1); 1.3–1.6 (2); less than or equal to 1.0 (3).
Character 68. Abdominal-metathoracic juncture: loosely articulated, membranous (0); tightly articulated, fused (1).
Character 69. Elytral and abdominal ventrite cuticle: moderately sclerotized, somewhat flexible (0); extremely thick and brittle (1).
Character 70. Surface of frons: apparently glabrous, micropunctures may be present (1); with short, sparse pelage (2); with dense pelage of visible setae (3).
Character 71. Elytral intervals: glabrous except for primary setae (micropunctures possible) (0); with very short pelage of secondary setae (1); with moderately short pelage of secondary setae (2); with long dense pelage distributed over surface (3).
Character 72. Dorsal pelage setae: fine, thin, pointed (0); thick, club-like (1) (Figs
Character 73. Head surface: glabrous except for macrosetae (0); with additional macrosetae posterad supraorbital setae (1).
Character 74. Frons microsculpture: evident, isodiametric to transverse (0); reduced, not traceable except in spots (1); absent (2).
Character 75. Frons: covered with distinct isodiametric sculpticells (0); covered with evident, stretched isodiametric sculpticells (1); covered with dense transverse microsculpture (1).
Character 76. Frons surface: smooth (0); with longitudinal microreticulations (1).
Character 77. Pronotal disc: with regular isodiametric mesh (0); with evident transverse microsculpture (1); with fine transverse lines (2); glossy, without evident microsculpture (3).
Character 78. Pronotal discal surface: without microcuticular patterns (0); with swirling microcuticular ridges (1).
Character 79. Elytral disc microsculpture: isodiametric mesh, may be in transverse rows (0); transverse mesh (1); evident transverse lines (2); not evident, surface glossy (3).
Character 80. Elytral microsculpture sculpticells: moderately raised to not raised (0); extensively raised into granulate microsculpture (1).
Character 81. Elytral disc: without window-like ivory spots (0); with apical window-like spot each elytron (1); with apical and basal window-like spot each elytron (2).
Character 82. Environmental patina: absent (0) (Figs
Character 83. Male abdominal apical setae: 2 (1 on each side) (0); 4 (2(rarely 3) on each side) (1); 8 (4 on each side) (2).
Character 84. Female apical abdominal setae: 4 or more (2 or more each side) (0); 2 (1 each side) (1).
Character 85. Apical visible male abdominal ventrite medially: convex to slightly, broadly concave (0); with distinct, deeper notch-like invagination (1).
Character 86. Metacoxae: trisetose (0); bisetose (1). A trisetose metacoxa has two lateral setae and one closer to body midline. The bisetose condition includes only the two lateral setae.
Character 87. Femora: moderately elongate (0) (Fig.
Character 88. Femora: concolorous from base to apex (0) (Fig.
Character 89. Fourth metatarsomere: truncate apically, without lateral lobes or with lobes short, obtuse (0) (Fig.
Character 90. Dorsal surface of tarsomeres: setose, setae long (0); sparsely covered with short setae (1); glabrous (2).
Character 91. Metatarsomere 5: with 2 rows of 6–8 long ventrolateral setae (0); with 2 rows of 4–5 long ventrolateral setae (1); with 2 rows of 3–(2) long ventrolateral setae (3); with two rows of 3 short ventrolateral setae (4).
Character 92 Abdominal tergite 8: complete, undivided medially by membranous cuticle (0); divided medially by narrowly membranous cuticle (1); divided medially by broadly membranous cuticle (2); divided medially by broad membrane, laterally extended as flaps (3).
Character 93. Male tergite VIII apodemes: narrow, the laterotergite stalked (0); broadly triangular, therefore the laterotergite triangular (1).
Character 94. Abdominal tergite 8 sclerotized margin: not incorporating spiracle (0); with spiracle incorporated (1).
Character 95. Male tergite IX ring sclerite: angulate apically, wishbone-shaped (0); hemicircular apically, horseshoe shaped (1).
Character 96. Male aedeagal median lobe: gracile (0) (Fig.
Character 97. Aedeagal median lobe: short to elongate but broader basally than toward apex (0); elongate, dorsal and ventral surfaces parallel for much of length (1).
Character 98. Aedeagal median lobe apex: gradually narrowed apicad ostium (0); parallel-sided, elongate apicad ostium (1); parallel-sided basally, curved into a hook apically (2).
Character 99. Male aedeagal median lobe apex: evenly attenuated or expanded (0) (Fig.
Character 100. Male aedeagal median lobe apex: of moderate length, attenuate or spoon-shaped (0) (Figs
Character 101. Male aedeagal median lobe apex: attenuated, evenly or not, but tip acuminate (0) (Fig.
Character 102 (unordered). Male aedeagal internal sac: evenly spiculated (0) (Fig.
Character 103. Female abdominal tergite IX: broadly convex (0); narrowly convex, apex tightly rounded (1).
Character 104. Female abdominal tergum X: broad, broadly extended laterally beyond gonocoxites (0); narrow, little extended beyond gonocoxites (1).
Character 105. Female bursa copulatrix: membranous (0) (Figs
Character 106. Female bursa copulatrix: membranous, inner surface smooth (0); sparsely covered with spicules on inner surface (1); densely covered with thin or thick spicules on inner surface (2).
Character 107. Female bursa copulatrix: vase-like, common oviduct short basad spermatheca (0); basally broad, with elongate oviduct basad spermatheca (1); basally broad, rectum-like, with very elongate extension to sp (2).
Character 108. Female spermathecal assembly: without apical bulb basally adjoining basal bulb (0); with apical bulb entering basad basal bulb (1).
Character 109. Spermathecal basal bulb: broadly joined to spermathecal sclerite (0); narrowly joined by a duct to spermathecal sclerite (1).
Character 110. Spermathecal gland reservoir: globose or elongate, not divided apically (0); linear, divided or not apically, ductules on elongate strand (1).
Character 111. Basal gonocoxite 1: with 1–3 apical fringe setae (0); with 5–12 apical fringe setae (1); with 14–16 apical fringe setae (2).
Character 112. Basal gonocoxite 1: not extended medially mesad inner margin of gonocoxite 2 (0); convexly extended medially mesad inner margin of gonocoxite 2 (1).
Character 113. Basal gonocoxite 1 basolateral apodeme: gradually thinned apically (0); very thick throughout length, bar-like (1).
Character 114. Apical gonocoxite 2: with 2 (or 1) lateral ensiform setae (0) (Fig.
Character 115. Lateral ensiform setae: arrayed along lateral margin of apical gonocoxite 2 (0); grouped near apex of apical gonocoxite 2 (1).
Character 116. Apical gonocoxite 2: acuminate apically (0); triangular, rounded apically (2).
Character 117. Gonocoxite 2: much narrower basally than length, falciform (0); narrower basally than width, acuminate to triangular (1).
Character 118. Dorsal ensiform seta of gonocoxite 2: present (0); absent (1).
Character 119 (inactive). Zoogeographic Region: New Caledonia, New Zealand, Australia, Asia, Africa incl. Madagascar, Palaearctic, Neotropical, Nearctic.
Cladistic analysis. The 79 taxon × 119 character matrix was developed in WinClada (
Classification of Odacanthini stat. n. The tribe Odacanthini is hereby proposed to consist of four monophyletic subtribes: 1, Actenonycina; 2, Homethina subtrib. n. (type genus Homethes Newman); 3, Pentagonicina; and 4, Odacanthina (Fig.
Actenonycina. The genus Actenonyx is included within Odacanthini based on the shared-derived abdominal configuration (Fig.
Homethina + Pentagonicina + Odacanthina. The presence of the spermathecal apical bulb and broader gonocoxae (characters 108, 117) are synapomorphies of this clade that excludes Actenonyx. The antennae are nearly uniformly elongate (character 5, states 1 and 2), and the apices of the elytra are generally separately rounded each side of the suture (character 61, state 1), though this condition is reversed in some Cyphocoleus, Aeolodermus emarginatus, and several species within Pentagonicina and Odacanthina.
Homethina subtrib. n. This subtribe is dramatically supported by the one segmented maxillary galea (character 2, state 1) that is appressed to the outer margin of the lacinia (character 3, state 1: Figs
The Homethina divide into two sister clades: Cyphocoleus versus five other genera (Fig.
Pentagonicina + Odacanthina. Cladistic support for monophyly of these two subtribes comes from six characters: 1, neck very constricted (character 17, state 2: reversed in Porocara Baehr of Odacanthini); 2, gena short before constricted neck (character 18, state 1: observed in Pentagonicina and Lasiocera Dejean of Odacanthina); 3, elytral subapical sinuation nearly obsolete (character 63, state 0: a condition observed in most pentagonicines and stem odacanthines from Lasiocera to Stenidia Brullé; 4, frons without visible microsculpture (character 74, state 1: extensively reversed to present in Pentagonica, some Scopodes spp., and several Odacanthini); 5, tarsomeres dorsally glabrous (character 90, state 2: reversed to setose in Pentagonica, some Scopodes, and several Odacanthini); 6, and finally, male aedeagal median lobe dorsal and ventral surfaces parallel near ostium, the ostial opening apical not dorsal (Figs
Pentagonicina. The three genera placed in this subtribe exhibit numerous synapomorphies, with the rank of the taxon being the sole change in this historically well-recognized group. The labrum is broadly convex (character 0, state 1), covering the dorsal surfaces of the short mandibles (character 1, state 0). The apical palpomeres are sparsely setose (character 4, state 1), and the mentum/gular suture is reduced laterally or absent altogether (characters 22 and 23, both state 1). These last characters are reversed in Scopodes tasmanicus and Parascopodes cyaneus. The pronotum bears a basal lobe that extends posterad between the hind angles (character 30, state 1) – again reversed in Parascopodes – but the pronotum is consistently angulate laterally (character 39, state 1), hence providing the basis for the taxon name.
Odacanthini. This very diverse, geographically cosmopolitan subtribe exhibits a broad array of characters and body forms, presumably associated with myrmeco-mimetic behavior while individuals are actively foraging. Focusing on the early-diverging lineages Lasiocera, Eucolliuris Liebke, Andrewesia Csiki and Porocara allows focus on the odacanthine groundplan. Based on this cladistic analysis, the common ancestor of Odacanthini is characterized by: 1, frons punctate (character 10, state 1); 2, submentum with two setae each side (character 19, state 0); 3, mentum with a median tooth (character 21, state 2); 4, surface of pronotal disc distinctly punctate (character 32, state 1); 5, elytral striae punctate (character 59, state 1); 6, frons without microsculpture between punctures, glossy (character 74, state 2); 7, pronotal disc (between punctures) without microsculpture, glossy (character 77, state 3); 8, femora with apex dark (character 88, state 2); 9, male aedeagal median lobe gracile (character 96, state 0: Figs
Atongolium Park & Will 2008: 100 (syn. n.)
