Research Article |
Corresponding author: James K. Liebherr ( jkl5@cornell.edu ) Academic editor: Dominique Zimmermann
© 2018 James K. 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 (2018) Cladistic classification of Mecyclothorax Sharp (Coleoptera, Carabidae, Moriomorphini) and taxonomic revision of the New Caledonian subgenus Phacothorax Jeannel. Deutsche Entomologische Zeitschrift 65(1): 1-63. https://doi.org/10.3897/dez.65.21000
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The 15 species of Mecyclothorax Sharp precinctive to New Caledonia are revised and shown by cladistic analysis to comprise a monophyletic lineage, here treated as subgenus Phacothorax Jeannel. The New Caledonian species of subgenus Phacothorax include Mecyclothorax fleutiauxi (Jeannel), M. najtae Deuve, and 13 newly described species: M. jeanneli sp. n., M. laterobustus sp. n., M. laterorectus sp. n., M. laterosinuatus sp. n., M. laterovatulus sp. n., M. manautei sp. n., M. megalovatulus sp. n., M. octavius sp. n., M. paniensis sp. n., M. picdupinsensis sp. n., M. plurisetosus sp. n., and two jointly authored species; M. kanak Moore & Liebherr sp. n., and M. mouensis Moore & Liebherr sp. n.. subgenus Phacothorax is one of five subgenera recognized within genus Mecyclothorax based on cladistic analysis of 65 exemplar taxa utilizing information from 137 morphological characters. The four other monophyletic subgenera include the precinctive Australian Eucyclothorax subgen. n. (type species Mecyclothorax blackburni [Sloane]), the precinctive Queensland Qecyclothorax subgen. n. (type species Mecyclothorax storeyi Moore), the precinctive New Zealand Meonochilus Liebherr & Marris status n., and the geographically widespread and very diverse nominate subgenus, distributed from St. Paul and Amsterdam Islands, eastward across Australia and New Guinea, and in the Sundas, Timor Leste, Lord Howe and Norfolk Islands, New Zealand, and the Society and Hawaiian Islands. The biogeographic history of Mecyclothorax can be derived from the parsimony cladogram time-calibrated by times of origin of particular geographic areas inhabited by resident representative species. Based on sister-group status of subgenus Phacothorax and subgenus Mecyclothorax, and occupation of Lord Howe Island–an island originating no earlier than 6 Ma–by the earliest divergent lineage within subgenus Mecyclothorax, the ancestor of present-day Phacothorax spp. is hypothesized to have colonized New Caledonia 6 Ma, subsequent both to Cretaceous Gondwanan vicariance as well as any Oligocene submergence. Area relationships among the New Caledonian Phacothorax point to earliest diversification incorporating the northern massifs, and most recent diversification on the ultramafic volcanic substrates in the south of Grand Terre. Flight wing loss has played an important role in shaping the various island faunas, both in their morphology as well as their diversity. The retention of flight capability in only a few of the many hundred Mecyclothorax spp. is presented in light of how populations evolve from macropterous colonizing propagules to vestigially winged specialists. Interspecific differences in genitalic structures for the sister-species pair M. fleutiauxi + M. jeanneli are shown to involve functional complementarity of male and female structures. Extensive geographic variation of male genitalia is demonstrated for several New Caledonian Mecyclothorax spp. This variation deviates from the geographically uniform male genitalia exhibited by species in the hyperdiverse Mecyclothorax radiation of Haleakalā volcano, Maui, suggesting that extensive sympatry occurring among species in that diverse species swarm selects for stability within this mate recognition system. Conversely, lower levels of sympatry characterizing the depauperate New Caledonian radiation permit the presence of more extensive male genitalic variation, this variation not selected against due to the lower likelihood of interspecific mating mistakes.
aptery, conservation prioritization, endemism, genitalic variation, revisionary systematics
The carabid beetle fauna of New Caledonia comprises elements ranging from the mundane to the bizarre. Typical of the more mundane representatives of the fauna would be the geographically widespread species Platycoelus melliei (Montrouzier), native to Australia and present in New Caledonia likely due to human transport (
This contribution sets out to establish the phylogenetic context of a taxon of New Caledonian carabid beetles initially proposed as the distinct genus Phacothorax
Taxonomic material. Revision of New Caledonian Mecyclothorax is based on 676 specimens deposited in 12 institutional or personal collections. The bulk of the material–518 specimens–was collected during various expeditions by Dr. Geoff Monteith and colleagues, Queensland Museum, Brisbane (QMB). Material from the Museum of Natural History, Wrocław University (MNHW) complements the QMB material in the careful annotation of geographic coordinates, elevation, and collecting method or situation. Use of label data from these two institutional collections provided the principal core of information for characterizing microhabitat occupation for the species. Additional material deposited in the following institutional collections was also studied: B. P. Bishop Museum, Honolulu (BPBM); Canadian National Collection, Agriculture Canada, Ottawa (CNC); Cornell University Insect Collection (CUIC); Essig Museum of Entomology, University of California, Berkeley (EMEC); Hungarian Natural History Museum, Budapest (HNHM); Muséum d’Histoire naturelle, Genève (MHNG); Muséum national d’Histoire naturelle, Paris (MNHN); Naturhistorisches Museum, Wien (NHMW); Queensland Museum, Brisbane (QMB); Martin Baehr personal collection (MBC); Pier M. Giachino personal collection (PMGC).
Material collected by Herbert Franz in 1970 (NHMW) were labeled only with a key code, with the codes deciphered in his field notebook (Jäch, pers. comm.).
All type specimens are labeled as such, with data for holotypes presented verbatim, including typographic spacing and capitalization. Individual lines on a label are separated by a slash “/” and different labels are indicated by a double slash “//”. The complete list of scientific names for species included in the cladistic analysis, with authorship, is provided in Table
Species-level taxa included in cladistic analysis. Mecyclothorax subgeneric classification based upon results of cladistic analysis. Type species indicated for Mecyclothorax generic-level taxa. Mecyclothorax andersoni represented by infraspecific aedeagal forms with either a short or long median lobe apex (
Moriomorphina [secondary outgroup] |
Neonomius laevicollis (Sloane) |
Amblytelina [primary outgroups] |
Amblytelus brevis Blackburn |
Amblytelus curtus (F.) |
Amblytelus matthewsi Baehr |
Dystrichothorax amplipennis (MacLeay) |
Epleyx lindensis Blackburn |
Paratrichothorax brevistylus Baehr |
Mecyclothorax Sharp [ingroup] |
subgenus Eucyclothorax Liebherr subgen. n |
Mecyclothorax (Eucyclothorax) blackburni (Sloane) [type species] |
Mecyclothorax (Eucyclothorax) curtus (Sloane) |
Mecyclothorax (Eucyclothorax) eyrensis (Blackburn) |
Mecyclothorax (Eucyclothorax) lophoides (Chaudoir) |
Mecyclothorax (Eucyclothorax) moorei Baehr |
Mecyclothorax (Eucyclothorax) peryphoides (Blackburn) |
Mecyclothorax (Eucyclothorax) punctatus (Sloane) |
Mecyclothorax (Eucyclothorax) sp. n. D |
subgenus Qecyclothorax subgen. n. |
Mecyclothorax (Qecyclothorax) impressipennis Baehr |
Mecyclothorax (Qecyclothorax) inflatus Baehr |
Mecyclothorax (Qecyclothorax) lewisensis Moore |
Mecyclothorax (Qecyclothorax) storeyi Moore [type species] |
subgenus Meonochilus Liebherr & Marris status n. |
Mecyclothorax (Meonochilus) amplipennis (Broun) [type species] |
Mecyclothorax (Meonochilus) bellorum (Liebherr & Marris) |
Mecyclothorax (Meonochilus) eplicatus (Broun) |
Mecyclothorax (Meonochilus) rectus (Liebherr & Marris) |
Mecyclothorax (Meonochilus) spiculatus (Liebherr & Marris) |
subgenus Phacothorax Jeannel |
Mecyclothorax (Phacothorax) fleutiauxi (Jeannel) [type species] |
Mecyclothorax (Phacothorax) jeanneli Liebherr sp. n. |
Mecyclothorax (Phacothorax) kanak Moore & Liebherr sp. n. |
Mecyclothorax (Phacothorax) laterobustus Liebherr sp. n. |
Mecyclothorax (Phacothorax) laterorectus Liebherr sp. n. |
Mecyclothorax (Phacothorax) laterosinuatus Liebherr sp. n. |
Mecyclothorax (Phacothorax) laterovatulus Liebherr sp. n. |
Mecyclothorax (Phacothorax) manautei Liebherr sp. n. |
Mecyclothorax (Phacothorax) megalovatulus Liebherr sp. n. |
Mecyclothorax (Phacothorax) mouensis Moore & Liebherr sp. n. |
Mecyclothorax (Phacothorax) najtae Deuve |
Mecyclothorax (Phacothorax) octavius Liebherr sp. n. |
Mecyclothorax (Phacothorax) paniensis Liebherr sp. n. |
Mecyclothorax (Phacothorax) picdupinsensis Liebherr sp. n. |
Mecyclothorax (Phacothorax) plurisetosus Liebherr sp. n. |
subgenus Mecyclothorax Sharp |
Mecyclothorax (Mecyclothorax) amingwiwae Liebherr |
Mecyclothorax (Mecyclothorax) andersoni Liebherr |
Mecyclothorax (Mecyclothorax) andersoni Liebherr (long aedeagus) |
Mecyclothorax (Mecyclothorax) baehri Guéorguiev |
Mecyclothorax (Mecyclothorax) brispex Liebherr |
Mecyclothorax (Mecyclothorax) goweri Moore |
Mecyclothorax (Mecyclothorax) gressitti Liebherr |
Mecyclothorax (Mecyclothorax) howei Moore |
Mecyclothorax (Mecyclothorax) kavanaughi Liebherr |
Mecyclothorax (Mecyclothorax) medioconstrictus Liebherr |
Mecyclothorax (Mecyclothorax) sedlaceki Darlington |
Mecyclothorax (Mecyclothorax) ambiguus (Erichson) |
Mecyclothorax (Mecyclothorax) basipunctus Louwerens |
Mecyclothorax (Mecyclothorax) globicollis (Mandl) |
Mecyclothorax (Mecyclothorax) lateralis (Laporte de Castelnau) |
Mecyclothorax (Mecyclothorax) lissus (Andrewes) |
Mecyclothorax (Mecyclothorax) marau Perrault |
Mecyclothorax (Mecyclothorax) minutus (Laporte de Castelnau) |
Mecyclothorax (Mecyclothorax) monteithi Moore |
Mecyclothorax (Mecyclothorax) montivagus (Blackburn) [type species] |
Mecyclothorax (Mecyclothorax) oopteroides Liebherr |
Mecyclothorax (Mecyclothorax) otagoensis Liebherr |
Mecyclothorax (Mecyclothorax) punctipennis (MacLeay) |
Mecyclothorax (Mecyclothorax) rectangulus Louwerens |
Mecyclothorax (Mecyclothorax) rotundicollis (White) |
Mecyclothorax (Mecyclothorax) sculptopunctatus (Enderlein) |
Laboratory techniques. Dissection and staining techniques are fully described in
Descriptive conventions. 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. The breadth of the eye socket on the head capsule varies from smaller, with the eye covering less of the extended ocular lobe (Fig.
