Corresponding author: James K. Liebherr (
Academic editor: Thorsten Assmann
The 101 species of
The area of Tahiti Island comprises 1045 km2, yet as is shown by this taxonomic revision, the island supports more than 100 precinctive carabid beetle species in the genus
This contribution returns to the question of Tahitian
This contribution first and foremost documents the currently known diversity of
Tahiti has recently seen the adventive introduction of various alien pest species, among these the little fire ant,
This study is based on 1,992 Muséum National d’Histoire Naturelle, Paris Bernice Pauahi Bishop Museum, Honolulu Cornell University Insect Collection Essig Museum of Entomology Collection, University of California, Berkeley Field Museum of Natural History, Chicago Natural History Museum, Basel U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC
This paper follows directly the laboratory protocols presented in
Treatments for all newly described species include an extended diagnosis, description of male and female genitalia if known, type specimen and species epithet information, and notes regarding geographical distribution, collecting method, or habitat. The extended diagnosis format includes an initial section that provides criteria to distinguish the species from all others known, followed by brief synopses of characters of the head, pronotum, elytra, microsculpture, and coloration. Holotype label data are transcribed verbatim, with individual lines of label text indicated by a slash, “/”, and separate labels indicated by a double slash; “//.” Many species epithets are derived from the Tahitian language, with meanings taken from
Various ratios of length and width are used to describe shapes of the head, pronotum and elytra, necessitating the availability of a measuring reticle in the microscope ocular. For the head these include: 1, the ocular ratio, the maximum head width across the outer surface of the compound eyes (MHW) divided by the minimum width of the frons between the eyes (mFW); and 2, the ocular lobe ratio, the diameter of the eye measured from above, divided by the distance from the front margin of the eye to the juncture of the ocular lobe and gena, measured from the same vantage point. Prothoracic dimensions presented as ratios include: MPW, maximum pronotal width; BPW, basal pronotal width, measured between the hind angles; APW, apical pronotal width measured between the two most anterior points at the pronotal front angles; and PL, pronotal length measured along the midline. Elytral dimensions include MEW, or maximum elytral width, and HuW, humeral width, measured between the most anteriorly positioned points, i.e., the humeral angles. Standardized body length comprises the sum of three values: 1, the length of the head from labral anterior margin to cervical ridge, the position of the ridge estimated from its lateral reaches when hidden medially under the pronotal margin; 2, median pronotal length; and 3, the length of the longer elytron from the basal ridge of the scutellum to its apex, measured parallel to the suture.
When more than one individual was available for the calculation of ratios, the number of individuals used for ratio calculation in that particular description is provided – as “(n = X)” – after the first mention of a ratio, with X representing the number of individuals measured. For larger series, a maximum of five individuals were so measured, with the largest individual, the smallest individual, and representatives of both sexes included in the sample of five. By this method the most disparate range of ratios was sought, although the ratios are used only for descriptive purposes and are not statistically evaluated. Measured specimens bear a numerical tag with values 1–5.
Massifs of the Tahiti Nui and Tahiti Iti volcanoes from which
Sharp’s four species were described with the type data limited to a statement of the numbers of specimens collected for each, along with ecological data and recording that E.C. Zimmerman was the collector. Following British Museum (Natural History) convention, he placed a round, red-bordered “Type” label on one of the specimens, and round yellow-bordered “Paratype” labels on the remaining members of the type series. The series were divided, with the “Type” and various numbers of “Paratypes” returned to Bishop Museum, where they were accessioned as holotypes and paratypes. However as no single specimen was designated as holotype in the description, all specimens of the type series for each species are syntypes. Given that Britton’s type series were divided between London and Honolulu, and given the subtle differences between the various Tahitian species, lectotypes are designated below to stabilize the nomenclature of these species. In all cases, the specimen bearing the round red-bordered “Type” label is chosen as the lectotype. The remaining syntypes become paralectotypes, and those in the Bishop Museum are so labeled.