Cyphocoleus heterogenus
When
Though quite variable in body proportions and appearance (Figs
1 | Elytral striae impunctate; elytral intervals of subequal convexity, although the 5th and 7th intervals may be indistinctly more convex in beetles of some species; body surface always smooth, without varnish-like exudate that may trap environmental debris (Figs |
2 |
– | Elytral striae punctate, either shallowly punctate with punctures elongate and most evident on discal striae 1–5, or distinctly punctate with deep round punctures in all striae; elytral intervals often carinate, especially the 3rd, 5th or 7th intervals; body surface of all mature individuals covered with varnish-like exudate that may trap environmental debris (Figs |
8 |
2 | Head capsule basally constricted posterad gena, neck constricted (Fig. |
3 |
– | Head capsule broad basally, not constricted behind eyes (Fig. |
1. C. lissus sp. n. |
3 | Pronotum broadest before midlength, pronotal front angles subangulate and slightly protruded (Figs |
4 |
– | Pronotum broadest in basal half, narrowed apically toward very narrow head, pronotal front angles narrowly rounded, not protruded (Fig. |
2. C. prolixus sp. n. |
4 | Pronotal front angles either minutely protruded, subangulate, or not protruded and obtusely rounded (Figs |
5 |
– | Pronotal front angles extremely protruded, acute (Fig. |
3. C. heterogenus Chaudoir |
5 | Pronotum broader (Figs |
6 |
– | Pronotum narrower (Figs |
7 |
6 | Eyes more convex, ocular ratio 1.47–1.55 (Fig. |
4. C. parovicollis sp. n. |
– | Eyes less convex, ocular ratio 1.41–1.44 (Fig. |
5. C. ovicollis Fauvel |
7 | Elytral basal groove nearly straight, meeting lateral margin at obtusely angled humerus (Fig. |
6. C. burwelli sp. n. |
– | Elytral basal groove more distinctly curved laterally to meet lateral marginal bead at anteriorly projected, right-angled humerus (Fig. |
7. C. angustatus sp. n. |
8 | Pronotum tubular, distinctly longer than broad, MPW/PL = 0.59–0.83, front angles acute, protruded, or represented by setose, horn-like, digitiform processes that are longer than their diameter (Figs |
9 |
– | Pronotum more orbicular, MPW/PL = 0.86–1.10, bordered laterally by convex lateral marginal bead that is much more elevated than anterior margin inside front angles; front angles obtuse, not or only slightly protruded (Figs |
13 |
9 | Elytra subellipsoid, basal carina recurved anterad to meet lateral margin at angulate humerus (Fig. |
10 |
– | Elytra broadly ellipsoid, disc convex, appearing inflated, humeri broadly rounded (Fig. |
8. C. montiethi sp. n. |
10 | Pronotal disc smooth each side of median impression; pronotal front and hind angles presented by horn-like, digitiform processes (Figs |
11 |
– | Pronotal disc transversely wrinkled each side of median impression; pronotal front angles acute, projection continuously margined by basal and lateral marginal bead (Fig. |
9. C. miricollis Fauvel |
11 | Eyes very small, little convex, 8 ommatidia along maximal horizontal diameter of eye; body size smaller, standardized body length 4.8–5.4 mm | 12 |
– | Eyes small but moderately convex, 12 ommatidia along maximal horizontal diameter of eye; body size larger, standardized body length 6.5–7.0 mm | 10. C. mirabilis (Park & Will) |
12 | Pronotum without lateral marginal expansion between very elongate anterior and posterior digitiform processes (Fig. |
11. C. moorei (Park & Will) |
– | Pronotum with elevated lateral marginal expansion between short, nub-like anterior pronotal process and elongate posterior digitiform process (Fig. |
12. C. lescheni sp. n. |
13 | Body size smaller, standardized body length 4.8–7.5 mm; pronotum much broader apically than basally, APW/BPW= 1.76–4.0 (Figs |
14 |
– | Body size larger, standardized body length 8.1–8.9 mm; pronotal apical width only slightly broader than basal width, APW/BPW = 1.28–1.40 (Fig. |
13. C. fasciatus sp. n. |
14 | Pronotum continuously bordered laterally and basally by uniformly elevated marginal bead that convexly crosses median base (Figs |
15 |
– | Pronotal lateral marginal bead sinuously adjoining median basal margin which bears a much less elevated basal bead or no bead at all (Figs |
18 |
15 | Pronotal disc distinctly wrinkled transversely, wrinkles extended nearly to lateral carina, lateral longitudinal impressions indistinct, broad and shallow (Figs |
16 |
– | Pronotal disc smoother, transverse wrinkles very shallow to obsolete, median disc delimited by distinct longitudinal impressions that define broadly planar lateral depressions (Figs |
17 |
16 | Elytral margin broadly extended laterally outside anterior series of lateral elytral setae, the basal margin distinctly, sinuously recurved anterad parascutellar seta, humeri broadly protruded anteriorly (Fig. |
14. C. cychroides Chaudoir |
– | Elytral margin only narrowly extended laterally outside anterior series of lateral elytral setae, the setae close to lateral marginal bead; elytral basal margin only slightly recurved, the broadly rounded humeri little protruded anteriorly (Fig. |
15. C. latipennis Fauvel |
17 | Eyes small in diameter but very convex, appearing “popeyed” (Fig. |
16. C. cordatus sp. n. |
– | Eyes small in diameter but little protruded, outer surface little convex (Fig. |
17. C. cardiopterus Chaudoir |
18 | Pronotal and elytral surface with evident microsculpture, the sculpticells isodiametric to transverse resulting in a matte to subiridescent surface; elytra and ventral body surface glabrous except for macrosetae, vertex glabrous to sparsely covered by a few short setae (environmental patina may need to be removed from elytra to assess this character) (Figs |
19 |
– | Cuticle of pronotum and elytra extremely glossy, surface microsculpture not traceable; body surface with well-developed pelage of microsetae, extremely dense and extremely evident on frons, elytra, and proepisterna (Fig. |
18. C. globulicollis Fauvel |
19 | Elytral striae deeply and distinctly punctate, their punctures expanding strial breadth (Figs |
20 |
– | Elytral striae punctate but punctures shallow and not expanding strial breadth (Figs |
21 |
20 | Pronotal median discal convexity delimited by deeply incised parasagittal impressions, the pronotal surface smooth, convex between parasagittal impressions and lateral marginal bead (Fig. |
19. C. flavipes Fauvel |
– | Pronotal median discal convexity bordered laterally by broad, shallow parasagittal impressions, transverse wrinkles of disc extended across impressions to reach lateral margin (Fig. |
20. C. bourailensis sp. n. |
21 | Legs and antennal scape flavous, distinctly contrasted to piceous body (Fig. |
21. C. subulatus sp. n. |
– | Legs and antennal scape fuscous, only slightly paler than piceous body (Fig. |
22. C. iledespinsensis sp. n. |
Male aedeagal median lobe and associated parameres of Cyphocoleus spp., dextral view: 51, C. lissus, Mt. Humboldt; 52, C. prolixus, Mt. Humboldt; 53, C. heterogenus, Yahoué; 54, C. heterogenus, internal sac everted, La Foa; 55, C. parovicollis, Me Maoya; 56, C. parovicollis, internal sac everted, Me Maoya; 57, C. ovicollis, Ningua Res.; 58–59, C. ovicollis, internal sac everted, Mt. Do; 58, laevoventral view; 59, dextrodorsal view. Scale bars, 1.0 mm: horizontal scale for figures with transverse orientation; vertical scale for figures with vertical orientation.
Female right gonocoxae, ventral view, for Cyphocoleus spp. illustrating apical fringe setae of basal gonocoxite 1, and two or three lateral ensiform setae, one dorsal ensiform seta, and two apical nematiform setae on apical gonocoxite 2; 66, C. lissus , Mt. Humboldt; 67, C. prolixus, Mt. Humboldt; 68, C. heterogenus, Yahoué; 69, C. parovicollis, Mandjélia; 70, C. ovicollis, Ningua Res.; 71, C. burwelli, Mt. Humboldt; 72, C. monteithi, Mt. Mou; 73, C. miricollis, Forêt Nord; 74, C. mirabilis, Mt. Koghi; 75, C. lescheni, apical gonocoxite damaged, Aoupinié. Scale bar, 0.10 mm.
Male aedeagal median lobe and associated parameres of Cyphocoleus spp., dextral view: 86, C. burwelli, internal sac everted, Ningua Res.; 87, C. angustatus, dextroventral view, internal sac everted, Mt. Panié; 88, C. monteithi, Parque Prov. Rivière Bleue, Haute Pourina; 89, C. miricollis, Mt. Mou; 90, C. fasciatus, dextral view, sac partially everted, Mt. Panié; 91–98, C. cychroides; 91, internal sac everted, Mandjélia; 92–93, Mandjélia; 94, internal sac everted, Hienghene; 95, Touho; 96, Tionaka Tal; 97, Ateou; 98, Pic d’Amoa. Scale bars, 1.0 mm: horizontal scale for figures with transverse orientation; vertical scale for figures with vertical orientation.
Dorsal habitus photographs of Cyphocoleus spp.: 99, C. miricollis, Forêt Nord; 100, C. mirabilis female, environmental patina intact, Mt. Koghi; 101, C. mirabilis male, environmental patina removed, Mt. Koghi; 102, C. moorei female, environmental patina removed from forebody and right elytron, E. Table Unio; 103, C. lescheni female, environmental patina removed, elytra reassociated with dissected specimen, Aoupinié; 104, C. fasciatus male, teneral specimen lacking environmental patina, Mt. Panié.
Female reproductive tract, ventral view, for Cyphocoleus spp.; 105, C. mirabilis, Mt. Koghi; 106, C. fasciatus, Mt. Panié; 107, C. cychroides, Pic d’Amoa; 108, C. latipennis, Mt. Panié; 109, C. cardiopterus, Mt. Mou; 110, C. globulicollis, Col des Roussettes; 111, C. flavipes, Nouméa; 112, C. subulatus, Forêt de Thi. For abbreviations see Table
Female right gonocoxae, ventral view, for Cyphocoleus spp. illustrating apical fringe setae of basal gonocoxite 1, and two lateral ensiform setae, one dorsal ensiform seta, and two apical nematiform setae on apical gonocoxite 2; 113, C. fasciatus, Mt. Panié; 114, C. cychroides, Pic d’Amoa; 115, C. latipennis Mt. Panié; 116, C. cordatus, Forêt Nord; 117, C. globulicollis, Col des Roussettes; 118, C. flavipes, Nouméa; 119, C. bourailensis, Bourail; 120, C. subulatus, Forêt de Thi. Scale bar, 0.10 mm.
Male aedeagal median lobe and associated parameres of Cyphocoleus spp., dextral view: 126–133, C. latipennis; 126, internal sac everted, Mandjélia; 127–128, Mandjélia; 129, Mt. Panié; 130, Pic d’Amoa; 131–132, Aoupinié; 133, Ouenarou; 134–135, C. cordatus; 134, Mt. Koghi; 135, C. cordatus, internal sac everted, Forêt Nord; 136, C. cardiopterus, Dzumac Rd. jct.
Male aedeagal median lobe and associated parameres of Cyphocoleus spp., dextral view: 141–152, C. globulicollis; 141, Aoupinié; 142, Me Maoya; 143, Col de Roussettes; 144, internal sac partially everted, Gelima; 145, Col d’Amieu; 146, Mt. Do; 147, internal sac everted, Ningua; 148–150, Mt. Dzumac Rd; 151–152, Mt. Koghi; 153, C. flavipes, Yahoué; 154, C. bourailensis, Bourail; 155–158, C. subulatus; 155, La Crouen; 156, Mt. Rembai; 157, Col d’Amieu; 158, Mt. Koghi; 159–160, C. iledespinsensis, Île des Pins; 159, dextral view; 160, ventral view.