Head and mouthpart structures of Mecyclothorax spp. A–C. Head capsule, dorsal view: A, M. goweri; B, M. fleutiauxi; C, M. laterorectus. D–I, Mentum, labial palps and ligula: D, M. peryphoides, ventral view; E, M. storeyi, ventral view; F, M. amplipennis, dorsal view; G, M. montivagus, dorsal view; H, M. kanak, ventral view; I, M. fleutiauxi, ventral view. Abbreviations include: aso, anterior supraorbital seta; pso, posterior supraorbital seta.
Elongation of the mandibles was quantified relative to the anterior margin of the labrum by the ratio of the distance from the base of the dorsal (anatomically anterior) condyle of the mandible to the mandibular apex divided by the distance from condylar base to the anterolateral margin of the labrum. Specimens with a non-retracted or angularly retracted labrum were used for this ratio when possible; otherwise the position of a non-retracted labrum was estimated.
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 proepisternum if visible in dorsal view; 2, APW, apical pronotal width, measured between the two most anterior points along the anterior pronotal margin; 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. Two measurements of the elytra were also used to describe body shape: 1, MEW, maximal elytral width; and 2, EL, elytral length, or the distance from the base of the scutellum to the elytral apex, measured parallel to the 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 the ratio MEW/EL. The ranges of all ratios are used for descriptive purposes only without any statistical connotations. When sufficiently available, at least five specimens were chosen for calculation of ratios, encompassing the largest and smallest specimen available, plus male and female representatives. For widespread species, five specimens from various localities were measured in order to give a view to any infraspecific trends in size or shape. The number of specimens used is presented at the start of each description. The number of male and female dissections used to elucidate male genitalic and female genitalic and reproductive tract characters is provided at the start of those sections of the species treatment.
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.
Dorsal body setation exhibited by each species is summarized in the diagnosis using a chaetotaxic formula expanded from that first proposed for Mecyclothorax by
Comprehensive descriptions to complement the diagnoses are provided for all newly described species. The various ratios that quantify proportions of body somites or their constituent structures–e.g., antennae, mandibles, eyes, pronota, elytra–are presented for all species. All character data were recorded in an Excel® spreadsheet, the characters grouped onto sheets according to body region. This spreadsheet is available from the author upon request. Also, many of the character states used as the basis for taxonomic description are also presented in the cladistic analysis (Fig.
Ingroup circumscription. Based on a tribal analysis of Moriomorphini (
Characters. Information from 137 binary or multistate characters was recorded for 65 moriomorphine taxa; 7 outgroup taxa and 58 terminals assignable to the genus Mecyclothorax. In the listing of characters below, multistate characters are presented as either ordered (+) or unordered (–). The initial state descriptions are nearly verbatim relative to the Winclada dataset (Suppl. material
0. 3rd antennomere (–): glabrous except for apical ring of setae (0); a few short setae basad distal ring (1); with 3–4 series of elongate setae along shaft (2); setose in apical half (3). This character is treated as unordered so as not to require that the state of a setose apical half of the antennomere be derived only from the state wherein a few setae accompany the apical ring of setae.
1. Antennomere 9 length/width ratio (+): 1.46 < x ≤ 1.92 (0); 2.0 < x < 2.8 (1); x ≥ 2.89–3.43 (2). The ratio values for the three states span the limits presented.
2. Labrum front margin (+): straight (0; Fig.
3. Mandibular length ratio (+): very short, ratio 1.18–1.55 (0); moderately elongate, ratio of 1.57–1.92 (1; Fig.
4. Apical 2 maxillary palpomeres (+): apparently glabrous (0; Figs
5. Anterior supraorbital seta: present (0; Fig.
6. Frontal grooves: very shallow, not evident between eyes (0); shallow to deep, evident between eyes (1).
7. Frontal grooves (–): subparallel (0; Figs
8. Frontal grooves: not extended onto, or much shallower on clypeus (0; Fig.
9. Neck impression (+) absent, dorsum of head convex (0; Figs 1BC, 8A–B); flat area between hind margins of eye (1); transverse depression behind hind margins of eyes (2; Figs
10. Ocular lobe assessed behind eye (dorsal view): protruded, meeting gena at obtuse angle (0; Fig.
11. Ocular lobe groove behind eye (+): distinct groove present (0; Fig.
12. Mentum tooth: sides acute to subparallel, apex tightly rounded (0; Fig.
13. Mentum width/length ratio (+): 2.3 < x < 3.11 (0); 3.14 < x < 3.5 (1); 3.70 < x < 4.0 (2). This ratio is defined as the maximum breadth of the mentum measured across its convex lateral margins divided by the distance from the mentum-submentum suture to the apex of a lateral lobe. The distributional limits of the states are reflected in the values presented.
14. Apex of ligula (+): apically rounded or with narrow concave apex between setae (0; Fig.
15. Apical two ligular setae: proximate, separated by 1–2 setal diameters (0; Fig.
16. Paraglossae (+): extended only slightly beyond apex of ligula (0); extended 1/2 distance from base to ligular apex beyond apex (1; Fig.
17. Ocular ratio (+): 1.31 < x ≤ 1.42 (0; Fig.
18. Ocular lobe ratio (+): 0.59 < x < 0.72 (0; Fig.
19. Eye convexity (EyL/EyD) (+): convex or popeyed, 2.0 < x < 2.53 (0); moderately convex, 2.55 < x < 2.89 (1; Fig.
20. Number of ommatidia across horizontal diameter of eye (+); 11–15 (0; Fig.
21. Pronotum: broad basally, MPW/BPW = 1.23–1.80 (0; Fig.
22. Pronotal lateral seta: present (0; Fig.
23. Pronotal basal seta: present (0; Fig.
24. Lateral pronotal setal position (if present) (+) with: articulatory socket in or adjoining lateral marginal depression (0); articulatory socket 1/2–2 setal diameters medially from marginal depression (1); articulatory socket 2–4 setal diameters medially from marginal depression (2).
25. Pronotal hind seta (if present): situated at hind angle (0); visibly anterad hind angle (1).
26. Pronotal hind angles (+): nearly right to obtuse, with sides subparallel anterad hind angle (0; Fig.
27. Pronotal hind angle: rounded to right-angled (0; Fig.
28. Pronotal basal margin: straight or convex mesad hind angles (0; Fig.
29. Pronotal median base (+): coplanar with disc medially (0; Fig.
30. Pronotum median base (+): smooth laterally mesad laterobasal depressions (0; Fig.
31. Pronotal basal margin: beaded medially (0; Fig.
32. Pronotal basal marginal bead (if present; i.e. state 0, #31): narrow, continuous with lateral bead (0); broader, discontinuous with lateral bead (1).
33. Pronotal median longitudinal impression (+): well defined, continuous on pronotal disc (0; Figs
34. Anterior transverse impression (+): obsolete to absent medially, anterior callosity not defined (0; Fig.
35. Anterior transverse impression (+): absent to obsolete laterally (0; Fig.
36. Pronotal anterior margin: unmargined (0); with marginal bead medially (1).
37. Pronotal front angles (+): not protruded, rounded behind (0; Fig.
38. Pronotal lateral marginal depression (at midlength) (+): very narrow, margin beaded (0; Fig.
39. Pronotal laterobasal depression (–): broad, smoothly extended to lateral margin (0; Fig.
40. Prosternal process: smooth ventrally, rounded behind (0; Fig.
41. Proepisternum (+): smooth (0; Fig.
42. Prosternum (+): medially convex anterad and between coxal cavities (0); medially depressed only between coxal cavities (1; Fig.
43. Prosternum (+): smooth medially (0; Fig.
44. Lateral portion of prosternum: impunctate except for anteapical furrow (0; Fig.
45. Prosternal anterior margin (+): without or with very shallow indistinct groove (0); with smooth depression or groove (1; Fig.
46. Prosternal anteapical groove (if present at all in some form; i.e. states 1–4, #45) (+): very short, restricted to dorsodistal surface of prosternum (0); longer, extended from dorsodistal margin ventrad onto lateral surface of prosternum (1); continuous ventrally but shallower or irregular between the lateral reaches of prosternum (2; Fig.
47. Proepimeron (+): with anterior and posterior grooves smooth (0); with anterior groove minutely punctate, posterior smooth (1); with both anterior and posterior grooves irregular or punctate (2).
48. Parascutellar seta: present (0); absent (1).
49. Scutellum: narrower, W/L = 0.83–1.86 (0; Fig.
50. Parascutellar striole: longer, terminated between sutural stria and suture where parallel (0; Fig.
51. Parascutellar striole (+): present, continuous, smooth or punctate (0; Fig.
52. Dorsal elytral setal count (+): 0 (0); 1 (1; Fig.
53. Accessory dorsal setae (+); absent (0); in fifth elytral interval (1); in fifth and seventh elytral intervals (2). The presence of accessory setae in the fifth elytral interval is documented for M. kavanaughi (
54. Position of anterior (or single) dorsal elytral seta (–): at anterior 1/5–1/3 of length (0); at midlength (1); in posterior third (2). A single dorsal elytral seta at elytral midlength occurs in M. storeyi and M. lewisensis (
55. Supracarinal elytral seta (basad subapical seta) (+): absent (0; Fig.
56. Elytral subapical seta: present (0); absent (1). This seta is plesiomorphically present in Mecyclothorax and its sister group comprising Amblytelus and allied genera, and is situated within the seventh stria (Fig.
57. Elytral apical seta: present (0); absent (1). This seta occurs just laterad the mesally curved apex of stria 2 near the elytral sutural apex (Fig.
58. Apex of elytral interval 8 (+): with same curvature as elytral apex (0; Fig.
59. Elytral shape (+): broadly subquadrate, sides subparallel, humeri broad (0; Fig.
60. Elytral shape (humeri moderately broad; i.e. state 2 of #59) (+): foreshortened, hemiovoid (0); elongate, ellipsoid (1; Fig.
61. Elytral dorsal curvature (+): moderately convex, sides sloping to margins, disc flat (0; Fig.
62. Elytral basal groove (+): hardly recurved (0; Fig.
63. Elytral humeral angle (+): rounded (0: Fig.
64. Elytral humeral angle: with evenly convex margin on base and lateral edge (0); with elevated humeral tooth at angle (1).
65. Elytral striae on disc (+): 1–8 present, complete (0; Fig.
66. Elytral striae on apex (+): all present, deep (0; Fig.
67. Elytral apex along suture: not upraised in a conjoined juncture (0; Fig.
68. Elytral strial punctation (+): absent (0; Fig.
69. Setal depressions of dorsal elytral setae (+): shallow, extended over 1/4 breadth of interval 3 or less (0; Fig.
70. Elytral stria 8 (+): smoothly impressed along midlength (0; Fig.
71. Elytral lateral marginal depression (+): very narrow throughout length (0; Fig.
72. Subapical sinuation: deep, distinct, more abruptly curved anteriorly (0; Figs
73. Mesepisternal punctures (+): none, surface smooth (0; Fig.
74. Mesosternal/mesepisternal suture: present (0; Fig.
75. Metathoracic flight wings (–): fully developed, venated, with reflexed apex (0; Fig.
76. Metepisternal width/length ratio (–): 0.30 < x < 0.60 (0); 0.60 < x < 0.87 (1); 0.87 < x < 1.46 (2; Fig.
77. Metepisternum-metepimeron suture: distinct, a distinct line (0; Fig.
78. Tarsomeres: dorsally glabrous (0); dorsally setose (1).
79. Metatarsomeres 4 (+): narrow, with short apical lobes (0); broadly triangular, broader apex with very short lobes (1); broad overall, bilobed, apical lobes broad, elongate (2).