This contribution aims to provide support for unambiguous identification of the
Setation. Carabid beetles exhibit macrosetae at specific positions on the external surface of the body, with the presence and number of seta extremely useful for diagnosing taxa. All
Head, pronotum, and elytra of
In the generalized condition the
The elytra bear setae at standardized positions across Tahitian
Elytral setation is completed by two series of lateral setae situated in the eighth elytral stria. The anterior series of lateral elytral setae consists of six to seven setae, more commonly seven, with specimens of several species exhibiting bilateral variation from six to seven setae. When seven setae are present, the third and fifth from the front are longest, with the anterior seta nearly as long. The posterior series of lateral elytral setae ranges from four to six setae, most commonly six. As in the anterior series, individuals may vary bilaterally by one or occasionally two setal counts. When six setae are present the second, fourth and sixth setae in the series are longer.
This contribution follows Perrault in presenting a setal formula (e.g.,
Microsculpture. The nature of reticulations in the cuticular surfaces is extremely useful for discriminating otherwise extremely similar species. This study follows the general terminology established by
Male genitalia. The aedeagal median lobe and internal sac, as well as the associated parameres, exhibit substantial variation across the species of Tahitian
Aedeagal orientation is complicated by torsion of the male intromittent organ during the evolutionary history of adephagan
The more generalized configuration of the median lobe entails an apex that is narrowly rounded (
Male aedeagal median lobe and associated parameres,
The parameres associated with the median lobe lie along the lobe’s ventral surface, and articulate at an apodeme situated at the juncture of the basal bulb and the narrower lobe shaft (
The median lobe internal sac in Tahitian
Alcohol preservation of the male specimen of
Spermatophore from male specimen of
Female reproductive tract. Although we lack sufficient taxonomic coverage to diagnose all closely related species using female reproductive tract characters, a survey of the female reproductive tract across the Tahitian
Female reproductive tract dissections,
Female reproductive tract dissections,
Female reproductive tract dissections,
Dorsal to the bursa copulatrix lies a spermatheca apically situated on a spermathecal duct, the latter entering the dorsal surface of the bursa copulatrix immediately dorsad the ventral juncture of the common oviduct and bursa. The spermatheca bears an appended spermathecal gland, the duct of which enters near the base of the fusiform spermatheca (
The female gonocoxae are bipartite, with a broader less sclerotized basal gonocoxite 1, and a more heavily sclerotized, triangular apical gonocoxite 2 (
Female right gonocoxae,
Female right gonocoxae,
An increased level of setation on gonocoxite 1 is observable in
The shape of gonocoxite 2 varies among species, with the base broadly extended laterally in many species (
In nearly all
This key to groups is based on
1 | Pronotum distinctly narrowed basally, the lateral margins sinuate, straight, or convex anterad the hind angles, the basal width not apparently larger than the apical width | 2 |
– | Pronotum subquadrate or trapezoidal, not or little narrowed basally, the basal width larger than the apical width, the hind angles distinctly angulate ( |
1. |
2 | Standardized body length less than 7.5 mm | 3 |
– | Body size larger, standardized body length 8.0–8.6 mm ( |
2. |
3 | Pronotal lateral margins convex to slightly sinuate for short distance anterad hind angles which are obtuse, blunt, or completely rounded | 4 |
– | Pronotal lateral margins sinuate to distinctly sinuate anterad the hind angles which are well developed, obtuse, right, or in some instances acute | 7 |
4 | Pronotal basal setae present, the pronotum quadrisetose | 5 |
– | Pronotal basal setae absent, only lateral setae present near midlength ( |
3. |
5 | Pronotal lateral margin straight to convex anterad rounded hind angle; a small toothlike projection may be associated with the articulatory socket of the basal seta | 6 |
– | Pronotal lateral margin straight to slightly sinuate anterad distinct, obtuse hind angle ( |