Dorsal body surface smooth, glossy, with cyanotic iridescent reflection on elytral disc due to dense transverse microsculpture, antennae and legs contrastedly pale than dark body; head stout, broadest at juncture with prothorax, eyes small in diameter with outer surface extremely convex (Fig.
(n = 5). Head capsule stout, frons broadly convex, the frontal grooves irregularly doubled anteriorly near clypeus; mandible length 2.0–2.1× distance from anterior margin of antennal articulatory socket to anterolateral margin of labrum; antennae filiform, moderately elongate, scape length 2.55× maximal breadth; eyes convex, but ocular ratio low due to very broad frons, MHW/mFW = 1.42–1.48; subgena without fixed macroseta. Pronotum orbicular, hind angles completely untraceable due to evenly convex lateral and basal margin; front angles briefly protruded; notum only slightly wider than long, MPW/PL = 1.03–1.07; median longitudinal impression shallow, finely incised on disc, terminated basally in a well-defined dimple; anterior transverse impression traceable as obsolete impressed line, anterior convexity flat; proepisternum not visible from above; prosternal process convex anteriorly, with deep median groove on ventral surface that continues onto posterior face. Elytral disc moderately convex, moderately broad, MEW/EL = 0.71–0.78; elytral intervals moderately and subequally convex across disc; striae indistinctly punctate to smooth, deep and well defined throughout length and in association with lateral elytral intervals; parascutellar seta present, articulatory socket not upraised; lateral elytral setae arranged as (5)6 + 1 + 7(8), the setal articulatory sockets not upraised above surface; both apical and subapical setae present; subapical sinuation moderately incised, convexly meeting lateral margin; apical margins of fused elytra rounded, the elytral apices slightly separated by curvature at suture. Mesepisternum smooth; metepisternum shorter than broad, dorsal length/diagonal width 0.80. Legs of moderate length, mt1 length/tibial length = 0.26; metacoxae bisetose; dorsal surfaces of tarsomeres with two dorsolateral rows of elongate setae; mt4 length to apex of outer lobe 1.6× median length, 3–4 ventrolateral setae each side. Abdomen with apical margin of apical ventrite broadly and very shallowly excavated to evenly convex; apical ventrite of male with one seta each side, of female with two setae each side. Microsculpture well developed on frons, consisting of isodiametric to slightly transversely stretched sculpticells; pronotum with dense transverse mesh over disc, but with isodiametric mesh medioapically on anterior margin; elytral intervals with dense transverse-line microsculpture; entire body surface with sparse pelage of fine microsetae, microsetae longer on frons and ventral body surface and shorter on pronotal disc and elytra. Coloration of head capsule rufopiceous; antennae pale throughout, brunneotestaceous; pronotum, proepipleuron and prosternum piceous; elytra rufopiceous basally, dark rufous apically; elytral epipleuron rufous; metepisternum piceous; abdomen rufobrunneous, ventrite 6 narrowly rufoflavous apically; femur, tibiae and tarsi flavous.
Male genitalia (n = 1). Male aedeagal median lobe robust, broadly parallel sided from base to apex of ostial opening, dorsoventral breadth at midlength 0.25× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 2.75× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype female (
Paratypes (5 specimens). NEW CALEDONIA: Aoupinié, top camp, rainforest, 850 m el., Berlese sieved litter, 21°11’S, 165°18’E, 23-xi-2001, Monteith (
The Latinized adjectival species epithet lissus is based on the Greek lissos, smooth (
Beetles of this species have been found at elevations from 700–1400 m, at localities ranging from Aoupinié to the north and Mt. Humboldt on the south (Fig.
These remarkable beetles are uniquely diagnosed by the extremely elongate head and prothorax combined with broad, ovoid elytra (Fig.
(n = 3). Head capsule broadest anteriorly, a broad flange extended laterally laterad the antennal articulation and before the eye, the posterior articulatory condyle of mandible defining broadest portion of head; neck impression broad and shallow just before juncture of head and prothorax; eyes small though little convex, 17 ommatidia across horizontal diameter of eye; frons broad relative to eyes, ocular ratio 1.23–1.36; supraorbital setae absent; gena setose, a large macroseta present just laterad gula at position slightly posterad hind margin of eye. Pronotum elongate, parallel sided, MPW/PL = 0.44–0.50, broadest near basal third of length; proepimeron bulging outward, visible in dorsal view just laterad and posterad broadest portion of notum; pronotal front angle rounded, not protruded at all; lateral margin of pronotum defined by marginal bead from front angle to just inside subangulate hind angle, median base margined by low, broadly concave ridge; median longitudinal impression very shallow, finely incised; anterior transverse impression shallow, perpendicular to median impression, defining broad, slightly elevated anterior callosity; prosternal process with medially depressed anterior and ventral surfaces, the posterior surface between procoxae convex. Elytra broadly ovoid, the disc relatively flat and sides steeply sloped; discal elytral intervals broadly convex, the associated striae deep, impunctate; parascutellar seta present, articulatory socket not upraised above surface; elytral humeri narrow, humerus distinctly angulate just mesad base of fifth stria; elytral lateral margin distinctly concave laterad anterior series of lateral setae; lateral elytral setae arranged as 6 + 1 + (7)8; subapical and apical elytral setae present; subapical sinuation broadly, shallowly concave, meeting lateral margin at rounded-obtuse angle; elytral apices acutely rounded, slightly separated from each other laterad fused suture. Mesepisternum impunctate; metepisternum subquadrate, dorsal length 1.08× diagonal width. Legs extremely elongate, femora very thin, mt1 length/tibial length = 0.27; metacoxa bisetose; dorsum of tarsomeres covered with extremely short, sparsely distributed microsetae; mt4 outer lobe 1.56× median length, 3–4 ventrolateral setae each side. Abdomen with apical margin of apical ventrite broadly and very shallowly excavated to evenly convex; apical ventrite of male with one seta each side, of female with two setae each side. Microsculpture of frons very transverse, densely packed lines only loosely connected into a mesh; pronotal and elytral discs covered with dense transverse lines causing silvery and cupreous iridescence; dorsal body surface bearing a sparse distribution of extremely short microsetae; pro- meso- and metasternum with pelage of longer, more densely distributed setae, abdominal ventrites apparently glabrous except for fixed macrosetae. Coloration of head capsule rufopiceous anterad eyes, piceous toward pronotum; antennae rufobrunneous, palps and maxillae slightly paler, brunneous; pronotum, proepipleuron and proepimeron piceous; elytral disc rufopiceous, elytral epipleura rufopiceous, metepisternum piceous; abdominal ventrites rufobrunneous, apical ventrite 6 narrow rufoflavous apically; femora and tibiae rufobrunneous, tarsi rufoflavous.
Male genitalia (n = 1). Male aedeagal median lobe robust, broadest near parameral articulations, slightly narrowed toward distal margin of ostial opening, dorsoventral breadth at midlength ̴0.3× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix elongate, columnar, distance from base of gonocoxites to spermathecal basal sclerite 4.5× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Paratypes (2 specimens). NEW CALEDONIA: Mt. Humboldt, moss forest, 1400 m el., pyrethrum trees & logs, 21°53’S, 166°24’E, 06–07-xi- 2002, Monteith & Burwell (
The elongate prothorax exhibited by beetles of this species (Fig.
This species is known only from Mt. Humboldt at 1400 m el. (Fig.
Beetles as large as C. prolixus, standardized body length 10.8–13.1 mm, but head and prothorax much broader, the head much more developed at the expense of pronotal length (Figs
(n = 5). Head capsule broad, robust, distance from clypeal-labral suture subequal to width across eyes; frons broadly convex, with small chevron-shaped indentation medially between front margins of eyes; eyes moderately large, convex, 28 ommatidia across horizontal diameter; eyes moderately small, convex, ocular ratio 1.35–1.42; only posterior supraorbital seta present; neck distinctly impressed just before juncture with prothorax; gena glabrous, subgenal seta absent. Pronotum distinctly cordate, margin sinuately constricted basally; lateral marginal bead present from acute front angles to just mesad tightly rounded hind angles; median base margin by fine bead; proepisternum slightly bulging, visible in dorsal view along middle half of prothorax; median longitudinal impression shallow, finely incised; anterior transverse impression broad, shallow, obliquely meeting median impression; anterior callosity slightly convex; prosternal process with fine median ridge anteriorly between procoxae, medially depressed on ventral and posterior face. Elytra broadly ovoid, disc moderately convex with side only moderately depressed relative to disc; elytral striae impunctate; parascutellar seta present, articulatory socket not upraised above elytral surface; elytra narrow basally, humeri obtuse-angulate at base of fifth stria; lateral margin straight to slightly concave lateral anterior series of lateral elytral setae; lateral elytral setae arranged as 6 + 1 + 7–8; subapical and apical elytral setae present; subapical sinuation distinctly invaginated, curvature greatest just mesad obtusely rounded juncture of sinuation and lateral margin. Mesepisternum impunctate; metepisternal dorsal length 0.86× diagonal width. Abdomen of males with apical margin of apical ventrite distinctly notched, the invagination smoothly rounded; apical ventrite of male with one seta each side, of female with two setae each side. Legs elongate, gracile, mt1 length/tibial length = 0.26; metacoxae bisetose; tarsomeres with short sparse setae dorsally; mt4 outer lobe 1.7× median length, 4–5 ventrolateral setae each side. Microsculpture of frons and vertex a stretched isodiametric mesh; pronotal disc and discal elytral intervals covered with fine transverse lines; dorsal surface of body apparently glabrous except for standard macrosetae; pro-, meso-, and metasternum and visible abdominal ventrites with sparsely distributed pelage of very short setae. Coloration of dorsum rufopiceous with cyanotic to cupreous iridescence due to microsculpture; antennae dark rufous, paler apically, palps brunneous; basal abdominal ventrites rufopiceous, ventrite 6 rufobrunneous, margin narrowly brunneous; femora rufopiceous, tibiae rufobrunneous, tarsi slightly paler, dark rufous.
Male genitalia (n = 8). Male aedeagal median lobe robust, broadly parallel sided in basal 3/4 of length, then evenly narrowed to projected, narrow tip, dorsoventral breadth at midlength 0.25× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 2). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 2.4× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Found throughout Grande Terre, from Mt. Ignambi and Mt. Panié on the north, to Forêt de Thi on the south (Fig.
This species and its following adelphotaxon are difficult to diagnose practically (Figs
(n = 5). Head capsule moderately broad, gena elongate anterad constricted neck (Fig.
Male genitalia (n = 4). Male aedeagal median lobe robust, broadly parallel sided to near distal margin of ostial opening, dorsoventral breadth at midlength 0.2× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 2). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 1.8× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Paratypes (30 specimens). NEW CALEDONIA: Mandjélia, above Pouébo 600–750 m el, 20°24’S, 164°32’E, 11–13-v-1984, Monteith & Cook (
The adelphotaxon status and great similarity of this species and C. ovicollis (Figs
This species is distributed in the northern half of Grande Terre, allopatric with its adelphotaxon, C. ovicollis, which is distributed to the south (Fig.