80. Suture between abdominal ventrites 1 and 2: straight to sinuous with ventrite 2 not or only slightly depressed (0); sinuous with ventrite 2 distinctly depressed inside curve (1).
81. Suture between abdominal ventrites 2 and 3: shallow, complete, traceable to margin (0); reduced, incomplete, absent laterally (1).
82. Abdominal ventrites 3–5 (–): with irregular linear plaques in lateral reaches (0); with round, smooth depressed plaques in lateral reaches (1); with surface punctate obscuring any other features (2).
83. Apical abdominal ventrite 6 (male) (–): with 1 seta each side (total of 2) (0); with 2 setae each side (total of 4) (1); with 3 setae each side (total of 6) (2); with 4 setae each side (total of 8) (3).
84. Apical abdominal ventrite (female) (–): with 2 setae each side (total of 4) (0); with 3 setae each side (total of 6) (1); with 4 setae each side (total of 8) (2).
85. Apical abdominal ventrite 6 (female) (–): glabrous medially between apical setae (0); with setose patch of 4–5 setae medially (1); with pair of subapical setae medially (2).
86. Microsculpture on frons (–): granulate isodiametric sculpticells (0); evident to shallow isodiametric mesh (1); isodiametric to transverse mesh in transverse rows (2); indistinct transverse mesh or lines (3); absent, surface glossy (4). All terms for microsculptural configurations are derived from
87. Microsculpture of pronotal disc (–): isodiametric in transverse rows mixed with transverse mesh, 2–3× broad as long (0); transverse mesh, sculpticells 3–4× broad as long (1); mix of transverse mesh and transverse lines (2); reduced, surface glossy, with indistinct transverse mesh or lines (3).
88. Microsculpture of pronotal base (–): isodiametric between basal punctures (0); transversely stretched isodiametric between punctures (1); evident transverse mesh, sculpticells 2–3× broad as long (2); indistinct, elongate transverse mesh or unconnected lines (3).
89. Microsculpture of elytral disc (–): a mix of distinct isodiametric and transverse sculpticells (0); a regular transverse mesh, sculpticell breadth 2–3× length (1); a mix of transverse mesh 3–4× broad as long and transverse lines (2); reduced, surface glossy, indistinct mesh plus lines over part (3).
90. Microsculpture on abdominal lateral base: swirling isodiametric and transverse sculpticells (0); reduced, surface glossy with indistinct sculpticells (1).
91. Body coloration (–): dark, elytral margins concolorous or indistinctly, narrowly paler (0); ferruginous, elytra bicolored with outer 4–5 intervals much paler (1); ferruginous, elytra bicolored, outer 3–4 intervals and apex much paler (2).
See Table
Key to abbreviations for structures of male genitalia, and female genitalia and reproductive tracts.
Abbreviation | Structure |
---|---|
aai | male aedeagal apical invagination |
af | apical face of male aedeagal median lobe |
afs | lateroapical fringe setae, female gc1 |
al | apical lobe of male aedeagal internal sac |
ans | apical nematiform setae, female gc2 |
bc | female bursa copulatrix |
bcd | female bursa copulatrix dorsal lobe |
co | female common oviduct |
des | dorsal ensiform seta, female gc2 |
dgd | defensive gland duct |
dgr | defensive gland reservoir |
dl | dorsal lobe of male aedeagal internal sac |
dp | dorsal plate of male aedeagal internal sac |
fl | flagellum of male aedeagal internal sac |
fs | flagellar sheath of male aedeagal internal sac |
fp | flagellar plate of male aedeagal internal sac |
gc1 | basal gonocoxite of female |
gc2 | apical gonocoxite of female |
gp | gonopore of male aedeagal internal sac |
hg | hindgut |
hs | helminthoid sclerite of female |
les | lateral ensiform setae, female gc2 |
la | ligular apophysis of female |
lp | left paramere of male aedeagus |
mac | dorsal macrospicules of male internal sac |
mbs | mediobasal shagreening of female gc1 |
ms | medial setae, female gc 1 |
mtIX | antecostal margin of male mediotergite IX |
ovo | ostial ventroapical operculum, male aedeagus |
r | ramus of female gc1 |
rp | right paramere of male aedeagus |
sd | female spermathecal duct |
sg | female spermathecal gland |
sp | female spermatheca |
spi | spiracle |
vss | ventral spicular sclerite of male internal sac |
92. Antecostal margin of male mediotergite IX (ring sclerite): angulate (0; Fig.
93. Distal terminus of male mediotergite IX antecostal margin: not extended (0; Fig.
94. Aedeagal ventral (right, except M. storeyi) paramere shape (+): elongate, paddle-like, apex broadly subparallel to rounded tip (0); quadrate-conchoid, dorsal and ventral margins subparallel to apex (1; Fig.
95. Aedeagal ventral (right, except for M. storeyi) paramere (+): densely setose on apicoventral margin, apical setae not present (0); densely setose on apicoventral margin, apical setae not differentiated (1; Fig.
96. Aedeagal ventral (right, except for M. storeyi) paramere: with dorsal surface glabrous or a single small seta subapically (0; Fig.
97. Aedeagal dorsal (left, except M. storeyi) paramere shape (+): broad basally, apical half elongate, margins subparallel (0); broad basally, apex short, narrowly rounded, subangulate (1; Fig.
98. Aedeagal dorsal (left, except for M. storeyi) paramere (+): glabrous apically, though setae may be present on the ventral surface (0; Fig.
99. Aedeagal median lobe ostial opening: without ventroapical operculum (0; Fig.
100. Aedeagal internal sac: unilobed, short to long (0; Fig.
101. Aedeagal internal sac (–): with flagellum associated with gonopore (0; Fig.
102. Short to elongate flagellar condition (i.e. state 0 of #101) (+): thin, lightly sclerotized, may be short, sinuous, or long and whiplike (0; Fig.
103. Apical flagellar plate (i.e. state 1 of #101): a donut-like roll, lightly sclerotized or spiculated, gonopore within convexity (0); a well-sclerotized and ridged plate, gonopore on convex side (1; Fig.
104. Aedeagal sac dorsal plate: absent, flagellum and/or flagellar sheath only (0; Figs
105. Aedeagal sac dorsal plate (i.e. state 1 #104): translucent, covered with microspicules (0; Fig.
106. Aedeagal internal sac surface dorsad position of dorsal plate: unarmored (0); with patch of robust macrospicules (1; Fig.
107. Aedeagal internal sac ventral sclerite patch: absent, area basad flagellum and sheath unsclerotized (0); present as distinct sclerite basad flagellum and sheath (1). This ventral sclerotic patch is observed in species of the outgroups, Dystrichothorax and Amblytelus.
108. Aedeagal internal sac ventrobasal spicular sclerite: absent (0); present (1; Fig.
109. Aedeagal median lobe shaft: gracile, aedeagus relatively narrow dorsoventrally, parallel-sided (0; Fig.
110. Aedeagal median lobe tip (–): extended narrowly beyond ostium (0; Figs
111. Broad extension of aedeagal median lobe (i.e. states 2, 3 #110) (+): parallel-sided dorsoventrally to rounded tip (0; Fig.
112. Aedeagal median lobe apical margin (+): smoothly convex to acuminate, no abrupt change in curvature along margin (0; Fig.
113. Aedeagal median lobe apex: smoothly extended past ostial opening (0; Fig.
114. Female bursa copulatrix shape: elongate, columnar, length/width > 2.0–3.25 (0; Fig.
115. Female bursa copulatrix surface (+): membranous, not wrinkled, transparent (0; Fig.
116. Female bursal copulatrix apicoventral surface: membranous as remainder of bursa (0); sclerotized into plate-like structure (1). Bursal sclerites are observed in females of M. kavanaughi (
117. Female bursa copulatrix: of approximately equal diameter throughout length (0; Fig.
118. Spermathecal duct placement (+): on bursa near common oviduct/bursa juncture (0; Fig.
119. Spermathecal duct placement near common oviduct (i.e. state 0, #118): on right side of bursa laterad common oviduct (0; Fig.
120. Female bursa copulatrix configuration (+): unipartite, without dorsal lobe (0; Fig.
121. Helminthoid sclerite at base of spermathecal duct (–): present as elongate sclerotized apodeme (0; Fig.
122. Ligular apophysis on common oviduct: absent (0; Fig.
123. Spermathecal duct: present, spermatheca stalked (0; Fig.
124. Spermathecal duct (i.e. state 0 of #123): subequal to spermathecal length (0; Fig.
125. Spermathecal shape (+): fusiform or ovoid bulb on narrow duct (0; Fig.
126. Spermathecal gland duct: entering on spermathecal duct at spermathecal base (0; Fig.
127. Ramus (gonocoxite VIII of
128. Lateroapical fringe of setae on gonocoxite 1 (+): composed of single seta (0); composed of 2 setae (1; Fig.
129. Medial setae of gonocoxite 1 (–): absent along entire mesal margin (or only 1–2 small seta apically) (0; Fig.
130. Mediobasal surface of gonocoxite 1: smooth (0; Figs
131. Apical gonocoxite 2 (+): broadly subtriangular, apex rounded, mitten-shaped (0); narrowly subtriangular, basal width < 0.4× length (1); more broadly subtriangular, basal width about half length (2); with lateral apodeme basally, basal width 0.6–0.7× length (3; Fig.
132. Lateral ensiform setal count for apical gonocoxite 2 (+): 1 (0); 2–3 (at least 2 unilaterally) (1; Figs
133. Lateral ensiform setae of apical gonocoxite 2: shorter, basal seta 0.25–0.40× gonocoxite length (0; Figs
134. Lateral ensiform setae of gonocoxite 2: on lateral margin of gonocoxite (0; Figs
135. Apical nematiform setae of gonocoxite 2: in fossa 1/4–1/5× length from apex (0; Figs
136. Mesal surface of laterotergite IX: setose (0); spiculose (1). Spiculose laterotergites diagnose the genera Amblytelus, Epelyx, and Dystrichothorax (
Cladistic methods. The character-state matrix was developed using Winclada (
Under all three numbers of ratchet iterations, eight equally parsimonious trees of 1203 steps were found (CI = 0.21, RI = 0.66), with those trees and the character-state changes summarized for presentation using the 1216-step strict consensus cladogram under fast optimization (Fig.