4. |
6 | Elytral striae very shallow, fine, discal elytral intervals nearly flat ( |
species 3–14 |
– | Elytral striae deep, well developed, punctate, the discal elytral intervals convex; pronotal hind angle obtuse, basal seta may be associated with a small jag or toothlike projection along base of lateral margin ( |
5. |
7 | Pronotal basal seta present, pronotum quadrisetose | 8 |
– | Pronotal basal seta absent, pronotum bisetose | 9 |
8 | Elytra narrowly ovoid, little convex, humeri very narrowly rounded; elytral disc brunneous, lateral margins and suture flavous ( |
6. |
– | Elytra broader, humeri more broadly rounded; elytral disc rufobrunneous to rufopiceous, lateral margins and suture rufoflavous; dorsal elytral setae present or absent ( |
7. |
9 | Pronotal lateral marginal depression narrower, broadest basally outside laterobasal depression, narrower anterad | 10 |
– | Pronotal lateral marginal depression subequally broad and explanate throughout length, surface broadly translucent in contrast to opaque disc ( |
8. |
10 | Pronotal lateral marginal depression narrow, especially in apical half of length where margin is beaded or narrowly upturned, pronotal margin distinctly sinuate basally anterad hind angle | 11 |
– | Pronotal lateral marginal depression evident throughout length, pronotal margin extended from disc, margin not beadlike ( |
3. |
11 | Pronotal and elytral bases broad, lateral margin joined to basal groove at nearly right angle at humerus, elytra subquadrate ( |
9. |
– | Pronotal and elytral bases narrow, humeral angle obtuse, subangulate, to rounded at humerus, margin narrowly and evenly rounded posteriorly, elytra ovate to obovate ( |
10. |
1 | Pronotum with two setae each side, one laterally near midlength and the second near or at hind angle | 2 |
– | Pronotum with one seta each side near midlength | 5 |
2 | Pronotum narrowed posteriorly, maximum width at the level of the lateral setae | 3 |
– | Pronotum not constricted basally, maximum pronotal width at the base | 4 |
3 | Both apical and subapical elytral setae present, setal formula 2202; pronotal lateral margin convex anterad hind angle, the angle marked by a minute jag just outside setal articulatory socket; elytral humeri broad, lateral margins subparallel outside anterior lateral setal series ( |
1. |
– | Apical elytral seta present, subapical seta absent, setal formula 2201; pronotal lateral margin briefly sinuate before hind angle, the angle projected; humeri rounded, the elytral lateral margins divergent outside anterior lateral setal series ( |
2. |
4 | Elytral microsculpture consisting of a broad mesh, not or little transverse; pronotal margins broadly rounded anteriorly, lateral marginal depression broadly explanate throughout length ( |
3. |
– | Elytral microsculpture consisting of a very narrow, transverse mesh; pronotal margins parallel for some distance basally, lateral marginal depression narrowed anteriorly ( |
4. |
5 | Pronotum narrowed basally, broadest at the level of lateral setae | 6 |
– | Pronotum not narrowed basally, broader or of subequal breadth across the base versus across positions of pronotal lateral setae | 8 |
6 | One supraorbital seta, setal formula 1101 | 7 |
– | Two supraorbital setae ( |
5. |
7 | Pronotal lateral margins straight or only slightly sinuate anterad the hind angles which are obtuse to right, rounded at the apex ( |
6. |
– | Pronotal lateral margins distinctly convergent anterad the hind angles, the sinuation angulate at point where margins anteriorly diverge, angles acute, laterally projected ( |
7. |
8 | One supraorbital seta each side, setal formula 1101 | 9 |
– | Two supraorbital setae each side, setal formula 2101 | 10 |
9 | Pronotal median base sloped anterad to meet disc, anterior portion of base smooth or with sparse shallow punctures ( |
8. |
– | Pronotal median base distinctly depressed relative to disc, juncture with disc lined with distinct longitudinal strigae and elongate punctures ( |
9. M. mapura Perrault |
10 | Pronotum distinctly transverse, MPW/PL = 1.49, lateral margin depression broad anteriorly ( |
10. |
– | Pronotum little transverse, MPW/PL = 1.26, lateral margin depression narrowed anteriorly ( |
11. |
Of the
Female reproductive tract. The lone female specimen was not dissected.