In juxtaposition to its cryptic adelphotaxon, this species can be diagnosed by the flatter eyes, ocular ratio 1.41–1.44, and presence of two apical abdominal setae in the males of this species – one seta each side of apical visible abdominal ventrite – versus four apical abdominal setae – two each side – in males of C. parovicollis. The pronotum is generally broader basally among individuals of this species – MPW/BPW = 2.04–2.29 – versus values of 1.87–2.10 in C. ovicollis. A male dissection will allow attribution of a series of specimens, with the male of C. ovicollis exhibiting an internal sac with a robust, mound-shaped tooth on the right side (Figs
(n = 5). (The description of C. parovicollis can serve for this species, with the following exceptions). Head capsule with mandibles elongate, length 2.4× distance from antennal articulatory socket to anterolateral margin of labrum. Elytra more broadly subellipsoid, MEW/EL = 0.71–0.73; three dorsal elytral setae uniformly present; lateral elytral setae arranged as 6 + 1(0) + 7(8). Mesepisternum dorsal length 0.8× diagonal width.
Male genitalia (n = 5). Male aedeagal median lobe robust, broadly parallel sided in basal half, then tapered to narrowly rounded tip, dorsoventral breadth at midlength 0.2× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 1.7× maximum breadth (dissection compressed under cover slip) (as in Fig.
Holotype male (
The known distribution northern portions of Province Sud, from Bourail and Col d’Amieu on the north to Mt. Do and Ningua Reserve to the south (Fig.
This species and the following, C. angustatus, comprise a second sibling species pair, these two characterized by the narrow body: i.e. pronotum and elytra. The pronotum of C. burwelli is more evenly ovate, with MPW/BPW = 1.72–1.83 versus values of 1.84–1.91 for C. angustatus. The pronotum is also relatively narrower overall, with MPW/PL = 0.68–0.72 compared to values of 0.72–0.77 recorded for C. angustatus. The elytra are narrower basally in C. burwelli, with MEW/HuW = 2.78–3.19 versus 2.60–2.79 for C. angustatus, and the humeral angle is obtuse versus very distinctly right angled to slightly acute in C. angustatus (Figs
(n = 4). Head capsule moderately elongate, trapezoidal, with broad flange basad posterior mandibular articulation, neck distinctly constricted (Fig.
Male genitalia (n = 1). Male aedeagal median lobe robust, broadly parallel sided in basal half, then tapered to rounded tip, dorsoventral breadth at midlength 0.3× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 1.4× maximum breadth (dissection compressed under cover slip) (as in Fig.
Holotype female (MNHB deposited in
Paratypes (3 specimens). NEW CALEDONIA: Ningua Reserve, near summit, 1300 m el., 21°45’S, 166°09’E, 13-xi-2001, Burwell & Monteith (
This species is named for Dr. Chris Burwell, Senior Curator of Insects, Queensland Museum, in recognition of the numerous specimens of Cyphocoleus he has collected: many representing previously undescribed species.
The distribution of this species ranges from Mt. Humboldt on the south, to Mt. Canala on the north (Fig.
As the second half of the sibling species pair also including C. burwelli, this species can be diagnosed by the more basally constricted pronotum, MPW/BPW = 1.84–1.91 versus values of 1.72–1.83 for C. burwelli. The pronotum is also relatively broader overall, with MPW/PL = 0.72–0.77 versus values of 0.68–0.72 for C. burwelli. The elytra are broader basally in C. angustatus, with MEW/HuW = 2.60–2.79 versus 2.78–3.19 C. burwelli. The humeral angle is positioned more anteriorly relative to the scutellum, and is very distinctly right angled to slightly acute in this species (Fig.
(n = 3). (The description of C. burwelli can serve for this species, with the following exceptions). Head capsule with mandibles exceedingly elongate; length 4.9× distance from antennal articulatory socket to anterolateral margin of labrum; antennae exceedingly elongate, gracile, scape length 4.9× maximal breadth. Pronotum margined basally with narrow, little upraised bead crossing median base between the distinctly margined, rounded hind angles. Elytra narrowly ovoid, extremely narrow basally; two dorsal elytral setae present at positions of middle and posterior seta; lateral elytral setae arranged as 6 + 1 + 7, or 6 + 8. Legs extremely elongate, gracile; metacoxae trisetose, two setae on lateral surface fore and aft, and a third large seta medial to posterior seta (short setae comprising part of ventral pelage also present); mt4 with length to apex of outer lobe 1.45× median tarsomere length. Coloration of legs paler than piceous thoracic and rufopiceous abdominal ventrites, femora dark rufous with piceous cast, tibiae rufobrunneous, and tarsi paler, brunneous to rufoflavous.
Male genitalia (n = 2). Male aedeagal median lobe robust, broadly parallel sided to near distal margin of ostial opening, dorsoventral breadth at midlength 0.2× distance from tip to base of closed basal bulb (Fig.
Holotype male (
Paratypes (7 specimens). NEW CALEDONIA: Mt. Panié, 730 m el., under stones amongst ants, 20°32’S, 164°44’E , 24-viii-1914, Montague (BMNH, 2), 28-viii-1914, Montague (BMNH, 1), 1300 m el., 20°34’S, 164°46’E, 03-xi-1988, Raven (
The narrow, elongate pronotum and very narrow elytral humeri (Fig.
This species is known only from Mt. Panié, with localities at 730 m elevation in 1914, and from 1300–1350 m elevation for more recently collected material (Fig.
This species (Fig.
(n = 5). Head capsule ellipsoid, genal surfaces behind eyes subparallel, then converging to distinctly constricted neck; frons with broad medial crest between eyes, fine arcuate wrinkles curving outward and posterad from crest, frontal grooves parallel, broadly depressed from position between hind margins of eyes to clypeus, broadest just posterad frontoclypeal suture; eyes small, convex, 20 ommatidia across horizontal diameter, ocular ratio 1.39–1.50; supraorbital setae absent; mandibles moderately elongate, length 2.4× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, scape length 2.6× maximal breadth; gena setose, subgenal seta present just laterad gula about 2/3 distance from hind margin of eye to constricted neck. Pronotum elongate, tubular, MPW/PL = 0.59–0.69, base constricted relative to apex, APW/BPW = 1.28–1.40; median longitudinal impression finely and shallowly incised, causing interruptions in orientation of transverse wrinkles in some instances; anterior transverse impression broad, shallow, extended laterally to behind front angles; proepisternum bulging, extended beyond marginal bead of notum, with distinct, deeply engraved vertical wrinkles; prosternal process distinctly depressed medially anterad procoxae, flat to narrowly depressed medially on ventral surface, narrowly convex on posterior surface. Elytra inflated, based inflexed at scutellum and depressed along suture, intervals 3–5 convexly elevated; striae smooth with very widely spaced micropunctures along length (view cleaned specimen); parascutellar seta present, articulatory socket papillate, elevated above surrounding elytral surface; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7, their articulatory sockets slightly upraised above surrounding cuticle, often appearing clear of cuticular varnish; subapical and apical elytral setae present, articulatory sockets papillate and clear of varnish; subapical sinuation moderately concave, meeting lateral margin in broad curve; elytral apices rounded, moderately separated by invaginated margin at fused suture. Mesepisternum impunctate, metepisternal dorsal length 1.1× diagonal width. Abdomen with apical ventrite broadly, slightly concave medially; males with one seta each side apical margin of apical ventrite, females with two setae each side. Legs gracile, moderately elongate, mt1 length/tibial length = 0.27; metacoxae bisetose; tarsomeres with dorsal surface glabrous; mt4 with length to apex of outer lobe 1.8× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture of head transverse, dense; pronotal and elytral disc with fine transverse lines; pro-, meso-, and metasternum plus abdominal ventrites with sparse pelage of extremely short microsetae, such extremely short setae also visible on head on frons, pronotum, and elytra (though cleaning the surface may remove setae). Coloration of head piceous, mandibles rufobrunneous, palps and antennae rufoflavous; pronotum, elytra and body ventrites and coxae piceous; femora piceous with rufobrunneous base and apex, tibiae rufopiceous dorsally, rufobrunneous on lateral and ventral surfaces, tibiae rufobrunneous.
Male genitalia (n = 6). Male aedeagal median lobe robust, broadly parallel sided from base to apex of ostial opening, dorsoventral breadth at midlength 0.25× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 1.8× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Paratypes (22 specimens). NEW CALEDONIA: Mt. Ouin, 1100 m el., pyrethrum trees & logs, 22°01’S, 166°28’E, 09-xi-2002, Burwell & Monteith (
I take great pleasure in naming this extremely distinctive species (Fig.
This species is known only from the southern portion of Grande Terre: localities range from Mt. Ouin on the north to Parc Provincial Rivière Bleue on the south (Fig.
This species shares the elongate prothorax with transversely wrinkled notum and acutely projected front angles with C. monteithi, but the elytra are very flat, and the elytral striae distinctly punctate, the punctures expanding strial breadth (Fig.
(n = 2). Head capsule ovoid with deeply constricted neck, genae distinctly incurved at back of head; frons with large, shallow chevron-shaped medial impression, frontal grooves broad and subparallel; eyes small, moderately convex, ocular ratio 1.30–1.50; supraorbital setae absent; mandibles moderately elongate, length 2.5× distance from antennal articulatory socket to anterolateral margin of labrum; gena glabrous. Pronotum narrow, tubular, MPW/PL = 0.63–0.67; median longitudinal impression deep, interrupting orientation of deep transverse wrinkles, terminated anteriorly as a deep pit; anterior marginal bead very broad, the deep groove just posterad bead interpreted as anterior transverse impression; proepipleuron and proepisternum bulging outward, visible in dorsal view for much of pronotal length, proepisternum smooth. Elytra broadly ellipsoid, MEW/EL = 0.68–0.69; humeri broadly angulate laterad depressed scutellum and parascutellar interval; parascutellar setae present, articulatory socket papillate, elevated above elytral surface; elytral striae deeply incised, punctate, the punctures separated by about 5× puncture diameter; elytral intervals distinctly convex, the sutural, third, fifth, and seventh interval subcarinate, most evidently so basally; dorsal elytral setae absent; lateral elytral margin straight to slightly concave laterad anterior series of lateral elytral setae; lateral elytral setae arranged as 6 + 6; subapical elytral seta present, apical seta absent; subapical sinuation distinctly concave, sinuation subangulately meeting lateral margin; elytral apices broadly rounded, margin deeply invaginated at fused suture. Mesepisternum impunctate; mesepisternum with dorsal length subequal to diagonal width. Abdomen of males and females with apical ventrite broadly, slightly concave medially; males with one seta each side of apical margin of apical ventrite, females with two setae each side. Legs gracile, moderately long; metacoxae bisetose; tarsomeres relatively broad and short, mt1 length 0.21× tibial length; mt4 with long, broad apical lobes, length to apex of outer lobe 2.67× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture reduced, frons with surface shiny, transversely stretched isodiametric mesh traceable over portions of surface; pronotum glossy with fine transverse lines visible over portions of surface; elytral glossy on disc with transverse lines visible apically; body surface – head, pronotum, elytra, and ventral surfaces – with sparse pelage of very short setae (these setae may be removed in part by cleaning). Coloration of head and pronotal disc rufopiceous; prothoracic ventrites piceous; elytra rufobrunneous; thoracic and abdominal ventrites rufopiceous, apical abdominal ventrite apically brunneous; femora and tibiae rufobrunneous, tarsi brunneous.