The converse monophyly of Mecyclothorax is supported in the resulting cladogram by many characters that reverse among the dense sampling of Mecyclothorax spp., however the presence of 4–5 setae medially on the apical abdominal ventrite (character 85, state 1) is an unreversed synapomorphy among the species analyzed here. The groundplan elytral striation for Mecyclothorax is more derived than observed among the “Amblytelus” genera, as striae 6 and 7, or 5 to 7 are reduced on the disc (character 65, states 2 to 5), versus all striae present or only the seventh reduced in the former. This strial reduction also affects the expression of striae on the elytral apex, with most Mecyclothorax taxa exhibiting reduction in striae 4–6 or 3–6 apically. The lone exception includes species of subgenus Meonochilus from New Zealand, a clade previously described as a distinct genus (
Thoracic structures of Mecyclothorax and Amblytelus spp. A–B. Pronotum, dorsal view: A, M. laterorectus; B, M. mouensis. C–F. Prosternum, ventral view: C, M. fleutiauxi; D, M. curtus; E, M. blackburni; F, M. moorei. G–J. Elytral apices, dorsal view: G, M. fleutiauxi; H, M. laterorectus; I, M. lissus; J, A. curtus. K–L. Elytral lateral margin showing eighth stria, lateral view: K, M. ambiguus; L, M. blackburni. Abbreviations include: aag, anteapical groove; acc, accessory elytral dorsal setae in interval 5; as, apical elytral seta; ca, carina laterad stria 7; ls, lateral elytral setae; ri, ridge mesad stria 8; sa, subapical elytral seta; sas, subapical sinuation of elytron; scs, supracarinal elytral setae; 8, medial portion of stria 8.
The classification of Mecyclothorax subgenera is based on the results of the cladistic analysis (Fig.
Mecyclothorax Sharp, 1903: 243 (type species Cyclothorax montivagus Blackburn by
Cyclothorax MacLeay, 1871: 104 (not Cyclothorax Frauenfeld, 1868; type species Cyclothorax punctipennis MacLeay by monotypy; synonymy
Thriscothorax Sharp, 1903: 257 (type species Cyclothorax unctus Blackburn by original designation; synonymy
Atelothorax Sharp, 1903: 269 (type species Atelothorax optatus Sharp by monotypy; synonymy
Metrothorax Sharp, 1903: 269 (type species Metrothorax molops Sharp by
Antagonaspis Enderlein, 1909: 488 (type species Antagonaspis sculptopunctata Enderlein by original designation; synonymy
Phacothorax Jeannel, 1944: 84 (type species Phacothorax fleutiauxi Jeannel by original designation; synonymy
Loeffleria Mandl, 1969: 54 (synonymy
Meonochilus Liebherr & Marris, 2009: 10 stat. n. (type species Tarastethus amplipennis Broun designated by
Eucyclothorax subgen. n. (type species Cyclothorax blackburni Sloane hereby designated).
Qecyclothorax subgen. n. (type species Mecyclothorax storeyi Moore hereby designated).
Species of this subgenus can be diagnosed by the synapomorphic presence of punctures on the prosternum (Fig.
All Australian Mecyclothorax species have been examined, with the following species assignable to this subgenus represented in the analysis: M. blackburni, M. curtus, M. eyrensis, M. lophoides, M. moorei, M. peryphoides, and M. punctatus (Table
To indicate membership of species of this subgenus in Mecyclothorax, and to reflect the position of this taxon as the adelphotaxon to the remainder of Mecyclothorax sensu lato, the initial syllable “eu”–the Ancient Greek prefix εύ, meaning “proper” or “true”–is combined with the first name proposed for the genus, Cyclothorax (
These robust-bodied species (Fig.
Given restriction of species in this subgenus to Queensland, the Queensland derived first syllable “qe” is combined with
Species of this subgenus share a robust, convex body shape with Qecyclothorax spp. (Fig.
The six species of this subgenus were revised by
This subgenus holds most of the species-level diversity in the genus, and within those radiations character diversity is rampant, making setal configurations, body form, or mensural characters useless for diagnosis. The best means to diagnose this group is through male genitalic characters, as no species of this subgenus studied to date exhibit a flagellum on the internal sac. Instead, the gonopore is surrounded by either a donut-shaped, soft expansion, or this area of the sac bears a scoop-like flagellar plate, well sclerotized and even ridged, with the gonopore present in membrane lying on the convex surface of this plate (Fig.
The immense diversity of species comprising this subgenus is represented by 25 species in the cladistic analysis. The various species known from Norfolk Island (
The species comprising this subgenus exhibit a remarkable diversity of body forms (Figs
There are 15 species assignable to subgenus Phacothorax, with all species restricted to New Caledonia.
1 | Pronotal base broad, hind angles well defined, MPW/BPW = 1.30–1.40 (Figs |
2 |
– | Pronotal base narrow relative to maximal width, hind angles either broadly rounded, or if well-defined, MPW/BPW = 1.66–3.91 (Figs |
4 |
2 | Anterior and posterior supraorbital setae present; pronotal lateral seta present | 3 |
– | A single posterior supraorbital setae present; pronotal lateral seta absent | 1. M. laterobustus Liebherr sp. n. |
3 | Sutural stria and striae 3–6 deep on disc, stria 2 shallower, nearly effaced between positions of dorsal elytral setae (Fig. |
2. M. laterosinuatus Liebherr sp. n. |
– | Sutural stria deep on disc, striae 2–6 nearly as deep, stria 2 smooth between positions of dorsal elytral setae (Fig. |
3. M. laterorectus Liebherr sp. n. |
4 | Head with only the posterior supraorbital seta present each side, the anterior supraorbital seta lacking (Fig. |
5 |
– | Head with two supraorbital setae each side (as in Fig. |
6 |
5 | Male aedeagal median lobe with apex prolonged beyond ostial opening to acuminate tip(Fig. |
4. M. fleutiauxi (Jeannel) |
– | Male aedeagal median lobe briefly projected beyond ostial opening to a narrowly rounded tip (Fig. |
5. M. jeanneli Liebherr sp. n. |
6 | Standardized body length larger, 4.7–5.9 mm | 7 |
– | Standardized body length smaller, 2.8–4.1 mm | 9 |
7. | Pronotum with only a single lateral seta each side at midlength; prosternum glabrous, metafemora with glabrous posterior margin | 8 |
– | Pronotum with single lateral seta on lateral reaches of disc at midlength, plus 13–14 accessory setae in the pronotal marginal bead, 4–5 setae anterad lateral seta, and 9 seta posterad; prosternum with sparse covering of elongate setae, metafemora with more than 10 fine, elongate setae along posterior margin | 6. M. plurisetosus Liebherr sp. n. |
8 | Elytral striae 1–8 fully developed, continuous and smooth from base to apex (Fig. |
7. M. megalovatulus Liebherr sp. n. |
– | Elytral surface smooth, striae at most suggested by obsolete longitudinal depressions on disc, any evidence of striae 1–7 absent apically (Fig. |
8. M. octavius Liebherr sp. n. |
9 | Pronotal hind angles very obtuse to obsolete, rounded, lateral margin immediately anterad angle at most slightly concave (Figs |
10 |
– | Pronotal hind angles obtuse to nearly right, lateral margin immediately anterad angle distinctly concave, the lateral margin sinuate (Fig. |
12 |
10 | Elytral striae 1–4 evident on disc, shallow to deep, contrasted with obsolete to absent striae 5–7 (Figs |
11 |
– | Elytral striae 1–7 all evident on disc, striae 3–4 shallower than striae 1–2, but striae 5–7 deep, smooth and continuous at elytral midlength (Fig. |
9. M. laterovatulus Liebherr sp. n. |
11. | Elytra orbicular–MEW/EL = 0.96–and very convex (Fig. |
10. M. najtae Deuve |
– | Elytra more ellipsoid–MEW/EL = 0.79–0.84–and moderately convex (Fig. |
11. M. manautei Liebherr sp. n. |
12 | Pronotal hind angles obtuse with rounded apex, not protruded (Fig. |
13 |
– | Pronotal hind angles protruded, nearly right (Fig. |
12. M. paniensis Liebherr sp. n. |
13 | Elytral lateral marginal depression narrow outside anterior series of lateral elytral setae, the depression piceous to its margin to match coloration of elytral disc (Fig. |
14 |
– | Elytral lateral marginal depression broad outside anterior series of lateral elytral setae, translucent brunneous outer reaches of explanate margin contrasted with piceous inner portion and elytral disc (Fig. |
13. M. mouensis Moore & Liebherr sp. n. |
14 | Male aedeagal median lobe rounded apically, with only a very small hitch in the apical margin or no hitch at all (Fig. |
15 |
– | Male aedeagal median lobe with distinct invagination along apical face (Fig. |
14. M. kanak Moore & Liebherr sp. n. |
15. | Male aedeagal median lobe slightly narrowed apically, with tip rounded and smooth, ostial opening asymmetrical apically, its ventroapical margin extended more toward tip than dorsoapical margin (Fig. |
14. M. kanak Moore & Liebherr sp. n., form Q |
– | Male aedeagal median lobe apex broadly downturned, tip broadly rounded, apical margin with or without a minute hitch, ostial opening broadly rounded, symmetrical apically relative to tip (Fig. |
15. M. picdupinsensis Liebherr sp. n. |
These beetles are very robust, with a broad pronotum and broadly convex elytra (Fig.
(n = 5). Head capsule broad, eyes moderately convex, ocular lobe meeting gena at very obtuse angle; 16 ommatidia along horizontal diameter of eye; ocular ratio 1.33–1.42, EyL/EyD = 2.34–3.0; frontal grooves deep, arcuately convergent and deepest just posterad clypeus, briefly and shallowly extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.67; ligular anterior margin rounded to ligular seta, concave between setae, the two setae separated by two setal diameters; paraglossae thin, extended as far beyond ligular margin as their basal length to margin; antennae moderately elongate, antennomere 9 length 2.0× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum moderately constricted basally, with obtuse-rounded hind angles and lateral margins briefly sinuate anterad angles (Fig.
Thoracic structures of Mecyclothorax spp. A–B. Lateral view of elytral epipleuron and meso-and metapleurites: A, M. laterorectus; B, M. fleutiauxi. C–D. Metathoracic flight wing: C, M. punctipennis, showing fully developed flight wing folded under elytron; D, M. lissus, stenopterous wing rudiment that extends beyond metanotum, rudimentary wing venation present at base of strap. Abbreviations include: mms, mesepisternal-mesosternal suture; msp, mesepimeron; mst, mesepisternum; mtp, metepimeron; mtt, metepisternum.
Paired left (above) and right (below) parameres of Mecyclothorax spp. (ectal view): A, M. blackburni; B, M. storeyi; C, M. amplipennis; D, M. montivagus; E, M. laterobustus. As the aedeagal assembly of M. storeyi males is inverted relative to other Mecyclothorax species (
Strict consensus cladogram of 16 equally most-parsimonious trees derived from cladistic analysis of 65 moriomorphine taxa, including the secondary outgroup Neonomius laevicollis, six primary outgroup taxa in Amblytelus and allied genera, and 58 ingroup terminals; 57 Mecyclothorax species plus two genitalic forms of M. andersoni. Species terminals are labeled with species name and three-letter abbreviation of relevant generic or subgeneric name. Full listing of included taxa along with subgeneric assignments is presented in Table
Mecyclothorax spp., dorsal view, with subgeneric assignments: A, M. (Eucyclothorax, subgen. n.) blackburni (type species); B, M. (Qecyclothorax, subgen. n.) storeyi (type species); C, M. (Meonochilus, status n.) amplipennis (type species); D, M. (Mecyclothorax) montivagus (type species); E, M. (Mecyclothorax) lissus.