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hiti el. 2000 m 2-VI- / 2006 lot 02 pyrethrin fog /
The species epithet honors the collector, Dr. Elin Claridge, who collected the specimen during the first and only entomological expedition to Pito Hiti.
The type specimen of this species was collected via application of pyrethrin fog to moss-covered vegetation. Thus within the microhabitat of epiphytic mosses, the beetles may be considered subarboreal in habits.
This is the smallest-bodied species in the
The single holotype female specimen was collected at 1000 m elevation on Mont Marau.
This is a species characterized by dramatically robust proportions, with the very broadly explanate and upraised pronotal lateral margins expanded to a very broad pronotal base (
The pronotal configurations of the male type of
The three specimens of this species have been collected at 1200 and 1300 m elevation on Mont Marau. This species is terricolous, with the 1200 m elevation specimen collected from under rocky debris, and those from 1300 m captured in a vinegar pitfall trap.
This species shares a transverse pronotal configuration (
The only known specimen was collected in litter at 1000 m elevation, on Pihaaiateta along the ridge to Pito Hiti.
Of the
This species has been collected at l100–1400 m elevation along the Mont Aorai ridge. It is a terricolous species, with one specimen collected in moss, and a second captured in a vinegar pitfall trap.
This species comprises beetles possessing only the posterior supraorbital seta and a transverse, bisetose pronotum; MPW/PL = 1.38 and setal formula 1101. The elytral humeri are distinctly angulate and the elytra are broadly ovoid, the lateral margins convex throughout their length (
Male aedeagal median lobe and associated parameres,
The type series was collected from mosses at 1200 m elevation on the Mont Aorai ridge. Subsequently, two individuals were extracted through the use of pyrethrin fog from moss growing on
The pronotum of this species is uniquely shaped among known members of this species group, with the lateral margins concave anterad slightly projected, acute hind angles (
Female reproductive tract. The single female specimen as not dissected.
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hiti el. 2070 m 2-VI- / 2006 lot 01 pyrethrin fog /
The species epithet honors Dr. Jean-Yves Meyer, botanist extradordinaire, who accompanied Dr. Claridge on the expedition to Pito Hiti. The epithet is derived in a manner parallel to that of
The single specimen was collected in a pyrethrin fog sample of moss-covered vegetation at 2070 m elevation on Pito Hiti, only 40 m elevation below the summit.
Of the four
Male genitalia. Aedeagal median lobe blunt apically, with a broadly rounded, slightly convex ventral face and more pointed dorsal projection (
Female reproductive tract. The bursa copulatrix of
Female reproductive tract dissections,
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1275 m / 10-IX-2006 lot 02 /
Allotype female (MNHN) labeled: SOCIETY IS: Tahiti / Tahiti Nui Mont Marau / 1280 m el. 6-XI-1999 / D.A. Polhemus pyr. fog / sta. 1
Paratypes: same data as allotype (CUIC, 2; NMNH, 1); Tahiti Nui, Mont Marau, Mt. Marau road, 1315 m el.,
The species epithet, poria, means fat or corpulent in Tahitian, the epithet signifying the broad outline of the body caused by the basally broadened pronotum and broadly based, laterally convex elytra.
All samples containing this species have come from the Mont Marau ridge between 1275 and 1315 m elevation. All specimens of the type series have been obtained by fogging moss-covered
Among
The holotype specimen was collected between 900 and 1200 m elevation on Mapura, the ridge NNW of Pihaaiateta of the Pito Hiti-Orohena massif.
This species is characterized by the broadest, most robust body, and the acutely protruded, glabrous, pronotal hind angles (
Perrault collected the female holotype, 20-xii-1977, at 1000 m elevation on Mont Marau. Earlier that same year – 29–30-vi-1977 – another female was collected “at night” from the summit of Marau, 1490 m elevation (W.C. Gagné and S.L. Montgomery; BPBM).
Unique among the
The holotype female was beaten from dead leaves at 1400 m elevation on Mont Marau.