Male genitalia (n = 1). Male aedeagal median lobe robust, broadly parallel sided from base to apex of ostial opening, dorsoventral breadth at midlength 0.25× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix broadly columnar, distance from base of gonocoxites to spermathecal basal sclerite 2.7× maximum breadth (dissection compressed under cover slip) (Fig.
Lectotype male (
Known from sites in the southern mountains of Province Sud, near Mt. Ouin, at Mt. Mou, and at Forêt Nord (Fig.
Atongolium mirabile Park & Will, 2008: 100.
This species comprises one-third of a species triplet characterized by anterior and posterior projections on the front and hind angles of the pronotum (Figs
(n = 3). Head capsule elongate, genae subparallel behind eyes and deeply converging at constricted neck; frons with small dimple-like depression medially between eyes, the dimple lined with isodiametric sculpticells; frontal grooves broad and deep, restricted to portion of frons anterad eyes; eyes small and slightly convex, a depressed groove continuous with groove at lateral edge of frons extended from eye dorsum to midpoint of eye hind margin; supraorbital setae absent; mandibles moderately elongate, length 2.0× distance from antennal articulatory socket to anterolateral margin of labrum; antennal scape broadly fusiform, length 3× maximal breadth; gena glabrous. Pronotum convex dorsally, with broadly spatulate processes at front and hind angles, the front processes flatter and broader in longitudinal orientation, the posterior processes more hornlike; lateral marginal carina extended and diminished behind posterior process, pronotal median base depressed but not margined; median longitudinal impression finely incised with surface sloping to meet at deepest portion; anterior transverse impression represented by oblique depressions that define an anterior pronotal collar about as wide as neck of head; proepipleuron and proepisternum bulging outward, visible in dorsal view in basal half of pronotal length; proepisternum smooth; prosternal process deeply depressed anterad procoxae, broadly depressed medially on ventral face, and depressed in an inverted “Y” pattern on posterior face resulting in a dorsomedial bump. Elytra only moderately convex overall, scutellum and sutural stria slightly depressed relative to stria 3; at midlength, sutural and fifth interval distinctly elevated, third interval slightly less so but it is also more elevated than intervals 2, 4, and 6; humeri very narrow, the basal groove present only from base of greatly elevated fifth interval to subangulate lateral margin just outside anterior series of lateral elytral setae; parascutellar and dorsal elytral setae absent; lateral elytral setae arranged 6 + 3 + 3, their articulatory sockets papillate, upraised above elytral surface: subapical elytral seta present, apical seta absent; subapical sinuation distinctly concave, the median portion of concavity upraised thereby elevating elytral margin; elytral apices tightly rounded, protruded, a deep invagination between them adjacent to the fused suture. Mesepisternum impunctate; metepisternal dorsal length subequal to diagonal width. Abdomen with very shallow, broad concavity medially on both male and female apical ventrite. Legs moderate, tarsomeres relatively short, mt1 length/tibial length = 0.21; mt4 length to apex of outer lobe 1.2× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture absent on vertex except in median dimple; surface of pronotal and elytral disc glossy, indistinct transverse lines over portions of elytral surface; pelage of club-shaped setae well developed; setae on head capsule and pronotal moderately expanded, twice as broad apically as basally; setae broad and paddle-like on prothoracic lateral carina and anterior and posterior pronotal processes; elytra with broad paddle-like setae on sutural, third, and fifth intervals, setae narrower apically on intervals 2, 4, and 6; elytral lateral margin with short paddle-like setae, their length about half that of longer setae on odd intervals; ventral body surface and legs covered with dense pelage of short microsetae, the setae on legs more robust than those on ventrites. Coloration of head capsule dark rufous; antennal segments 1–4 rufobrunneous, outer antennomeres flavous with rufous cast; pronotum, elytra, and thoracic sclerites and abdominal ventrites dark rufous with a piceous cast; femora rufobrunneous, tibiae darker, rufopiceous, tarsi brunneous.
Male genitalia. Male aedeagal median lobe robust, broadly parallel sided in basal half, evenly tapered in distal half to subacuminate tip, dorsoventral breadth at midlength 0.22× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 2× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Known from across the southern third of New Caledonia, from Col d’Amieu on the north to Rivière des Pirogues on the south (Fig.
Atongolium moorei Park & Will, 2008: 101.
This second of the species triplet with anterior and posterior pronotal processes is unique in the absence of a lateral carina joining the processes, the pronotum fused to proepipleuron without any evidence of the shared suture (Fig.
(n = 2). Head capsule narrow elongate, genae parallel behind eyes, basally constricted to narrowly constricted neck (Fig.
Male genitalia. Male aedeagal median lobe robust, broadly parallel sided in basal half, ventral margin slightly convex near midlength, dorsoventral breadth at midlength 0.2× distance from tip to base of closed basal bulb (
Female reproductive tract. Not examined (see
Holotype male (
Known from slightly north of the distribution of C. mirabilis along the SarraméaCanala Road near Table Unio, though with a broad sympatric overlap with that species ranging from Col d’Amieu to Ningua Reserve (Fig.
Among the triplet of species with setose pronotal processes, C. lescheni can be diagnosed by: 1, presence of a carinate ridge between the anterior and posterior pronotal processes; 2, small, flat eyes, eight ommatidia crossed on a horizontal diameter; 3, elytra with seventh interval the most carinate, its convexity marking the border between a moderately flat disc bearing six intervals, and a much more vertical elytral margin with a pelage-seta bearing eighth interval. Standardized body length 5.0–5.4 mm.
(n = 2). Head capsule elongate, genae parallel behind eyes, basally constricted to well-defined neck; frons convex between eyes, frontal grooves linear and moderately convergent anterad, widest at frontoclypeal suture; supraorbital setae absent; mandibles moderately elongate, length 1.9× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, apical antennomeres filiform; scape fusiform, broadened into a paddle-like shape, length 2.5× maximal breadth; gena glabrous except for pelage, subgenal seta absent. Pronotum orbicular; anterior pronotal process short, apically obtuse, the process flattened longitudinally; posterior pronotal process columnar, peg-like, slightly more than twice as long as broad; lateral ridge joining two process a low rounded carina; pronotal median base constricted, surface undulated with median and two lateral depressions separate by low rounded longitudinal ridges, lateral depressions bordering outside by similar low, rounded lateral longitudinal ridges; median longitudinal impression finely incised, moderately deep; anterior transverse impression interpreted as a punctulate transverse groove that defines a rounded anterior collar, the collar broadest medially, terminated laterally anterad pronotal processes; proepipleuron and proepisternum bulging outward, visible in dorsal view; proepisternum smooth; prosternal process medially concave anteriorly near front of procoxae, deeply medially depressed on ventral face, broadly depressed medially on posterior face. Elytra narrow, elongate, with scutellum and parascutellar interval depressed relative to very carinate seventh interval, the carination of that interval greatest at base; parascutellar and dorsal setae absent; humeri extended anterad at base of seventh interval, basal margins straight and narrowly diverging posterad humeral angle to position laterad anterior three lateral elytral setae where the margins become subparallel posteriorly; lateral elytral setae arranged as 6 + 3 + 4, the setal articulatory sockets papillate, raised as mounds above the surrounding cuticle; subapical elytral seta present, apical seta absent; subapical sinuation distinctly concave, margin upraised at middle of concavity, the sinuation subangulately meeting lateral margin. Mesepisternum impunctate; metepisternal dorsal length 1.3× diagonal width. Abdomen with apical margin of apical ventrite moderately concave medially in females, the middle pair of the four apical setae just laterad median concavity. Legs moderately elongate, gracile; mt1 length/tibial length = 0.21; mt4 length to apex of outer lobe 1.2× median length, with 4–5 ventrolateral setae each side. Microsculpture of head and pronotal disc glossy, without microsculpture; elytral disc with intervals glossy, indistinct isodiametric (?) sculpticells in deepest portions of striae; pelage well developed, comprising club-like to paddle-like setae; head capsule densely covered with club-like setae that are longest on posterior portion of head capsule, about half a long on frons, and very short, scale-like anterad and below eyes; pronotal disc pelage consisting of setae much like present on posterior portion of head, pronotal processes and lateral ridges with apically broader, more paddle-like setae; elytral intervals lined with longitudinal series of club-like setae, the setae broader apically on odd intervals (sutural, 3, 5, 7), and shorter and narrower apically on even intervals; elytral lateral margin densely lined with short, thick setae; ventral body surface, femora and tibiae covered with very fine, apically narrowed microsetae; tarsomeres with and additional paired dorsolateral setae, four on mt1 and two on mt2–4. Coloration of head and pronotal disc rufous, pronotal lateral ridge somewhat darker due to deeper cuticle; elytra and abdominal ventrites rufoflavous, elytral lateral margin dark rufous (as in pronotal lateral carina); legs flavous.
Female reproductive tract (n = 1). Basal portion of one right gonocoxa available for study; basal gonocoxite with apical fringe of five setae; apical gonocoxite apparently subtriangular based on basal fragment, with two lateral ensiform setae.
Holotype female (
Paratype female (
We take great pleasure in naming this species for Dr. Richard A. B. Leschen, Curator, New Zealand Arthropod Collection and the collector of the two type specimens currently representing this species. Dr. Kipling W. Will is accorded author status based on his recognition of this species as undescribed from amongst the material collected during the 2007 New Zealand Arthropod Collection-University of California, Berkeley Essig Museum expedition.
Both types of this species were collected from leaf litter by Berlese extraction. The long club-shaped setae afford the means for a thick layer of environmental patina to adhere to the body surface.
This species (Fig.
