Corresponding author: James K. Liebherr (
Academic editor: Dominique Zimmermann
The 15 species of
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
This contribution sets out to establish the phylogenetic context of a taxon of New Caledonian carabid beetles initially proposed as the distinct genus
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.
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Head and mouthpart structures of
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:
Standardized body length used to describe body size is the sum of three measurements:
Dorsal body setation exhibited by each species is summarized in the diagnosis using a chaetotaxic formula expanded from that first proposed for
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.
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
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
55. Supracarinal elytral seta (basad subapical seta) (+): absent (0; Fig.
56. Elytral subapical seta: present (0); absent (1). This seta is plesiomorphically present in
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
95. Aedeagal ventral (right, except for
96. Aedeagal ventral (right, except for
97. Aedeagal dorsal (left, except
98. Aedeagal dorsal (left, except for
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,
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
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
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
Thoracic structures of
The classification of
Species of this subgenus can be diagnosed by the synapomorphic presence of punctures on the prosternum (Fig.
All Australian
To indicate membership of species of this subgenus in
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
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
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. |
3 | Sutural stria and striae 3–6 deep on disc, stria 2 shallower, nearly effaced between positions of dorsal elytral setae (Fig. |
2. |
– | Sutural stria deep on disc, striae 2–6 nearly as deep, stria 2 smooth between positions of dorsal elytral setae (Fig. |
3. |
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. |
– | Male aedeagal median lobe briefly projected beyond ostial opening to a narrowly rounded tip (Fig. |
5. |
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. |
8 | Elytral striae 1–8 fully developed, continuous and smooth from base to apex (Fig. |
7. |
– | Elytral surface smooth, striae at most suggested by obsolete longitudinal depressions on disc, any evidence of striae 1–7 absent apically (Fig. |
8. |
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. |
11. | Elytra orbicular–MEW/EL = 0.96–and very convex (Fig. |
10. |
– | Elytra more ellipsoid–MEW/EL = 0.79–0.84–and moderately convex (Fig. |
11. |
12 | Pronotal hind angles obtuse with rounded apex, not protruded (Fig. |
13 |
– | Pronotal hind angles protruded, nearly right (Fig. |
12. |
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. |
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. |
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. |
– | 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. |
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
Male aedeagal median lobe and parameres of
Paired left (above) and right (below) parameres of
Female reproductive tract, gonocoxites and associated laterotergites,
Strict consensus cladogram of 16 equally most-parsimonious trees derived from cladistic analysis of 65 moriomorphine taxa, including the secondary outgroup
Paratypes (10 specimens). NEW CALEDONIA: Ningua Reserve, camp, 1100 m el.,
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
New Caledonian
(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 aedeagal median lobe and parameres, and ring sclerite–mediotergite plus antecostal margin, tergite IX–of
Paired left (above) and right (below) parameres of
Female reproductive tract, gonocoxites and associated laterotergites,
Left female gonocoxa, ventral view,
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.
Geographical distributions of
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
Paratypes (21 specimens). NEW CALEDONIA: Hienghène [vicinity, code PA 63], 150 m el.,
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.
Though markedly different from all other New Caledonian species, this and the following
(n = 10). The description for
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
(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.
Recorded geographical distribution of
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.
New Caledonian
(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 aedeagal median lobe and parameres, and ring sclerite–mediotergite plus antecostal margin, tergite IX–of
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,
(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.
Paired left (above) and right (below) parameres of
Paratypes (3 specimens). NEW CALEDONIA: Mt. Panié, 1300-1600 m el.,
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.
Geographical distributions of
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.
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
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.
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
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.
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
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.
New Caledonian
(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.
Paratypes (3 specimens). NEW CALEDONIA: Mt. Humboldt, moss forest, 1400 m el.,
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.
Paratypes (11 specimens). NEW CALEDONIA: Mt. Panié,
The adjectival ending -ensis is elided with the type locality Mt. Panié to obtain the species epithet
This species’ distribution is restricted to Mt. Panié (Fig.
This species, along with
(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
Paratypes (33 specimens). NEW CALEDONIA: Mts. Koghis, [code PA 58],
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
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
Male aedeagal median lobe and parameres, and ring sclerite–mediotergite plus antecostal margin, tergite IX–of
Paired left (above) and right (below) parameres of
Female reproductive tract, gonocoxites and associated laterotergites,
Left female gonocoxa of
(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.
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.
Geographical distributions of
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
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
This final member of the species triplet also including
(n = 5). The description of
Paratypes (11 specimens). NEW CALEDONIA: Pic du Pin[code PA 48],
This species name represents the genitive adjectival form of the type locality, Pic du Pin;
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
Given an age of origin for New Caledonian
The distributional ranges of the 15 New Caledonian
Distributional attributes of New Caledonian subgenus
The pattern of speciational diversification for New Caledonian
New Caledonian species of subgenus
The ecological situations within which species of subgenus
The sister-group relationship between
The initial divergence event within
Species now classified in subgenus
The third lineage to diverge from the main
The nominate subgenus
The species included to represent diversity of subgenus
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
Given the preponderance of brachypterous lineages branching off the
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
Infraspecific variation of the male genitalia can be documented for several species of New Caledonian
The amount and manner of variation in the
Number of
This study was made possible through the kind loan offer of significant numbers of New Caledonian
Data file for cladistic analysis of
NONA format data file
Date-locality data for
specimen records