1 | Vertex and neck impression glossy, surface smooth, without evident sculpticells; pronotal lateral margins explanate, translucent throughout length, median base with distinct rounded punctures in lateral reaches; two dorsal elytral setae in third interval ( |
12. |
– | Vertex and neck impression with evident, regular transverse-mesh microsculpture; pronotal lateral margins narrowed anteriorly, surface largely opaque with isolated spots of translucent cuticle (best viewed from ventrolateral aspect); third elytral interval lacking dorsal elytral setae ( |
13. |
Relative to
This species is known only from the holotype specimen collected by the botanist F.R. Fosberg, from dead
The largest bodied Tahitian
The holotype female was collected by J. Gourvès at 1900 m elevation on Mont Aorai, whereas the male first reported above was collected by E.M. Claridge at 2070 m on Pito Hiti (EMEC). These two localities are an estimated 5.5 km apart taking the ridge distance between them. The microsculpture of the male is somewhat less developed than that of the female holotype. For the present the two specimens are considered conspecific, with the discovery of a male specimen from Mont Aorai the best arbiter for establishing the conspecificity or distinctiveness of the Aorai and Pito Hiti populations.
Male aedeagal median lobe and associated parameres,
Male aedeagal median lobe and associated parameres,
1 | Basal pronotal seta absent | 2 |
– | Basal pronotal seta present (rare setal configuration observed in 2 of 49 specimens) (Aorai) | 14. |
2 | Pronotal hind angles rounded, either broadly rounded, or more narrowly rounded with the lateral margin straight or only slightly sinuate anterad the angle | 3 |
– | Pronotal hind angles distinctly indicated, basal margin sinuate anterad hind angle, either very briefly sinuate with hind angle denticulate, or more broadly sinuate with the hind angle obtuse | 12 |
3 | Striae deep, distinct; elytral coloration flavous to piceous, but without metallic reflection | 4 |
– | Discal elytral striae shallow ( |
14. |
4 | Pronotal lateral depression and elytral marginal depression posterad humerus broad and deep, anterior lateral elytral seta more than twice distance to elytral margin as breadth of elytral interval 9 | 5 |
– | Pronotal lateral depression and elytral marginal depression posterad humerus narrower, distance of anterior lateral elytral seta to elytral margin no more than breadth of elytral interval 9 | 7 |
5 | Pronotal lateral margin broadly and evenly reflexed along entire pronotal length; elytral humeri angulate, elytral base narrow, MEW/HuW = 2.25–2.50 | 6 |
– | Pronotal lateral margin broadly elevated basally, narrow anteriorly, pronotum appearing cychroid; elytral humeri rounded, elytral base broad ( |
15. |
6 | Pronotal anterior transverse impression complete medially ( |
16. |
– | Pronotal anterior transverse impression effaced medially ( |
17. |
7 | Apex of elytral interval 7 without projected tubercle; pronotal hind angles completely rounded | 8 |
– | Apex of elytral interval 7 with a projected tubercle ( |
18. |
8 | Elytral striae not or indistinctly punctate basally | 9 |
– | Elytral striae distinctly punctate basally, intervals convex ( |
19. |
9 | Elytral microsculpture irregularly isodiametric | 10 |
– | Elytral microsculpture transverse | 11 |
10 | Elytral striae finely punctate basally; elytral lateral marginal depression moderately explanate outside anterior series of elytral lateral setae ( |
20. |
– | Elytral striae smooth, impunctate; elytral lateral marginal depression very narrowly reflexed outside anterior series of elytral lateral setae ( |
21. |
11 | Elytral microsculpture consisting of fine transverse lines with little tendency to form a mesh (Marau) | 22. |
– | Elytral microsculpture a transverse mesh (Aorai) | 23. |
12 | Pronotal hind angles obtuse to obtuse-rounded, pronotal base narrow, MPW/BPW = 1.55–1.95 | 13 |
– | Pronotal hind angles toothlike, little projected ( |
24. |
13 | Pronotal lateral margins divergent immediately anterad obtuse-rounded hind angles, margin only slightly concave resulting in little-developed sinuation | 14 |
– | Pronotal basal margins subparallel laterad laterobasal depression, the hind angles prominent, distinctly obtuse, lateral pronotal margin distinctly sinuate | 15 |