Male genitalia (n = 1). Antecostal margin of male mediotergite IX distally angulate, little extended (Fig.
Female reproductive tract (n = 1). Bursa copulatrix elongate, length more than twice circumference, surface translucent, membranous (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA / Ningua Res.camp / 12-13Nov 2001 / G.B.Monteith // QM Berlesate 1039 / 21°45’Sx166°09’E / Rainforest, 1100m / Sieved litter // QUEENSLAND / MUSEUM LOAN / Date: Nov. 2003 / No. LEN-1686 (green label) // HOLOTYPE / Mecyclothorax / laterobustus / J.K.Liebherr 2017 (black-bordered red label).
Paratypes (10 specimens). NEW CALEDONIA: Ningua Reserve, camp, 1100 m el., 21°45’S 166°09’E, rainforest, sieved litter, 12-13-xi-2001, lot 1039, Monteith (QMB, 4), near summit, rainforest, 1300 m el., 21°45’S 166°09’E, berlesate, sieved litter, 13-xi-2001, lot 1052, Monteith (QMB, 3); Plateau de Dogny, rainforest, 910 m el., 21°37’S, 165°53’E, berlesate sieved litter, 16-xi-2002, lot 1085, Burwell (QMB, 3).
The very broad and robust body form of these beetles (Fig.
This species is known from two localities, Ningua Reserve and Plateau de Dogny (Fig.
These beetles are very similar in external appearance to those of M. laterorectus (Fig.
(n = 5). Head capsule broad, eyes convex, ocular lobe meeting gena at obtuse angle very close to eye posterior margin; 12–14 ommatidia along horizontal diameter of eye; ocular ratio 1.43–1.50, ocular lobe ratio 0.84–0.89, EyL/EyD = 2.50–2.65; frontal grooves nearly straight from posterior terminus inside anterior supraorbital seta to deepest point just posterad clypeus, briefly and shallowly extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.8; ligular anterior margin narrowly rounded to ligular seta, concave between setae, the two setae separated by two setal diameters; paraglossae thin, extended as far beyond ligular margin as their basal length to margin; antennae elongate, antennomere 9 length 2.25× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum distinctly constricted basally, cordate, hind angles obtuse rounded, lateral pronotal margins subparallel anterad hind angles, then distinctly divergent anteriorly (Fig.
Male genitalia (n = 5). Antecostal margin of abdominal mediotergite IX angulate distally, an elongate distal extension present (Fig.
Male aedeagal median lobe and parameres, and ring sclerite–mediotergite plus antecostal margin, tergite IX–of Mecyclothorax (Phacothorax) spp.: A–B, M. laterorobustus, right view, dorsal view in situ (Ningua); C, M. laterosinuatus, right view (Col d’Amieu); D, M. laterosinuatus, right view (Touho TV tower); E, M. laterosinuatus ring, dorsal view; F, M. laterorectus, right view (Mt. Panié); G–H, M. laterorectus, right vew, dorsal view in situ (Mandjélia); I–J, M. fleutiauxi, right view, dorsal view in situ (Mt. Do); K–L, M. fleutiauxi, right view, sac inverted and everted (Me Maoya); M, M. fleutiauxi, right view (Ningua); N–O, M. jeanneli, right view, dorsal view in situ (Mt. Humboldt, 1350 m); P, M. jeanneli, right view (Mt. Humboldt, 1300 m); Q, M. jeanneli, right view, sac everted (Mt. Dzumac); R, M. jeanneli, right view (Rivière Bleue). See Table
Female reproductive tract (n = 2). Bursa copulatrix elongate, length about twice circumference, surface translucent, membranous (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA / Aoupinie top camp / 2-3Nov 2001 / G.B.Monteith // QM Berlesate 1060 / 21°11’Sx165°18’E / Rainforest, 850m / Sieved litter // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // New Caledonia Mecyclothorax revision / measured specimen 2 / J.K.
Paratypes (83 specimens; BPBM, MNHW, PMGC, QMB): see Suppl. material
Like the preceding and immediately following species, beetles of this species exhibit a basally broad pronotum (Fig.
This species is broadly distributed along mid-latitudinal Grand Terre, from Touho TV tower on the north to Mt. Rembai on the south (Fig.
This third of the species characterized by robust body and a cordate, broad-based pronotum with well-defined hind angles (Fig.
(n = 5). Head capsule broad, eyes very convex, popeyed, ocular lobe meeting gena at obtuse angle close to eye posterior margin; 14 ommatidia along horizontal diameter of eye; ocular ratio 1.37–1.45, ocular lobe ratio 0.81–0.90, EyL/EyD = 2.56–2.89; frontal grooves nearly straight from posterior terminus inside anterior supraorbital seta to deepest point just posterad clypeus, briefly and shallowly extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.7; ligular anterior margin narrowly rounded to ligular seta, concave between setae, the two setae separated by one to two setal diameters; paraglossae thin, extended as far beyond ligular margin as their basal length to margin; antennae moderately elongate, antennomere 9 length 2.0× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum distinctly constricted basally, cordate, hind angles protruded and nearly right, lateral margins distinctly, briefly sinuate anterad angles (Figs
Male genitalia (n = 3). Antecostal margin of abdominal mediotergite IX angulate, little extended distally (Fig.
Female reproductive tract (n = 1). Bursa copulatrix elongate, length more than twice circumference, surface translucent, wrinkled, membranous (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA / Mt. Panié / 16 May 1984 / G. monteith & D. Cook // Q.M. BERLESATE No. 650 / 20.35S X 164.47E / Rainforest, 900 m / Litter // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // genitalia in polyethylene vial with glycerine // HOLOTYPE / Mecyclothorax / laterorectus / J.K.Liebherr 2017 (black-bordered red label).
Paratypes (21 specimens). NEW CALEDONIA: Hienghène [vicinity, code PA 63], 150 m el., 20°41’S 164°57’E, 02-ix-1970, Franz (NHMW, 4); Mandjélia, 700 m el., 20°24’S 164°32’E, 12-v-1984, rainforest, Q.M. berlesate 648, sieved litter, Monteith & Cook (QMB, 2), montane rainforest, 700-780 m el., 20°24’S 164°32’E, Pandanus, 20-xi-2008, Wanat (MNHW, 1), 750-780 m el., 20°24’S 164°32’E, 01-xi-2007, sieved litter, Wanat (MNHW, 5), summit, 750 m el., 20°24’S 164°32’E, hand collecting, 06-07-xi-2001, lot 8750, Burwell & /Monteith (QMB, 1), rainforest, sieved litter, 06-07-xi-2001, lot 1055, Monteith (QMB, 3), flight intercept trap, 29-xi-2003–11-i-2004, lot 11486, Monteith (QMB, 1), 780 m el., 20°24’S 164°32’E, berlesate, sieved litter, 12-xii-2004, lot 11941, Monteith (QMB, 2); Mt. Panié, rainforest, 900 m el., 20°33’S 164°45’E, Q.M. berlesate 650, litter, 16-v-1984, Monteith & Cook (QMB, 1); Roche d’Ouaïème, near Hienghène [PA 65], 500-700 m el., 20°37’S 164°52’E, 03-ix-1970, Franz (NHMW, 1).
As in the preceding two species (Fig.
This species is distributed in the northern reaches of the Chaîne Centrale, from Mandjélia south to near Hienghène (Fig.
Phacothorax
fleutiauxi
Mecyclothorax fleutiauxi , Liebherr & Marris 2009: 10.
Though markedly different from all other New Caledonian species, this and the following M. jeanneli (Fig.
(n = 10). The description for M. jeanneli (below) serves to describe the external anatomy of this species, with the following exceptions: 18–20 ommatidia along horizontal diameter of eye; ocular ratio 1.43–1.52, ocular lobe ratio 0.89–0.94, EyL/EyD = 3.09–3.33; MPW/PL = 1.24–1.30; MEW/EL = 0.68–0.73; either one or two dorsal elytral setae associated with third interval; if one, the posterior seta is absent (in 2 of 10 specimens scored).
Male genitalia (n = 17). Antecostal margin of abdominal mediotergite IX robust distally, broadly angulate and truncate (Fig.
Female reproductive tract (n = 5). Bursa copulatrix bilobate, an elongate ventral lobe with length more than twice circumference, and a short dorsal lobe that is about as long as broad and which bears the spermathecal duct at its apex (Fig.
Type – Holotype male (MNHN): La Foa à Canala / fev. 1907 La Foa // TYPE (red label) // Phacothorax / Fleutiauxi / Jeannel (male dissected with genitalia separately mounted on card with label “Phacothorax / fleutiauxi ? Jeann.”). So that the separated genitalia can remain unambiguously associated with the male holotype body, a black-bordered red holotype label has been added to that pin.
Additional taxonomic material (145 specimens; EMEC, MNHW, QMB): see Suppl. material
This species is recorded from middle latitudes of Grand Terre, from Aoupinié on the north to Ningua Reserve on the south (Fig.
The diagnosis for M. fleutiauxi serves to summarize external diagnostic characters for this species (Fig.
(n = 10). Head capsule narrowly elongate, with large, moderately convex eyes, ocular lobe meeting gena at obtuse angle, a broad shallow groove indicating juncture; 20 ommatidia along horizontal diameter of eye; ocular ratio 1.42–1.51, EyL/EyD = 2.8–3.4; frontal grooves sinuously canaliculate, deepest posterad frontoclypeal suture at a line between posterior margin of antennal fossae, briefly prolonged onto clypeus (as in Fig.
Male genitalia (n = 23). Antecostal margin of abdominal mediotergite IX robust distally, broadly angulate and obliquely truncate (Fig.
Recorded geographical distribution of M. fleutiauxi (including population samples A–E northwest of dashed boundary) and M. jeanneli (including population samples F–K southeast of dashed boundary): numbers of males sampled for each locality indicated in lower left of boxes. Sampled population localities for M. fleutiauxi include: A, Me Maoya; B, Rembai; C, Gelima; D, Mt. Do; E, Ningua. Sampled population localities for M. jeanneli include: F, Mt. Humboldt, 600 m; G, Mt. Humboldt, 1300–1600 m; H, Mt. Ouin; I, Mt. Dzumac; J, Rivière Bleue; K, Mts. Koghis.
Female reproductive tract (n = 8). Bursa copulatrix bilobate, a broad ventral lobe complemented by narrower dorsal lobe of subequal length, dorsal lobe bearing spermathecal duct at apex (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA.9944 / 22°11’Sx166°01’E [sic, crossed out and hand corrected to 31’E] / Mt Koghi, 750m / 29Nov2000.GB Monteith / Pyrethrum, trunks&logs // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // HOLOTYPE / Mecyclothorax / jeanneli / J.K.Liebherr 2017 (black-bordered red label). The type locality longitude is incorrect on the label, and is corrected (
Paratypes (156 specimens; EMEC, HNHM, MNHW, NHMW, QMB): see Suppl. material
This species epithet is a patronym honoring Dr. René Jeannel, whose extensive body of literature dominates 20th Century carabidology. Dr. Jeannel’s early and perspicacious appreciation of Wegenerian historical biogeography was ground breaking in Entomology (
This species is distributed in the southern end of Grand Terre, from Mt. Humboldt on the north to Forêt Nord on the south (Fig.