(n = 5). Head capsule broad, stout, genae parallel behind convex, pop-eyed compound eyes; neck constriction broad, moderately deep dorsally but not evidenced laterally; ocular ratio 1.39–1.45, eyes with 20 ommatidia across horizontal diameter; frons medially rugose between eyes, frontal grooves distinctly doubled, the deep canaliculi separated by a distinct rounded ridge, groove convergent to frontoclypeal suture; supraorbital setae absent; mandibles moderately elongate, length 2.1× distance from antennal articulatory socket to anterolateral margin of labrum; antennae elongate, scape length 2.9× maximal breadth; gena setose, subgenal seta present. Pronotum orbicular, front angles briefly but acutely projected; lateral marginal bead subcarinate, elevated, terminated posteriorly in blunt, obtuse tooth that continues medially as a diminished basal marginal bead, though median base not margined; pronotal base constricted, set off from disc by median and two oblique lateral depressions; median longitudinal impression deep, moderately broad, undulated longitudinally with transverse wrinkles of disc not all aligned across impression; anterior transverse impression represented by an oblique series of longitudinal depressions that extend anterad at their ends as two to three longitudinal canaliculi crossing the anterior callosity; two linear parasagittal impressions present, these grooves extended from basal tooth ¾ pronotal length toward front angles; pronotal flanges laterad parasagittal impressions irregularly wrinkled, surface smoother than median pronotal disc; proepipleuron and proepisternum bulging outward, visible in dorsal view; proepisternum irregularly and obliquely wrinkled, the wrinkles oriented more or less longitudinally; prosternum rugose, the surface with deep, irregular wrinkles that are most distinct near anterior margin; prosternal process distinctly margined anteriorly by ridges surrounding procoxal cavities, broadly medially depressed on ventral face, narrowly carinate dorsally on posterior face. Elytra ovoid, disc elevated relative to depressed lateral margins, appearing somewhat cordate due to basal elevation of third to seventh intervals versus depressed scutellum and parascutellar interval; parascutellar seta present, situated at base of sutural interval, elevated on papillate articulatory socket; basal groove broadly rounded laterad depressed and inflexed scutellum; sutural, and elytral intervals 2, 4, and 6 moderately convex on disc, intervals 3, 5, and 7 elevated, subcarinate basally, the intervals broader where they are more convex; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7; subapical elytral seta present, apical seta absent; subapical sinuation well developed, concavity near lateral margin of sinuation extended ventrad to subangulately meet lateral margin, thus apical elytral margin not elevated inside sinuation; elytral apices broader rounded, separated by inflexed margin adjacent to fused suture. Mesepisternum impunctate, metepisternal dorsal length 0.9× diagonal width; metasternum with deep, laterally ovoid median fossa at base of metasternal process. Abdomen with broad, shallow medial concavity at apex of apical ventrite in both males and females; males with one seta each side of apical ventrite, females with two setae. Legs elongate, gracile; metacoxae bisetose; mt1 length/tibial length = 0.27; mt4 length to apex of outer lobe 2.33× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture of head stretched isodiametric mesh, visible over portions of vertex, microsculpture obscured in rugose area of frons and in frontal grooves; pronotal disc glossy, indistinct sculpticells traceable only in deepest portions of wrinkles; elytral disc glossy, obsolete transverse lines visible over portions of apex; sparsely distributed pelage of short microsetae visible on head capsule, prothorax, meso- and metasternum, and coxae; elytra with pelage of microsetae arranged in mediolongitudinal series along elytral intervals; legs sparsely covered with very short, fine microsetae in addition to the larger fixed setae, these short setae also arranged in irregular longitudinal series on the dorsal surfaces of tibiae and tarsomeres. Coloration (specimen clear of environmental patina is teneral; Fig.
Male genitalia (n = 2). Male aedeagal median lobe robust basally, broadly parallel sided in basal half, evenly tapered in distal half to extremely elongate, curved, proboscis-like apex, dorsoventral breadth at midlength 0.16× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-shaped, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 1.5× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Paratypes (26 specimens). NEW CALEDONIA: Mt. Panié, 950–1300 m el., 20°34’S, 164°46’E, 14–16-v-1984, Monteith & Cook (
The species epithet fasciatus is derived from the Latin to “envelop with bands (
This species is known only from Mt. Panié at elevations ranging 950–1600 m (Fig.
Among all the smaller-bodied species with orbicular pronotum and cordate elytra (Figs
(n = 5). Head capsule narrowly ovoid, genae gradually narrowed to moderately constricted neck; eyes only slightly convex, oriented toward upper surface of head, ocular ratio quite variable, 1.20–1.56, about 18 ommatidia across horizontal diameter of eye; frons with paired, longitudinal depressions that surround a low median ridge; frontal grooves broad, deep, parallel, extended from just anterad hind margin of eye onto clypeus; mandibles elongate, length 2.1× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, scape length 2.75× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles rounded, only slightly protruded; median base continuously margined, but bead straight along base; median longitudinal impression finely incised, slightly irregular as it traverses transverse wrinkles; anterior transverse impression a deep, oblique depression, defining a flat anterior callosity with indistinct longitudinal wrinkles; proepisternum slightly extended beyond marginal bead, narrowly visible in dorsal view; proepisternum mostly smooth, with indistinct vertical wrinkles; prosternal process broadly, medially depressed anteriorly, with paired, shallow longitudinal depressions ventrally, posterior face broadly convex. Elytra broadly cordate, basal groove sinuously incurved near depressed scutellum, continued in broad curve laterally around humerus, a slight hitch in the curve near base of third stria; discal intervals moderately convex, all intervals of subequal convexity; elytral striae deep, punctate, the punctures larger basally where they expand strial breadth; parascutellar seta present, articulatory socket papillate; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7, the articulatory sockets papillate; both subapical and apical elytral setae present, the articulatory socket for apical seta smaller; subapical sinuation well developed, lateral portion of concavity joined to depressed lateral margin in broad curve; elytral apices conjoined, no invagination of apical margin at fused suture. Mesepisternum impunctate; metepisternal dorsal length 1.1× diagonal width; metasternum with broad, shallow median fossa at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two setae each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.27; mt4 length to apex of outer lobe 1.6× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture on vertex stretched isodiametric mesh, the head capsule glossy; pronotal disc with evident transverse mesh visible within transverse wrinkles, the surface matte; elytral intervals covered with transverse-line microsculpture resulting in silvery iridescence; pelage little developed, glossy head capsule glabrous, pronotum and elytra with sparse distribution of exceedingly short microsetae, visible only in lateral view; ventrites and legs glabrous except for macrosetae. Coloration of body dorsum piceous, mouthparts, scutellum, and apical elytral margin rufopiceous; body venter including elytral epipleura rufopiceous; femora dark rufous; tibiae piceous medially, rufoflavous at base and apex; tarsi rufoflavous.
Male genitalia (n = 10). Male aedeagal median lobe robust, broadly parallel sided over basal 2/3 of length, the lobe shaft angled ventrally ventral ostial opening, dorsoventral breadth at midlength 0.20–0.25× distance from tip to base of closed basal bulb (Figs
Female reproductive tract (n = 2). Bursa copulatrix vase-shaped, abruptly narrowed distally near spermatheca, distance from base of gonocoxites to spermathecal basal sclerite 3× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Known from Province Nord. The recorded distribution is bounded by Canala on the south, and Col d’Amoss along the northeastern coast (Fig.
This species is characterized by presence of the posterior supraorbital seta, a configuration shared, among the nine Cyphocoleus species with orbicular pronotum and cordate elytra (Figs
(n = 5). Head capsule narrowly ellipsoid, genae gradually narrowed to moderately constricted neck; eyes only slightly convex, ocular ratio 1.46–1.53, 13–18 ommatidia across horizontal diameter of eye; frons with paired, elongate-ellipsoid depressions that surround a narrow elevated median carina; frontal grooves deepest and broadest just mesad eyes, sinuously convergent toward clypeus; mandibles elongate, length 2.3× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, scape length 2.75× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles rounded, slightly protruded; median base depressed relative to disc, lateral margin sinuate anterad median basal margin that is straight across base; median longitudinal impression deep, broad, extended to anterior margin; anterior transverse impression defined by shallow groove where discal wrinkles terminate, anterior callosity crossed by numerous longitudinal wrinkles; proepisternum extended beyond marginal bead, visible in dorsal view, smooth; prosternal process broadly, medially depressed anteriorly and on ventral face, narrowly carinate posteriorly. Elytra broadly ovoid, depressed mediobasally between parascutellar setae; basal groove subangulate basad sutural stria, lateral margin broadly and evenly extended outside humeral angle; discal intervals moderately convex, all intervals of subequal convexity; elytral striae deep, punctate, the punctures limited at the deepest portion of strial depression (and so easy to miss in uncleaned specimens); parascutellar seta articulatory socket papillate; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7, the articulatory sockets papillate; both subapical and apical elytral setae present, the articulatory socket for apical seta smaller; subapical sinuation moderately developed, lateral portion of concavity joined to depressed lateral margin in broad curve; elytral apices conjoined, no invagination of apical margin at fused suture. Mesepisternum impunctate; metepisternal dorsal length 1.0× diagonal width; metasternum with broad, shallow median fossa at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two setae each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.27; mt4 length to apex of outer lobe 1.8× median tarsomere length, 3–4 ventrolateral setae each side. Pelage little developed, glossy head capsule, pronotum, and elytra with only sparsely distributed, very short microsetae. visible only in lateral view; pro-, meso-, and metasternum with evident pelage of microsetae, the setae longer than on dorsal surface; second and third abdominal ventrites with short, sparse microsetae medially, other ventrites and legs glabrous except for macrosetae. Coloration of body dorsum piceous, mouthparts rufopiceous; ventral surfaces of head and prothorax and thoracic sternites piceous, elytral epipleuron and abdominal ventrites to rufopiceous to dark rufous; femora dark rufous except for paler, rufoflavous apex, tibiae and tarsi rufobrunneous.
Male genitalia (n = 14). Male aedeagal median lobe gracile, median lobe constricted near midlength, dorsoventral breadth at midlength 0.13–0.16× distance from tip to base of closed basal bulb (Figs
Female reproductive tract (n = 3). Bursa copulatrix narrowly elongate, distance from base of gonocoxites to spermathecal basal sclerite 7× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype female (
This is the most broadly distributed Cyphocoleus species, distributed along the length of Grande Terre (Fig.
This species and its adelphotaxon C. cardiopterus (Figs
(n = 5). Head capsule relatively broad, little narrower than prothorax, genae slightly convex anterad distinct, moderately constricted neck; eyes distinctly convex, oriented toward upper surface of head, ocular ratio quite variable, 1.43–1.50, 16 ommatidia across horizontal diameter of eye; frons with paired, rugose depressions that bracket median ridge, the ridge narrowed apically due to anterior broadening of depressions; frontal grooves broad, deep, expanded anterad to surround median frontal tubercle, their surfaces arcuately wrinkled; mandibles very elongate, length 2.6× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderate, scape length 2.5× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles angulate, anterior pronotal margin broadly concave; median base continuously margined, bead elevated across base well above scutellum; median longitudinal impression deep, finely incised at depth, straight, extended basally to deep transverse groove just inside basal marginal bead, extended anteriorly nearly to well-defined anterior marginal collar that extends to front angles; anterior transverse impression by deep groove defining anterior limit of shallow transverse wrinkles, anterior callosity smooth, slightly convex behind anterior marginal collar; proepisternum extended beyond marginal bead, visible in dorsal view; proepisternum mostly smooth, with irregularities associated with shallow isodiametric microsculpture; prosternal process deeply, narrowly depressed medially before procoxae, narrowly medially depressed ventrally, posterior face with expanded triangular surface ventrally, median ridge invaginated dorsally. Elytra broadly cordate, basal margin sinuously recurved laterad depressed parascutellar striole, basal groove with deep pit at base of fourth stria; humerus broadly extended laterally at bases of striae 5–8, then evenly curved posterad outside anterior series of lateral elytral setae; discal intervals broadly convex, striae deep, all intervals of subequal convexity; parascutellar seta present, situated twice as far from basal margin than from suture, articulatory socket papillate; dorsal elytral setae absent; lateral elytral setae arranged as (5)6 + (1) + (7)8, the articulatory sockets papillate; both subapical and apical elytral setae present, their articulatory sockets subequal; subapical sinuation shallowly concave, lateral portion of concavity joined to depressed lateral margin in broad curve, medial portion near suture nearly straight; elytral apices conjoined or nearly so, no or little invagination of apical margin at fused suture. Metepisternal dorsal length 0.9× diagonal width; metasternum with deep, circular median fossa at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two setae each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.24; mt4 length to apex of outer lobe 1.6× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture on vertex dense transverse mesh; pelage little developed, head capsule, pronotum, and elytra with sparse distribution of exceedingly short microsetae, visible only in lateral view; pro-, meso-, and metasternites, and mediobasal portions of visible abdominal ventrites 2 and 3 with exceedingly short setae, other ventrites and legs glabrous except for macrosetae. Coloration of head capsule and prothorax piceous; elytra dark rufous with silvery iridescent reflection due to microsculpture; pterothoracic ventrites and elytral epipleura rufopiceous, abdominal ventrites rufous; femora dark rufous with piceous cast; tibiae rufobrunneous; tarsi rufoflavous.