14 | Elytra ovate, maximum width near midlength ( |
25. |
– | Elytra obovate, humeri extremely narrow, maximum elytral width well behind midlength ( |
26. |
15 | Discal elytral striae well developed, associated intervals broadly and evenly convex; both apical and subapical elytral seta present, setal formula 2122; standardized body length 4.5–5.3 mm | 16 |
– | Discal elytral striae shallow, associated intervals only slightly convex ( |
27. |
16 | Pronotum broadly transverse, MPW/PL = 1.33, basally constricted, MPW/BPW = 1.72 ( |
28. |
– | Pronotum less transverse, MPW/PL = 1.25, broader basally, MPW/BPW = 1.59 ( |
29. |
Among species of the
This species was previously known (
This is the broadest bodied species of the
Male genitalia. Aedeagal median lobe dorsoventrally expanded apically, with a slightly convex ventral expansion and blunt dorsal tooth (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2000 m 2-VI- / 2006 lot 02 pyrethrin fog /
The species epithet aano means wide or extensive in Tahitian, the name signifying the very broad body characteristic of this species.
This species is known only from the holotype obtained in a pyrethrin fog sample of moss-covered vegetation at 2000 m on Pito Hiti.
This species is diagnosable within the species group by the broadly reflexed lateral margins of the pronotum and elytra, the explanate and laterally reflexed margins extending the lengths of both structures (
Like
The type series (
This species is uniquely characterized by a large tuberculate projection that distorts and upraises the eighth elytral interval and seventh stria, dorsad and immediately anterad the subapical sinuation (
The single female specimen was collected near the summit of Mont Aorai at 1900 m elevation.
This species is best diagnosed by the pronotal lateral depressions that are narrowly reflexed anterad the lateral seta, and the narrowly incised, distinctly punctate discal elytral striae (
This species is recorded from Mont Teatara, Tahiti Iti, between 900 and 1100 m elevation. Two male specimens collected in 2006 have associated ecological data; one was found in association with dead, fermenting
This is a gracile species with a narrow orbicular prothorax, MPW/PL 1.25 (
This species’ known distribution ranges 1100–1400 m elevation on Mont Marau. One specimen was collected in a Malaise trap, indicating the well-developed climbing abilities, and a second was beaten from a mix of
Beetles of this species appear superficially similar to, and are distributed sympatrically with those of
This species is known to occupy habitats on Mont Aorai from 1255 to 1940 m elevation. All associated ecological data place it in the axils of dead, decaying
This and the following species,
This species is known from between 1125 and 1400 m elevations on Mont Marau. It has been collected by beating dead tree fern fronds, rotten
Like
This species is known only from the two type specimens collected at 1900 m elevation near the summit of Mont Aorai.
This species is placeable in the
This species was collected at 1100 m elevation on Mont Mauru in association with moss-covered
This species is among the five
This species has been found from 900–1165 m elevation on Mont Teatara, Tahiti Iti. Specimens have been obtained by beating dead and living fern fronds, within the axils of
This species is similar to
This species’ known distribution is restricted to the highest elevations on Mont Aorai; 1650–2000 m. It has been collected by beating moss on trees and shrubs, indicating that the species possesses at least partially arboreal habits.
Beetles of this species are similar to those of
This species has been collected from 1125–1490 m elevation on Mont Marau, always in association with ferns. These situations include beating dead
Befitting its name, this species is characterized by the palest cuticle observed in all Tahitian
Georges Perrault collected the two known specimens together between 800 and 1000 m elevation on Mont Teatara, Tahiti Iti.
Beetles of this species appear quite similar to those of the sympatrically distributed
This species is known from two female specimens collected between 800 and 1000 m elevation on Mont Teatara, Tahiti Iti.