These beetles (Fig.
(n = 3). Head capsule trapezoidal, neck broad, with small, moderately convex eyes, ocular lobe meeting gena at obtuse angle very close to eye posterior margin; 18–19 ommatidia along horizontal diameter of eye; ocular ratio 1.28–1.42, ocular lobe ratio 0.88–0.91, EyL/EyD = 2.9–3.1; frontal grooves very deep, arcuately convergent at midlength, extended deeply onto clypeus; mandibles moderately elongate, mandibular ratio 1.8; ligular lateroanterior margin rounded to ligular seta, the two setae separated by one to two setal diameters; paraglossae thin, extended as far beyond ligular margin as half of basal length to margin; antennae very elongate, antennomere 9 length 3.4× maximal breadth; antennomere 3 with sparse fine setae near apex in addition to apical ring of longer setae. Pronotum transversely ovoid, median base depressed relative to disc, lateral margins evenly curved to meet narrow median peduncular collar, pronotal lateral margin obtusely concave at the juncture (Fig.
Male genitalia (n = 1). Antecostal margin of abdominal mediotergite IX angulate, an elongate “butterfly-net handle” extension distally (Fig.
Male aedeagal median lobe and parameres, and ring sclerite–mediotergite plus antecostal margin, tergite IX–of Mecyclothorax (Phacothorax) spp.: A–B, M. plurisetosus, right view, dorsal view in situ (Aoupinié); C–D, M. megalovatulus, right view, dorsal view of righ sclerite (Mt. Panié); E–F, M. octavius, right view, dorsal view in situ (Me Maoya); G–H, M. laterovatulus, right view, dorsal view in situ (Aoupinié); I–J, M. manautei, right view, dorsal view in situ (Mt. Humboldt, 1400 m); K–L, M. paniensis, right view, dorsal view in situ (Mt. Panié); M–N, M. mouensis, right view, dorsal view in situ (Mt. Mou). See Table
Female reproductive tract (n = 1). Bursa copulatrix unilobate, elongate, length about 3× circumference, walls thin, slightly wrinkled (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA 8716 / 21°11’Sx165°18’E.850m / Aoupinie, top camp, / 2-3Nov2001.C.Burwell& / GMonteith.pyr.trees,logs // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // New Caledonia Mecyclothorax revision / measured specimen 3 / J.K.
Paratypes (2 specimens): same locality and labeling as holotype (QMB, 1); Aoupinié summit, 1000 m el. [984 m;
The presence of accessory setae on the pronotal lateral margins (Fig.
This species is known only from the upper elevations of Mt. Aoupinié (Fig.
This and the following species, M. octavius, can be diagnosed by their large body size; in this species standardized body length = 5.4–5.8 mm. Individuals of this species have the elytral striae well developed, with striae 1–8 deep and continuous throughout their length (Fig.
(n = 4). Head capsule trapezoidal, neck broad, eyes broad and little convex, ocular lobe meeting gena at very obtuse angle well behind eye posterior margin; 20 ommatidia along horizontal diameter of eye; ocular ratio 1.37–1.44, ocular lobe ratio 0.74–0.82, EyL/EyD = 2.81–2.86; frontal grooves narrow, well incised, sinuously convergent to just posterad clypeus, extended briefly onto clypeus; mandibles moderately elongate, mandibular ratio 1.83; ligular margin rounded to ligular seta, concave between the two setae, setae separated by one to two setal diameters; paraglossae thin, extended as far beyond ligular margin as half of basal length to margin; antennae elongate, antennomere 9 length 2.9× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum vase-shaped, narrow basally, lateral margins only slight concave anterad rounded hind angles, median base convex, not depressed relative to disc, without marginal bead (Fig.
Male genitalia (n = 1). Antecostal margin of abdominal mediotergite IX angulate, broad and little distended (Fig.
Female reproductive tract (n = 1). Bursa copulatrix elongate, length more than twice basal circumference, apical portion expanded laterally toward right, surface thick apically, wrinkled, densely stained relative to bursal base (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA 8764 / 20°34’Sx164°46’E / Mt Panie refuge, 1300 m / 8-9Nov.2001.C.Burwell / Pyrethrum, trees & logs // QUEENSLAND / MUSEUM LOAN / DATE: Sept 2002 No. LE 02.43 (green label) // New Caledonia Mecyclothorax revision / measured specimen 3 / J.K.
Paratypes (3 specimens). NEW CALEDONIA: Mt. Panié, 1300-1600 m el., 20°35’S 164°46’E, 15-v-1984, Monteith & Cook (QMB, 1), refuge, 1300 m el., summit, 1600 m el., 20°34’S 164°46’E, rainforest, sieved litter, 9-xi-2001 lot 8769, Burwell (QMB, 1), track, 1500 m el., 20°34’S 164°46’E, pyrethrum trees & logs, 09-xi-2001, lot 8768, Burwell (QMB, 1).
Large body size and ovoid pronotum and elytra (Fig.
This species is only known from 1300–1600 m elevation on Mt. Panié, in the northern portion of the Chaîne Centrale (Fig.
This species (Fig.
(n = 1). Head capsule elongate, distinctly broader at the small, moderately convex eyes, ocular lobe gradually curving to meet gena well behind eye posterior margin; 20 ommatidia along horizontal diameter of eye; ocular ratio 1.45, ocular lobe ratio 0.71, EyL/EyD = 2.4; frontal grooves deep, sinuously convergent to just posterad clypeus, extended deeply onto clypeus; mandibles moderately elongate, mandibular ratio 1.8; ligular lateroanterior margin rounded to ligular seta, the two setae separated by one to two setal diameters; paraglossae thin, extended twice as far beyond ligular margin as their basal length to margin, apex observably spiculate (100×); antennae elongate, antennomere 9 length 2.3× maximal breadth; antennomere 3 with glabrous except for apical ring of setae. Pronotum transverse, vase-shaped, median base depressed relative to broadly convex disc, lateral margins evenly curved until joined to long median peduncular collar, pronotal lateral margin obtusely concave at the juncture (Fig.
Male genitalia (n = 1). Antecostal margin of abdominal mediotergite IX angulate, not extended (Fig.
Type –Holotype male (MNHN): NEW CALEDONIA / Me Maoya, summit / plateau.12Nov2002. / Monteith & Burwell // QM Berlesate 1080 / 21°22’Sx165°20’E / rainforest, 1400 m/ / sieved litter // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // New Caledonia Mecyclothorax revision / measured specimen 1 / J.K.
Given that the body form comprises a pair of ovoids (Fig.
The single specimen of this species was collected from sieved litter collected at the 1400 m summit of Mt. Maoya (Fig.
This and the following species M. najtae (Fig.
Standardized body length 4.1 mm.
(n = 1). Head capsule broad, foreshortened, eyes small, moderately convex, ocular lobe meeting gena at very obtuse angle; 15–16 ommatidia along horizontal diameter of eye; ocular ratio 1.39, ocular lobe ratio 0.89, EyL/EyD = 3.13; frontal grooves nearly straight from posterior terminus inside anterior supraorbital seta to deepest point just posterad clypeus, briefly and shallowly extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.8; ligular anterior margin narrowly rounded to ligular seta, concave between setae, the two setae separated by one to two setal diameters; paraglossae thin, extended as far beyond ligular margin as their basal length to margin; antennae elongate, antennomere 9 length 2.5× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum vase-shaped, hind angles broadly subangulate, lateral margins only slightly concave anterad angles (Fig.
Male genitalia (n = 1). Antecostal margin of abdominal mediotergite IX angulate, not extended (Fig.
Type – Holotype male (MNHN): NEW CALEDONIA 11665 / 21°11’S 165°16’E. / Aoupinie, summit. 1000 m / 2Oct2004. G.Monteith / pyrethrum, trees&logs // QUEENSLAND / MUSEUM LOAN / DATE: July 2005 / No. LE 05.24 // New Caledonia Mecyclothorax revision / measured specimen 1 / J.K.
Though this species is separated in the dichotomous key and therefore in the species treatment sequence from its broad-bodied phylogenetic relatives (Fig. 9AC), this species is given the compound adjectival epithet laterovatulus to signify the broad body and ovoid pronotum coupled with rounded pronotal hind angles (Fig.
The lone holotype of this species was collected on Aoupinié summit at 1000 m elevation via application of pyrethrin spray to trees and logs (Fig.
Mecyclothorax
najtae
This, the second of the small-bodied species with very obtuse pronotal hind angles and basally straight lateral pronotal margins (Fig.
(n = 1). As a complement to
Type – Holotype female (MNHN): HOLOTYPE (red label) // Nouvelle-Caledonie / Menazi 1020 m / 18.x84 Tillier”/”Bouchet // Mecyclothorax / najtae sp. n. / det. T. Deuve // New Caledonia Mecyclothorax revision / measured specimen 1 / J.K.
The type locality northwest of the summit of Menazi (Fig.
This species can be diagnosed by small body size–standardized body length 3.5–3.9 mm–and the transversely ovoid pronotum with evenly convex lateral margins basally, i.e., hind angles lacking (Fig.
(n = 4). Head capsule quadrate, eyes very convex, popeyed, ocular lobe meeting gena at obtuse angle close to eye posterior margin; 14 ommatidia along horizontal diameter of eye; ocular ratio 1.48–1.52, ocular lobe ratio 0.88–0.89, EyL/EyD = 2.2–2.4; frontal grooves deep, slightly convergent to deepest portion just posterad clypeus, deeply extended onto clypeus; mandibles elongate, mandibular ratio 1.92; ligular anterior margin rounded to ligular seta, concave between setae, the two setae separated by one to two setal diameters; paraglossae thin, extended twice as far beyond ligular margin as their basal length to margin; antennae elongate, antennomere 9 length 2.25× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum transversely ovoid, lateral margin evenly convex anterad base, hind angles absent though suggested by presence of short peduncular collar at base of pronotum (Fig.
Male genitalia (n = 1). Antecostal margin of abdominal mediotergite IX broadly angulate, not extended (Fig.
Female reproductive tract (n = 1). Bursa copulatrix length slightly greater than circumference, its surface thin, translucent, only slightly wrinkled (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA 11138 / 21°53’SX166°24’E.1400m. / Mt Humboldt, moss forest. / 6-7Nov2002. Monteith & / Burwell.pyreth,trees&logs // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // New Caledonia Mecyclothorax revision / measured specimen 1 / J.K.
Paratypes (3 specimens). NEW CALEDONIA: Mt. Humboldt, moss forest, 1400 m el., 21°53’S 166°24’E, pyrethrum trees & logs, 06-07-xi-2002, lot 11138, Monteith/Burwell (QMB, 2), beyond summit, 1500 m el., 21°53’S 166°25’E, 07-xi-2002, lot 11122, Burwell (QMB, 1).