Male genitalia (n = 4). Male aedeagal median lobe robust, broadly parallel sided over basal 3/4 of length, dorsoventral breadth at midlength 0.33× distance from tip to base of closed basal bulb (Figs
Female reproductive tract (n = 1). Bursa copulatrix vase-like, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 2.75 maximum breadth (dissection compressed under cover slip) (as in Fig.
Holotype male (
Paratypes (18 specimens). NEW CALEDONIA: Rivière Bleue, Haute Pourina, 800 m el., pyrethrum trees & logs, 22°06’S, 166°38’E, 22-xi-2000, Monteith (
The species epithet cordatus – the Latin adjective heart-shaped (
This species is restricted to the southern portion of Grande Terre, allopatrically south of the distribution of its adelphotaxon, C. cardiopterus (Fig.
The diagnosis of C. cordatus should be consulted to determine the characters that uniquely circumscribe this species and its adelphotaxon (Fig.
(n = 5). (The description of C. cordatus can serve for this species, with the following exceptions to accompany the diagnostic characters presented above). Lateral elytral setae arranged as 6 + (1) + 7(8), articulatory sockets slightly upraised above cuticular surface, not papillate; elytral disc rufopiceous, without iridescent reflection.
Male genitalia (n = 1). Male aedeagal median lobe robust, broadly parallel sided over basal 2/3 of length, dorsoventral breadth at midlength 0.3× distance from tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 1). Bursa copulatrix vase-like, basally stalked, distance from base of gonocoxites to spermathecal basal sclerite 2.75 maximum breadth (dissection compressed under cover slip) (Fig.
Holotype female (
This species is known from Mt. Mou and near Mt. Dzumac, occupying a distribution allopatric and to the north of its adelphotaxon, C. cordatus (Fig.
(n = 5). Beetles comprising this species (Fig.
Head capsule parallel-sided, appearing “reduviidoid”, the small, little-convex eyes only slightly extended beyond the genal profile in dorsal view; neck only slightly depressed medially, genae only indistinctly narrowed laterally before pronotal front angles; frons with narrow median longitudinal crest surrounded by flattened, rugose, lateral areas that lie mesad the deep, sinuous frontal grooves; frontoclypeal suture reduced, difficult to trace, frontal grooves apparently extended onto clypeus; mandibles elongate, length 2.1× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, scape length 2.5× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles protruded, angulate, the angle right to slightly acute; lateral marginal carinae extended to base, slightly sinuate just before depressed median base, the marginal bead across base irregularly interrupted by convex basal wrinkles of disc; median longitudinal impression deep and broad, separating discal wrinkles, extended nearly to anterior margin but deepest at juncture with transverse anterior impression; anterior transverse impression a depressed oblique border defining an irregularly surfaced anterior callosity bearing distinct longitudinal wrinkles; proepisternum bulging, distinctly extended beyond marginal bead in dorsal view; proepisternal surface dimpled, the irregularities mostly short vertical wrinkles near dorsal margin with proepipleuron, short longitudinal wrinkles ventrally; prosternum deeply depressed medially from pronotal anterior marginal bead to prosternal process, process broadly, slightly depressed on ventral face, posterior face with depressed ventral triangle and rounded medial ridge above. Elytra broadly ovoid, parascutellar region depressed and inflexed, basal margin sinuously subangulate basad stria 4, then broadly extended and posteriorly curved; sutural intervals elevated and fused into median callosity on middle of disc, intervals 3, 5, and 7, and to a lesser degree intervals 4 and 6, subcarinate, very convex on elevated basal portion of disc immediately posterad apex of parascutellar striole; striae deep, not visibly punctate except in extremely clean specimens where very small elongate punctulae can be seen in the deepest portion of the striae, these elongate depressions separated from each other by twice their length or more; parascutellar seta present, articulatory socket distinctly papillate, very upraised; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 6, the articulatory sockets distinctly papillate; both subapical and apical elytral seta present, the articulatory socket for apical seta much smaller, not papillate; subapical sinuation distinctly developed, lateral portion of concavity joined to depressed lateral margin in tightly rounded curve; elytral apices rounded, slightly separated by shallow invagination at fused suture. Mesepisternum impunctate; metepisternal dorsal length 1.1× diagonal width; metasternum with deep, circular median fossa at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two setae each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.24; mt4 length to apex of outer lobe 2.0× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture on vertex densely transverse, shallow, difficult to trace on the glossy surface; higher areas between pronotal transverse wrinkles glossy, depressions of wrinkles and lateral flanges outside parasagittal groove covered with granulate isodiametric sculpticells, the surface matte; pelage well developed, head capsule, pronotum and elytra with densely distributed, evident microsetae, the setae on each elytral interval arranged in two irregular, longitudinal rows; prosternum with elongate pelage setae, meso- and metasternum largely glabrous, and meso- and metacoxae, and abdominal ventrites 2–3 medially with very short microsetae. Coloration of body dorsum piceous; thoracic sternites and elytral epipleura piceous, abdominal ventrites dark rufous; femora and tibiae rufopiceous, their articulation rufoflavous, tarsi rufobrunneous.
Male genitalia (n = 21). Male aedeagal median lobe variably robust, from gracile (Figs
Female reproductive tract (n = 3). Bursa copulatrix columnar, distance from base of r gonocoxites to spermathecal basal sclerite 3× maximum breadth (dissection compressed under cover slip) (Fig.
Lectotype female (
This species is distributed in the southern two-thirds of Grande Terre, being known from Aoupinié on the north to Montagne des Sources on the south (Fig.
Like C. globulicollis (Fig.
(n = 5). Head capsule broad, genae short, neck only slightly constricted; eyes moderately convex but profile not exceeding curvature of gena on posterior portion of eye, ocular ratio 1.47–1.56, 15–16 ommatidia across horizontal diameter of eye; frons with well-developed median carina that is surrounded by transversely wrinkled median spot; frontal grooves narrow between eyes, sinuously extended anterad to a deep, rounded depression just posterad frontoclypeal suture; mandibles elongate, length 2.2× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, scape length 2.75× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles rounded, only slightly protruded, very obtuse; lateral marginal bead distinctly sinuate before depressed median base that is margined medially by an irregularly depressed bead continuous with the base of the median longitudinal impression; median longitudinal impression deep, broad, extended nearly to anterior marginal bead, with pit-like depression near juncture with anterior transverse impression; anterior transverse impression a shallow, irregular border on the anterior margin of the discal transverse wrinkles, defining a rugose, longitudinally wrinkled anterior callosity; proepisternum bulging, extended beyond marginal bead, visible in dorsal view, smooth; prosternal process convex anterad procoxae, slightly depressed medially on ventral face, broadly convex posteriorly. Elytra broadly cordate, basal margin slightly concave to base of fourth stria, then posteriorly curved across broad humeri; discal intervals moderately convex, all intervals of subequal convexity; elytral striae broad, deep, punctate, the punctures expanding strial breadth to elytral apex; parascutellar seta present, articulatory socket broadly papillate; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7, the articulatory sockets papillate; both subapical and apical elytral setae present, the articulatory socket for apical seta smaller, not papillate; subapical sinuation distinctly concave, lateral portion of concavity joined to depressed lateral margin in tightly rounded curve; elytral apices conjoined, margin not invaginated at fused suture. Mesepisternum impunctate; metepisternal dorsal length subequal to diagonal width; metasternum with broad, shallow median fossa at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two seta each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.23; mt4 length to apex of outer lobe 1.9× median tarsomere length, 3–4 ventrolateral setae each side. Pelage little developed, thoracic sternites with only a few very short microsetae, and abdominal ventrites and legs apparently glabrous except for macrosetae. Coloration of body dorsum piceous; prosternum piceous; meso- and metathoracic sternites and pleurites rufopiceous; abdominal ventrites rufous with piceous cast, apical margin of apical ventrite paler, brunneous to flavous; femora dark rufous basally and apically, but with flavous band just beyond midlength; tibiae rufous, matching femoral apex; tarsi brunneous.
Male genitalia (n = 3). Male aedeagal median lobe very short, broad, longer left paramere extended more than half distance from lobe tip to base of closed basal bulb (Fig.
Female reproductive tract (n = 2). Bursa copulatrix columnar, distance from base of gonocoxites to spermathecal basal sclerite 3.5× maximum breadth (dissection compressed under cover slip) (Fig.
Lectotype female (
This species is known only from the disjunct localities of Fonwhary in the northern portion of Province Sud, and Yahoué and Nouméa in the south of New Caledonia (Fig.
This species shares the well-developed dorsal microsculpture, transversely wrinkled pronotum, and punctate elytral striae with C. flavipes (Fig.
(n = 5). Head capsule broad, genae short, neck only slightly constricted; eyes moderately convex but profile not exceeding curvature of gena on posterior portion of eye, ocular ratio 1.44–1.61, 16 ommatidia across horizontal diameter of eye; frons with chevron-shaped median elevation surrounded by hemicircular depressions, the broad part of the chevron continued anteriorly as median flat area bordered by frontal grooves; frontal grooves narrow with lateral marginal carina to mesad eyes, sinuously extended anterad to a broad, rounded depression just posterad frontoclypeal suture; mandibles moderately elongate, length 1.9× distance from antennal articulatory socket to anterolateral margin of labrum; antennae of moderate proportions, scape length 2.4× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles rounded, only slightly protruded, very obtuse; lateral marginal bead slightly sinuate before depressed median base that is margined medially; median longitudinal impression finely incised, interrupted in part by transverse wrinkles, extended nearly to anterior marginal bead; anterior transverse impression a shallow, irregular border on the anterior margin of the discal transverse wrinkles, defining an irregularly, longitudinally wrinkled anterior callosity; proepisternum not extended beyond marginal bead in dorsal view, anteriorly with irregularly, arcuately wrinkled from front to dorsal margin; prosternal process slightly concave anterad procoxae, broadly and deeply depressed medially on ventral face, broadly convex posteriorly. Elytra broadly cordate, basal margin concave from scutellum to third stria, then extended anteriorly across broadly rounded humerus; small pits at the base of the parascutellar and third stria; discal intervals moderately convex, all intervals of subequal convexity; elytral striae finely incised at depth, deep, distinctly punctate, the punctures greatly expanding strial breadth throughout their length; parascutellar seta present, articulatory socket papillate; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7, the articulatory sockets papillate; both subapical and apical elytral setae present, the articulatory socket for apical seta smaller, not papillate; subapical sinuation distinctly concave laterally, subangularly meeting lateral margin, margin slightly convex medially in dorsal view; elytral apices conjoined, margin not or minutely invaginated at fused suture. Mesepisternum impunctate; metepisternal dorsal length subequal to diagonal width; metasternum with broad, shallow median depression at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two setae each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.25; mt4 length to apex of outer lobe 1.8× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture well developed on head, isodiametric sculpticells on median elevation, vertex and neck with more transverse sculpticells defining a regular mesh; elevated portions of pronotal disc with shallow transverse-mesh microsculpture, wrinkled lined with granulate isodiametric sculpticells; elytral intervals with mix of transverse mesh, sculpticell breadth 3× length, and transverse lines loosely joined into a mesh; pelage little developed, head apparently glabrous, prothoracic and elytral disc with extremely short microsetae visible only in lateral view. Coloration of body dorsum piceous; prosternum piceous; meso- and metathoracic sternites, and abdominal ventrites dark rufous, apical margin of apical ventrite paler, brunneous to flavous; femora and tibiae dark rufous to match abdomen, tarsi slightly paler, brunneous.