Male aedeagal median lobe and associated parameres,
Male aedeagal median lobe and associated parameres,
1 | Elytral striae superficial, shallow to obsolete, smooth to indistinctly punctate, elytral intervals flat; if intervals appear slightly convex, elytral margins contrastedly paler than elytral center | 2 |
– | Elytral striae deeper, elytral intervals moderately convex to convex, elytral center and lateral margin concolorous though marginal depression may be narrowly paler | 5 |
2 | Center of each elytron brunneous to piceous, the sutural interval and lateral margin broadly and contrastedly paler, rufoflavous; elytral striae 1–7 evident though stria 7 may be shallower | 4 |
– | Entire elytral surface concolorous, dark rufobrunneous, only the lateral marginal depression and sutural interval paler, the margin rufoflavous and sutural interval dark rufous; elytral striae extremely shallow, finely incised | 3 |
3 | Pronotum narrow, MPW/PL = 1.16 ( |
30. |
– | Pronotum transverse, MPW/PL = 1.31 ( |
31. |
4 | Elytral striae smooth basally, impunctate though with slight irregularities along length ( |
32. |
– | Elytral striae indistinctly punctate in basal portion of disc ( |
33. |
5 | Elytra with only the apical seta present | 6 |
– | Elytra with two setae apically, the apical seta near the apex of stria 2, and the subapical seta associated more basally with stria 7 | 13 |
6 | Pronotum distinctly transverse, MPW/PL = 1.25–1.33 | 7 |
– | Pronotum little transverse, MPW/PL = 1.17–1.25 | 9 |
7 | Two supraorbital setae over each eye; standardized body length 5.3–5.8 mm | 8 |
– | A single, posterior supraorbital seta over each eye; standardized body length 4.6–4.8 mm | 34. |
8 | Elytra subquadrate, broad basally, MEW/HuW = 2.04 ( |
35. |
– | Elytra narrowed basally, lateral margins narrowly rounded posterad angulate humeri, MEW/HuW = 2.10–2.20 ( |
36. |
9 | Pronotal margins linearly narrowed anterad hind angles ( |
10 |
– | Pronotal lateral margins more broadly explanate anterad the obtuse-rounded hind angles ( |
37. |
10 | Elytral microsculpture consisting of transverse mesh with a tendency to form transverse rows; two dorsal elytral setae in the third interval | 11 |
– | Elytral microsculpture consisting of transverse lines without tendency to form a distinct mesh; dorsal elytral setae absent (Marau) | 38. |
11 | Pronotal disc with distinct transverse microsculpture, the sculpticell margins distinct, resulting in a gratelike surface reflection; pronotal lateral marginal depression moderately broad ( |
12 |
– | Pronotal disc glossy, with indistinct elongate transverse microsculpture, the sculpticells not visible in areas of reflected light; pronotal lateral marginal depression very narrow laterally ( |
39. |
12 | Elytra subquadrate, lateral margins extended laterally outside humeral angle, lateral marginal depression moderately broad, edge upraised and translucent near anterior series of lateral elytral setae ( |
40. |
– | Elytra ovoid, lateral margins curved posterad outside humeral angle, lateral marginal depression narrower, edge little upraised and not translucent near anterior series of lateral elytral setae ( |
41. |
13 | Pronotal lateral margins straight or concave before well-developed obtuse hind angles; elytral microsculpture distinct; parascutellar striole completely indicated | 14 |
– | Pronotum lateral margins convex before scarcely indicated, blunt hind angles, the laterobasal seta producing a small projection, the pronotal base narrow, MPW/BPW = 1.39 ( |
42. |
14 | Elytral humerus rounded ( |
15 |
– | Elytral humerus angulate, the elytral margin raised as a distinct ridge ( |
43. |
15 | Pronotal lateral margins straight anterad blunt, obtuse hind angles ( |
16 |
– | Pronotal lateral margins sinuate anterad the distinct hind angles ( |
4. |
16 | Body size larger, standardized body length 5.6–6.4 mm; two dorsal elytral setae in third interval | 17 |
– | Body size smaller, standardized body length 4.3 mm; dorsal elytral setae absent (Marau) | 38. |
17 | Pronotum narrower basally, MPW/BPW = 1.37–1.41, lateral margin convex anterad obtuse-rounded hind angle ( |
45. |
– | Pronotum broader basally, MPW/BPW = 1.32, lateral margin slightly sinuate anterad obtuse hind angle, concavity of margin defined by jag at basal setal articulation ( |
40. |
Of the four species in this group with shallow elytral striae (
Male genitalia. Aedeagal median lobe robust, shaft broad; apex moderately extended beyond ostium, tip gently downturned (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hiti el. 2080 m 2-VI- / 2006 lot 03 pyrethrin fog /
Paratype: Tahiti Nui, Pito Hiti, 2090 m el.,
The species epithet papau is the Tahitian adjective for shallow, as well as the noun for shallows. The name signifies the very shallow elytral striae observed in individuals of this species.