This species epithet is a patronym honoring Joseph Manauté, Directeur du Parc Provincial de la Rivière Bleue chez Province Sud, who provided helicopter support for the Queensland Museum expedition to Mt. Humboldt, allowing Geoff Monteith and Chris Burwell to collect the type series of this species as well as other interesting and important taxa (
The species is known only from elevations 1400–1500 on Mt. Humboldt (Fig.
These beetles can be diagnosed by the parallel lateral margins at the base of the pronotum, resulting in very slightly obtuse hind angles that protrude laterally (Fig.
(n = 5). Head capsule elongate, eyes small, convex, ocular lobe-genal juncture evenly curved, a very shallow groove indicating limit of ocular lobe; 14 ommatidia along horizontal diameter of eye; ocular ratio 1.35–1.42, ocular lobe ratio 0.77–0.82, EyL/EyD = 2.0–2.49; frontal grooves narrowly incised, straight and convergent to deepest portion at frontoclypeal suture, deeply extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.71; ligular anterior margin narrowly rounded, the two ligular setae separated by one to two setal diameters; paraglossae thin, extended twice as far beyond ligular margin as their basal length to margin; antennae elongate, antennomere 9 length 2.62× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum distinctly cordate, lateral margins slightly convergent anterad protruded, obtusely-angulate hind angles, the lateral margin immediately divergent anterad subparallel lateral margins at pronotal base (Fig.
Male genitalia (n = 1). Antecostal margin broadly angulate, not extended (Fig.
Female reproductive tract (n = 1). Bursa copulatrix length slightly greater than circumference, its surface thin, translucent, not wrinkled (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA / Mt Panie summit / Nov 2001 / C. Burwell // QM Berlesate 1058 / 20°34’Sx164°46’E / Rainforest, 1600m / Sieved litter // QUEENSLAND / MUSEUM LOAN / DATE: Sept 2002 No. LE 02.43 (green label) // HOLOTYPE / Mecyclothorax / paniensis / J.K.Liebherr 2017 (black-bordered red label).
Paratypes (11 specimens). NEW CALEDONIA: Mt. Panié, 20°34’S 164°46’E, 08-x-1977, J. Balogh (HNHM, 1), 1300-1600 m, 15-v-1984, Monteith & Cook (QMB, 1), E trail, 1350–1629 m el., 20°35.3’S 164°46.2’E, rainforest, 24-xi-2010, Wanat & Ruta (MNHW, 3), refuge, 1300 m el., 20°34’S 164°46’E, rainforest, sieved litter, 8–9–xi–2001, lot 1056, Burwell (QMB, 2), summit, 1600 m el., 20°34’S 164°46’E, rainforest, sieved litter, xi-2001, lot 1058, Burwell (QMB, 2), summit, 1600, 20°35’S 164°46’E, 18-xi-2000, 9939, Bouchard, Burwell & Monteith (QMB, 2).
The adjectival ending -ensis is elided with the type locality Mt. Panié to obtain the species epithet paniensis, an adjective in the genitive case.
This species’ distribution is restricted to Mt. Panié (Fig.
This species, along with M. kanak and M. picdupinsensis, comprises a triplet of cryptic sibling species best determined using the male aedeagal median lobe. To that end, this species is characterized by a median lobe with the apex dorsally and ventrally subparallel beyond the ostial opening, and with the lobe’s apical face narrowly flattened (Fig.
(n = 5). Head capsule elongate, eyes small, very convex, ocular lobe-genal juncture evenly curved, a very shallow groove indicating limit of ocular lobe; 11 ommatidia along horizontal diameter of eye; ocular ratio 1.43–1.48, ocular lobe ratio 0.73–0.77, EyL/EyD = 2.17–2.38; frontal grooves narrowly incised, sinuously convergent to pit at frontoclypeal suture, briefly extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.73; ligula narrowed to a moderately broad, slightly convex anterior margin, the two ligular setae separated by one setal diameter; paraglossae thin, extended twice as far beyond ligular margin as their basal length to margin, apex visibly spiculate (100×); antennae moderately elongate, antennomere 9 length 2.0× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum very transverse, distinctly cordate, lateral margins sinuately concave anterad obtuse-rounded hind angles (Figs
Male genitalia (n = 8). Antecostal margin of mediotergite IX angulate, not extended (Fig.
Female reproductive tract (n = 1). Bursa copulatrix length subequal to circumference, its surface thin, translucent, slightly wrinkled (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA / Mt Mou summit / 24 May 1984 / G. Monteith & D. Cook // Q.M. Berlesate 659 / 22.04S X 166.21E / Rainforest, 1200 n / Litter // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // HOLOTYPE / Mecyclothorax / mouensis / B.P. Moore & J.K. / Liebherr 2017 (black-bordered red label).
Paratypes (33 specimens). NEW CALEDONIA: Mts. Koghis, [code PA 58], 22°11’S 166°31’E, 30-viii-1970, Franz (NHMW, 3), ~5 km N Noumea, 550 m el., 22 10.5’S, 166 30.3’E, forest litter, 25-xi-2009, Schuh (NHMW, 1), forest, 520 m el., 22°10.7’S 166°30.3’E, sifted litter, 25-x-2008, Wanat (MNHW, 9), La Roussette, 600 m el., 22°10.8’S 166°30.7’E, sifted litter, 27-x-2008, Wanat (MNHW, 4), track entrance, 500 m el., 22°11’S 166°31’E, berlesate, 06-v-2005, lot 12264, Monteith (QMB, 4), 500 m el., 22°10’S 166°31’E, 26-viii-1978, S. & J. Peck (CNC, 1), 400-500 m el., 22°11’S 166°31’E, primary forest, 18–19-x-1998, Löbl (MBC, 1; MHNG, 6); Mt. Mou, summit, 1200 m el., 22°04’S, 166°21’E, rainforest litter, 24-v-1984, Monteith & Cook (QMB, 2), top camp, 1150 m el., 22°04’S, 166°21’E, berlesate, sieved litter, 27-xii-2004, lot 12013, Monteith (QMB, 2). Non-type female specimen: Forêt Cachée, 250 m el., 22°11.5’S 166°47.2’E, sifted litter, 26-x-2008, Wanat (MNHW, 1).
As in the species above, this species epithet elides the type locality, Mt. Mou, with the genitive, adjectival ending -ensis. As Dr. Barry P. Moore had both diagnosed this species from the following M. kanak through the use of male genitalic characters, and had chosen a holotype specimen, he is given senior author status for this species.
This species is restricted to the southern portion of Grande Terre, from Mt. Mou on the west, Mts. Koghis north of Noumea, and with a easternmost record from Forêt Cachée based on a single non-type female specimen (Fig.
This second of the species triplet also including M. mouensis and M. picdupinsensis can be diagnosed externally by characters listed in the M. mouensis diagnosis. This species and the following M. picdupinsensis are adelphotaxa based on possession of the synapomorphous hitch on the apical surface of the male aedeagal median lobe (Fig.
Male aedeagal median lobe and parameres, and ring sclerite–mediotergite plus antecostal margin, tergite IX–of Mecyclothorax (Phacothorax) spp.: A–B, M. kanak, right view, dorsal view in situ (Mt. Humboldt, 630 m); C–D, M. kanak, right view, dorsal view in situ (Mt. Humboldt, 1400 m); E, M. kanak, form Q, right view (Mt. Humboldt, 1300 m); F–G, M. kanak, right view, dorsal view in situ (Mt. Dzumac); H–I, M. kanak, right view, dorsal view in situ (Rivière Bleue); J–K, M. kanak, right view, dorsal view in situ (Col de Yaté); L–N, M. picdupinsensis, right views, dorsal view of ring sclerite. See Table
Female reproductive tract, gonocoxites and associated laterotergites, Mecyclothorax (Phacothorax) spp: A, M. manautei; B, M. paniensis; C, M. mouensis; D, M. kanak (Mt. Dzumac; common oviduct artificially darkened to show configuration); E, M. kanak (Pic du Grand Kaori); F, M. picdupinsensis. See Table
(n = 24). Head capsule elongate, eyes small, very convex, ocular lobe-genal juncture evenly curved, a very shallow groove indicating limit of ocular lobe; 12–14 ommatidia along horizontal diameter of eye; ocular ratio 1.41–1.54, ocular lobe ratio 0.74–0.81, EyL/EyD = 2.16–2.45; frontal grooves narrowly incised, sinuously convergent to pit at frontoclypeal suture, extended onto clypeus; mandibles moderately elongate, mandibular ratio 1.60; ligula narrowed to a moderately broad, slightly convex anterior margin, the two ligular setae separated by 1–2 setal diameters; paraglossae thin, extended twice as far beyond ligular margin as their basal length to margin, apex visibly spiculate (100×); antennae moderately elongate, antennomere 9 length 2.1× maximal breadth; antennomere 3 glabrous except for apical ring of setae. Pronotum very transverse, distinctly cordate, lateral margins sinuately concave anterad obtuse-rounded hind angles (Fig.
Male genitalia (n = 25). Antecostal margin of abdominal mediotergite IX angulate, not extended (Fig.
One of a series of nine males from Mt. Humboldt, 1300 m el., 6–7-xi-2002, Monteith & Wright, lot 1076 (QMB) exhibits a broad aedeagus with a rounded tip, the ostial opening more asymmetrical apically (Fig.
The geographic pattern of aedeagal configuration shows that populations in the southern end of the range include males with a longer, narrower, and more curved aedeagal apex (Fig.
Female reproductive tract (n = 4). Bursa copulatrix length 1.1–1.4× circumference, its surface thin, translucent, slightly to not wrinkled (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA / Mt Dzumac / 28 May 1987 / R.Raven // Q.M. Berlesate No. 800 / 22°03’S. 166°28’E. / Rainforest 900m / Litter // QUEENSLAND / MUSEUM LOAN / DATE: Nov. 2003 / No. LEN-1686 (green label) // HOLOTYPE / Mecyclothorax / kanak / B.P. Moore & J.K. / Liebherr 2017 (black-bordered red label).
Paratypes (181 specimens; CUIC, EMEC, MBC, MHNG, MNHW, NHMW, PMGC, QMB): see Suppl. material
This species is named to honor the Kanak people of New Caledonia. As the species epithet kanak is not derived from Latin, it is to be treated as indeclinable (I.C.Z.N. 1999, Article 31.2.3). Dr. B. P. Moore diagnosed this species from the preceding sibling species M. mouensis, and as such it is appropriate that he receive senior authorship for the species.
This species is densely and abundantly distributed across the southern reaches of Grande Terre, with records from Mt. Humboldt south to Forêt Nord (Fig.
Recorded geographical distribution of M. kanak (including population samples A–I for which representative male aedeagi are figured): numbers of males sampled for each locality indicated in lower left of boxes. Sampled population localities include: A, Mt. Humboldt, 1300–1400 m; B, Mt. Humboldt, 600 m; C, Mt. Dzumac; D, Rivière Bleue, 800 m; E, Rivière Bleue, 400 m; F, Mts. Koghis; G, Col de Yaté; H, Pic du Grand Kaori; I, Forêt Nord.