Male genitalia (n = 3). Male aedeagal median lobe gracile, parallel sided, dorsoventral breadth at midlength 0.18× distance from tip to base of closed basal bulb (Figs
Female reproductive tract (n = 1). Bursa copulatrix columnar, distance from base of gonocoxites to spermathecal basal sclerite at least 2× maximum breadth (dissection broken); bursal walls thin, translucent, lumenal surface smooth in portion examined; basal gonocoxite with apical fringe of five setae (Fig.
Holotype female (
Paratypes (7 specimens found in header-labeled series including holotype and labeled paratype; General Coleoptera Collection,
The species epithet is based on the type locality Bourail, with the -ensis suffix denoting this locality.
As this species is known only from a single series of specimens with the header specimens labeled Bourail, we know only the general collecting locality: Bourail (Fig.
Among species with orbicular pronotum and broadly cordate elytra, this species (Fig.
(n = 5). Head capsule ovoid, neck moderately constricted with genae moderately convergent behind eyes in dorsal view; eyes moderately convex, outer profile extended beyond curvature of gena behind eye, 16–18 ommatidia across horizontal diameter of eye; frons with well-developed median carina that is surrounded by transversely wrinkled median flat spot; frontal grooves narrow, sinuous anteriorly from between eyes onto clypeus, frontoclypeal suture obsolete; mandibles moderately elongate, length 1.9× distance from antennal articulatory socket to anterolateral margin of labrum; antennae moderately elongate, scape length 2.5× maximal breadth; gena setose, subgenal seta present. Pronotum with front angles tightly rounded, slightly protruded, nearly right; lateral marginal bead distinctly sinuate before depressed median base that is margined medially by narrow bead; median longitudinal impression narrowly and finely incised, extended from just inside basal marginal bead nearly to anterior marginal bead, transverse wrinkles on either side of impression not aligned; anterior transverse impression very shallow, irregularly impressed in association with shallow longitudinal wrinkles on anterior callosity; proepisternum not visible in dorsal view, smooth ventrally, with fine vertical wrinkles along dorsal margin; prosternal process slightly convex medially anterad procoxae, with shallow medial depression ventrally, broadly convex posteriorly. Elytra broadly cordate, basal margin slightly concave to base of fourth stria, then posteriorly curved across broad humeri; discal intervals convex, intervals 3–5 slightly more convex, broader basally; elytral striae deep, narrow, impunctate; parascutellar seta present, articulatory socket papillate; dorsal elytral setae absent; lateral elytral setae arranged as 6 + 7, the articulatory sockets papillate; both subapical and apical elytral setae present, the articulatory socket for apical seta smaller, not papillate; subapical sinuation distinctly concave, lateral portion of concavity joined to depressed lateral margin in broad curve, medial margin convex to suture; elytral apices conjoined, margin not or only slightly invaginated at fused suture. Mesepisternum impunctate; metepisternal dorsal length subequal to diagonal width; metasternum with broad, shallow median fossa at base of metasternal process. Abdomen with apical margin of apical ventrite shallowly concave medially in both males and females; males with one seta each side of apical margin of apical ventrite, females with two seta each side. Legs moderately elongate; metacoxae bisetose; mt1 length/tibial length = 0.25; mt4 length to apex of outer lobe 1.6× median tarsomere length, 3–4 ventrolateral setae each side. Microsculpture well developed on pronotal disc, a combination of upraised isodiametric and transverse sculpticells, the surface matte; elytral microsculpture an irregularly present transverse mesh, sculpticell borders shallow, difficult to trace in glossier portions of surface; pelage little developed, head apparently glabrous, pronotal and elytral disc with sparsely distributed, extremely short microsetae, visible only in lateral view; prosternum with only a few very short microsetae, pterothoracic and abdominal ventrites and legs apparently glabrous except for macrosetae. Coloration of body dorsum and venter, including elytral epipleuron, uniformly piceous; legs paler, rufobrunneous with piceous cast except for flavous median femoral band (a minority of specimens lack this pale band and so have concolorous, rufobrunneous leg segments).
Male genitalia (n = 7). Male aedeagal median lobe elongate, heavily sclerotized, parallel sided from base to ostial opening; dorsoventral breadth at midlength 0.13× distance from tip to base of closed basal bulb (Figs
Female reproductive tract (n = 3). Bursa copulatrix columnar, distance from base of gonocoxites to spermathecal basal sclerite 4× maximum breadth (dissection compressed under cover slip) (Fig.
Holotype male (
Paratypes (239 specimens). NEW CALEDONIA: Aoupinié, top camp, 850 m el., pyrethrum trees & logs, 21°11’S, 165°18’E, 02–03xi2001, Burwell & Monteith (
The Latin adjectival subulatus, awl-shaped or pointed (
This species is known from the southern half of Grande Terre, with a northerly outlying locality, based on current specimens, at Aoupinié (Fig.
(n = 1). This species can be characterized as a dark, fuscous version of C. subulatus, its adelphotaxon (Fig.
(The description of C. subulatus can serve for this species, with the following exceptions to accompany the diagnostic characters presented above). Head capsule narrowly ovoid, neck distinctly constricted with genae moderately convergent behind eyes in dorsal view; 17 ommatidia across horizontal diameter of eye; frons with narrow, elevated median carina that is surrounded by ovoid, transversely wrinkled median flat spot; frontal grooves narrow, sinuous anteriorly from between eyes onto clypeus, separated from supraocular groove by narrow ridge, frontoclypeal suture obsolete laterally, broadly and shallowly evident medially; mandibles moderately elongate, length 2.1× distance from antennal articulatory socket to anterolateral margin of labrum. Pronotum with front angles tightly rounded to angulate, slightly protruded, obtuse; median longitudinal impression narrowly and deeply incised; anterior transverse impression indicated laterally as oblique depressions extended toward front angles, absent medially where anterior wrinkle is extended anteriorly onto anterior callosity, very fine longitudinal irregularities traversing callosity. Elytral subapical sinuation distinctly concave, lateral portion of concavity joined to depressed lateral margin in tightly rounded curve; elytral apices rounded, narrowly separated by shallow invagination at fused suture, Abdomen with apical margin of apical ventrite shallowly concave medially in female, two setae each side of concavity. Microsculpture on elytral intervals much reduced, surface glossy, a transverse mesh or transverse lines present in depressed undulations of surface. Coloration of body dorsum and venter, including elytral epipleuron, uniformly piceous; legs rufopiceous.
Male genitalia (n = 1). Male aedeagal median lobe elongate, heavily sclerotized, parallel sided from base to ostial opening; dorsoventral breadth at midlength 0.13× distance from tip to base of closed basal bulb (Figs
Holotype male (
The species epithet is based on the type locality Île des Pins; the -ensis suffix denoting this locality.
The lone holotype is labeled only “I. des Pins” leaving the ecological situation and exact locality within which this species may occur a mystery. To extrapolate from the known habitat of its sister species, C. subulatus, we can predict that beetles of this species occupy dead branches with fungi, rotten logs with and without epiphytes, and the margins of small streams.
Cyphocoleus radiation. The New Caledonian Cyphocoleus fauna exhibits substantial disparity in body forms. However when these forms are aligned using cladistic principles, the disparate body plans become explainable as the sequential evolution of salient characters, with these characters transforming the end products of the lineage in dramatic fashion. The groundplan for Cyphocoleus is built on a common ancestor who lacked flight wings, and thus evolved thickened cuticle and the robust body form as observed in Cyphocoleus lissus (Fig.
The two successive branching lineages to come from the common ancestor – the clade of five species subtended by C. heterogenus, and C. prolixus (Fig.
The sister group to the C. prolixus lineage (Fig.
Subsequent to evolution of the environmental patina, the Cyphocoleus splits into two disparate clades. The first C. miricollis clade includes beetles that are smaller–body lengths 4.8–7.7 mm–but with significantly punctate striae on the elytra (Figs
The adelphotaxon to the C. miricollis clade represents the final iteration of body form in Cyphocoleus, i.e. beetles with orbicular pronotum and broadly cordate elytra (Figs
At 8.1–8.9 mm body length, Cyphocoleus fasciatus is the largest-bodied taxon in this clade, evolutionarily retaining a larger body size similar to that of C. monteithi (8.5–10.5 mm). All subsequently evolved taxa in the orbicular pronotum + cordate elytra clade measure 4.8–7.5 mm body length, with body sizes among all of the species broadly overlapping. Beetles representing this body form have been collected from a broad variety of ecological situations. This is especially so for the abundantly collected C. subulatus, where individuals have been found associated with dead wood, fungi, epiphytes, and stream margins with associated flood debris. Thus this body form is associated not only with the greatest number of known species, but also with the largest variety of microhabitats.
Environmental Patina Function. The development of a varnish-like patina that obscures the deeper features of the cuticle, such as pronotal transverse wrinkles and elytral striae, evolved once within Cyphocoleus in the common ancestor of C. monteithi and its adelphotaxon (Fig.
Cenozoic History of New Caledonia. There is little controversy whether New Caledonia is associated with the Lord Howe Rise, and that this region of eastern Gondwana progressively rifted from the eastern margin of Australia from Late Cretaceous to Paleocene, 90–61 Ma (
The biogeographic pattern exhibited by Homethina is consistent with amphiantarctic vicariance of Gondwana. Initial isolation of the Australian and South American mammal faunas is hypothesized to have started 64 Ma, with vicariance completed by 50 Ma (
Diversification history of odacanthine subtribes and generic-level taxa within Homethina placed in temporal context defined by major geological events and molecular dating estimates (
What if we were to accept the geological scenario whereby New Caledonia subsided completely during crustal extension of the Australian Plate, and only reemerged 37 Ma? This could be rectified with the molecular dating estimates by hypothesizing the extinction of lineages related to Cyphocoleus, these extinct lineages “marking time” at “an undisclosed location” during the interval between the Cretaceous origin of Odacanthini and an Eocene reemergence of New Caledonia (Fig.
I thank the following institutions and curators for access to material that proved essential for the taxonomic revision of Cyphocoleus, cladistic analysis of Odacanthini, and outgroup analysis including Lachnophorini and various taxa of Lebiitae (institutions providing Cyphocoleus material indicated by asterisk after coden, other institutions cited in Suppl. material
Checklist of taxa represented in cladistic analysis of Odacanthini
Data type: Taxonomic checklist
NONA format data file for cladistic analysis of Odacanthini
Data type: Phylogenetic analysis data file
Specimen records for previously described Cyphocoleus spp.
Data type: Species distributions and specimen repositories