This final member of the species triplet also including M. mouensis and M. kanak can be diagnosed from the former by characters listed in the M. kanak species diagnosis. As stated above, this species cannot be diagnosed from M. kanak based on external characters. The basis for species recognition lies both in the form of the male aedeagal median lobe apex–i.e., broadly rounded with a minute hitch present (Fig.
(n = 5). The description of M. kanak can serve to describe this species, with substitution of the following ranges of ratios of the measured specimens: ocular ratio 1.44–1.52; ocular lobe ratio 0.71–0.80; EyL/EyD = 2.08–2.38; MPW/BPW = 2.30–2.44; MPW/PL = 1.33–1.38; APW/BPW = 1.32–1.36; MEW/EL = 0.85–0.89.
Male genitalia (n = 2). Antecostal margin of abdominal mediotergite IX angulate, not extended (Fig.
Female reproductive tract (n = 1). Bursa copulatrix length subequal to circumference, its surface thin, translucent (Fig.
Types – Holotype male (MNHN): NEW CALEDONIA 11787 / 22°15’Sx166°49’E, 280m / Pic du Pin, site 1,rainfor. / 26Nov2004, berlesate / G.Monteith,P.Grimbacher // QUEENSLAND / MUSEUM LOAN / DATE: July 2005 / No. LE 05.24 // New Caledonia Mecyclothorax revision / measured specimen 1 // HOLOTYPE / Mecyclothorax / picdupinsensis / J.K.Liebherr 2017 (black-bordered red label) /
Paratypes (11 specimens). NEW CALEDONIA: Pic du Pin[code PA 48], 22°15’S 166°49’E, 28-vi-1970, Franz (NHMW, 3), base, 280 m el., 22°14.9’S 166°49.7’E, sifted rainforest, litter, 26-xii-2006, Wanat & Dobosz (MNHW, 1), 22°15’S 166°49’E, sifted litter, 22-x-2008, Wanat (MNHW, 1), site 1, rainforest, 280 m el., 22°15’S 166°49’E, berlesate, 26-xi-2004, lot 11787, Monteith & Grimbacher (QMB, 5), 21-xii-2004, lot 12045, Monteith (QMB, 1).
This species name represents the genitive adjectival form of the type locality, Pic du Pin; picdupinsensis. This final species name includes reference to the New Caledonian Pine, Araucaria columnaris J. R. Forster and W. J. Hooker, and partially recapitulates the final species named in
This species is restricted to the vicinity of Pic du Pin, southern Grande Terre (Fig.
The question regarding time of origin of the New Caledonian Phacothorax radiation is answered by the results of the cladistic analysis. As sister taxa (Fig.
Given an age of origin for New Caledonian Phacothorax of 5–6 Ma, this lineage colonized New Caledonia over water. Subgenus Phacothorax is bracketed on the cladogram by subgenus Mecyclothorax with early divergent taxa in New Guinea, and subgenus Meonochilus of New Zealand. Any biogeographic relationship of Phacothorax and Meonochilus is likely based on overwater dispersal. Zealandic relationships tying together New Caledonia and New Zealand have been demonstrated for stick insects (
The distributional ranges of the 15 New Caledonian Mecyclothorax spp. include a significant proportion of narrowly endemic species. Species richness is concentrated in relatively few localities for this taxonomic data set (Fig.
Distributional attributes of New Caledonian subgenus Phacothorax spp.: A, recorded species richness (open circles) at principal collecting localities represented in this revision (A–S), and species shared between adjacent localities (filled circles and italicized numerals) – see text regarding parallel networks of collection localities in southern end of Grand Terre; B, chorologically oriented cladogram of subgenus Phacothorax spp. (Fig.
The pattern of speciational diversification for New Caledonian Mecyclothorax indicates earlier diversification on the older, continental northern portion of Grand Terre, outside the areas of recently deposited ultramafic rocks. The most morphologically similar and therefore difficult to diagnose cryptic species comprise the clade of four species subtended by M. manautei; also including M. kanak, M. mouensis, and M. picdupinsensis (Figs
New Caledonian species of subgenus Phacothorax appear phylogenetically fickle with regard to occupation of ultramafic soils. In addition to the juxtaposition of M. paniensis on continental rocks versus its four-species adelphotaxon on ultramafic soils, the sister species M. jeanneli and M. fleutiauxi, though as closely related as possible phylogenetically, exhibit a stark dichotomy with regard to occupation of ultramafic soils; the former distributed strictly on such soils, the latter mostly not (Fig.
The ecological situations within which species of subgenus Phacothorax have been collected include both semiarboreal and ground-level microhabitats. The former situations are represented by specimens collecting through the application of pyrethrin insecticide to the mossy surfaces of tree trunks and downed logs (G. Monteith pers. comm.). The area of application spanned the lower trunk or log upwards to where the insecticide fog stopped being effective: less than 4 m or so. Occupation of ground-level situations are guaranteed for specimens collecting by sieving leaf litter with or without subsequent Berlese extraction. Using these criteria, 8 of the 14 species for which data are available have been collected only in sieved litter: M. laterobustus, M. laterorectus, M. laterosinuatus, M. megalovatulus, M. mouensis, M. octavius, M. paniensis, and M. picdupinsensis. Four species have been collected only through pyrethrin fogging: M. fleutiauxi, M. jeanneli, M. manautei, and M. plurisetosus. Two species have been collected using both methods: the very abundantly collected M. kanak, and M. megalovatulus. Occupation of ground-level microhabitats is the groundplan ecological condition for this radiation, with this behavior symplesiomorphically shared with the other basal lineages of Mecyclothorax: subgenera Eucyclothorax, Qecyclothorax, and Meonochilus (see below). The strict utilization of semiarboreal microhabitats by M. plurisetosus, M. fleutiauxi, and M. jeanneli is ecologically synapomorphous, as these three species constitute a triplet of species terminating one of the clades of the radiation (Fig.
The sister-group relationship between Amblytelus and allied genera (
The initial divergence event within Mecyclothorax involved separation of the taxa newly proposed as subgenus Eucyclothorax (Fig.
Species now classified in subgenus Qecyclothorax constitute evolutionary products of the next divergence event along the Mecyclothorax line (Fig.
The third lineage to diverge from the main Mecyclothorax stem, those species in subgenus Meonochilus (Fig.
The nominate subgenus Mecyclothorax is by far the most diverse subgeneric taxon recognized here. Males for all species of this clade for which males are known, exhibit a modification of the male aedeagal internal sac whereby the apex of the sac bears either a roll of thickened cuticle, or a hardened, scooplike sclerite that is associated with the gonopore (Fig.
The species included to represent diversity of subgenus Mecyclothorax are a small subset of all known species. Only one of the 108 French Polynesian Mecyclothorax (
The interpretation of carabid beetle flight wing loss under Dollo Parsimony is consistent with flight wing polymorphism in carabid beetles and the observed transformation over ecological time from populations comprising predominantly macropterous individuals to those within which nearly all individuals lack wings. Such a change was documented over only two seasons in a population of Trechus obtusus colonizing a newly available polder in the Netherlands (
Given the preponderance of brachypterous lineages branching off the Mecyclothorax radiation, with isolated radiations occupying New Zealand, New Caledonia, New Guinea, and Polynesia, it is tempting to suggest that the many independent origins of island lineages during the evolution of Mecyclothorax may have been accomplished by 100% brachypterous propagules. Such a conclusion is contrary to the evidence for at least some of the radiations. The Hawaiian Mecyclothorax include the most generalized species, M. montivagus, individuals of which retain stenopterous wing stubs 3.3× long as broad with rudimentary Sc, R, M, and Cu veins present, indicating this species’ recently evolved brachypterous condition. Similarly, species of the generalized M. striatopunctatus group in Tahiti–e.g. M. curtisi
Isolation of closely related carabid beetle species is mediated by physical incompatibility between the male aedeagal intromittent organ and the female bursa copulatrix (
Occurrence of either bilobed male aedeagal internal sacs or bilobed female bursae has been documented repeatedly for species in subgenus Mecyclothorax. Males of M. bilobatus Liebherr from Haleakalā, Maui have bilobed internals sacs, with the large scoop-like flagellar plate characteristic of the subgenus located at the apex of the apical lobe (
Infraspecific variation of the male genitalia can be documented for several species of New Caledonian Phacothorax recognized in this revision. Males of M. laterosinuatus exhibit uniform flagellar structures associated with the internal sac (Fig.
The amount and manner of variation in the M. kanak male aedeagus is much greater than that observed within species of the very speciose Mecyclothorax radiation from Haleakalā, Maui (
Number of Mecyclothorax spp. present in latitude × longitude quadrats for two Hawaiian Islands: A, Haleakalā volcano, Maui (area 1450 km2) with species recorded within 1´ × 1´ cells; B, Hawaiˋi Island (10,432 km2) with species recorded within 5´ × 5´ cells. A total of 116 species of Mecyclothorax are known to occur on Haleakalā (
This study was made possible through the kind loan offer of significant numbers of New Caledonian Mecyclothorax specimens made by Dr. Geoff Monteith, Queensland Museum. Receipt of additional specimens from other curators resulted in access to specimens from nine institutional and two personal collections. I thank the following curators and collection managers (institutional coden presented in Material and Methods follows parenthetically): James Boone (BPBM); Yves Bousquet (CNC); Jason Dombroskie, Cornell University Insect Collection (CUIC); Peter Oboyski and Kipling W. Will (EMEC); Győző Szél, Hungarian Natural History Museum, Budapest (HNHM); Giuluo Cuccodoro, Muséum d’Histoire naturelle, Genève (MHNG); Thierry Deuve and Azadeh Taghavian (MNHN); Pawel Jałoszyński (MNHW); Manfred Jäch (NHMW); Susan Wright (QMB); Martin Baehr personal collection (MBC); Pier M. Giachino personal collection (PMGC). Additional specimens were used for the cladistic analysis of Mecyclothorax and allied genera, with the following curators and collection managers thanked for access to that material: Robert L. Davidson, Carnegie Museum of Natural History, Pittsburgh; Borislav Guéorguiev, National Museum of Natural History, Sofia; Peter Hudson, South Australian Museum, Adelaide; Beulah Garner, The Natural History Museum, London; Kate Lemann, Australian National Insect Collection, Canberra; John Marris, Lincoln University Entomological Research Collection, Canterbury, New Zealand; Catriona McPhee, Museums Victoria, Melbourne; Philip D. Perkins, Museum of Comparative Zoology, Harvard University, Cambridge; Roberto Poggi, Museo Civico di Storia Naturale “G. Doria”, Genova; Chris Reid, Australian Museum, Sydney; Danny Shpeley, Strickland Entomological Museum, University of Alberta, Edmonton. I thank Manfred Jäch for providing a copy the pertinent pages of Herbert Franz’s field notebook that detail the itinerary and stops during his 1970 New Caledonian expedition. Photographic equipment essential to this research project was underwritten by National Science Foundation award DEB-0315504. I thank Borislav Guéorguiev and Kipling W. Will for their comprehensive critical editorial reviews, which substantially improved the manuscript.
Data file for cladistic analysis of Mecyclothorax Sharp.
Data type: NONA format data file
Date-locality data for Mecyclothorax (Phacothorax) species represented by more than 50 specimens.
Data type: specimen records