The two specimens were collected in pyrethrin fog samples of moss-covered vegetation 20–30 m elevation below the summit of Pito Hiti.
Like
Female reproductive tract. The female holotype was not dissected, but the gonocoxae are exerted and therefore visible. Basal gonocoxite 1 with apical fringe of 2–3 setae laterally, and 4–5 smaller setae along medial margin including apex; apical gonocoxite 2 with two lateral ensiform setae, the apical lateral seta stouter but of the same length as the basal seta, and a single dorsal ensiform seta as broad as the apical seta; apical sensory furrow bearing two nematiform setae.
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hiti el. 2110 m 2-VI- / 2006 lot 04 pyrethrin fog /
The species epithet is the Tahitian word mānina, meaning smooth, polished, or unwrinkled, signifying the smooth polished elytra observed in the holotype.
This was the sole specimen collected on the actual summit of Pito Hiti, 2110 m elevation. It was found via pyrethrin fogging of moss-covered vegetation. Only 30 m lower in elevation, the same collecting technique resulted in specimens of
Among the
Male genitalia. Aedeagal median lobe shaft moderately slender, the apex beyond the ostium recurved first ventrally then dorsally, the apex expanded dorsoventrally, moreso ventrally, and flattened as an apical face (
Female reproductive tract. Bursa copulatrix curved and folded to the right apically, length about 2× maximal breadth compressed on microslide (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hiti el. 2070 m 2-VI- / 2006 lot 01 pyrethrin fog /
Allotype female (MNHN) labeled as holotype.
Paratypes: same data as holotype (CUIC, 2; EMEC, 3); Tahiti Nui, Pito Hiti, 2000 m el.,
Based on synapomorphies of the body coloration and male genitalia, this species is hypothesized to represent the adelphotaxon of
All specimens have been collected from 2000 to 2080 m elevation on Pito Hiti via pyrethrin fogging of moss-covered vegetation.
Individuals of this species share the pale elytral margins and sutural interval surrounding piceous elytral discs with
This species is known from two male specimens collected at 1900 m elevation near the summit of Mont Aorai.
Beetles of this species (
Male genitalia. Aedeagal median lobe broad in basal half, evenly narrowed to apex of ostium, lobe apex narrowly extended beyond ostium, expanded into a narrowly spatulate tip with blunt dorsal expansion (
Female gonocoxae. The female allotype was not dissected, but examining the distended abdomen (
Holotype male (MNHN) labeled: FRENCH POLYNESIA: Tahiti Iti / Mont Atara, 815 m el. /
Allotype female (MNHN) labeled as holotype // ALLOTYPE / Mecyclothorax / ramagei / J.K. Liebherr 2013 (black-bordered red label).
The species epithet is a patronym honoring the collector, Thibault Ramage, entomological consultant, Concarneau, France.
This species is distributed on the Tahiti Iti volcano. The type series was obtained by sifting mossmats on tree trunks, and extracting the insects from the siftate using a Winkler extractor (T. Ramage pers. comm.).
Among the larger-bodied species of the
This species was previously known from two male specimens
Among species of the
Male genitalia. Aedeagal median lobe shaft slightly broader at midlength, ventral surface slightly convex (
Female reproductive tract. Bursa copulatrix broad with rounded apex, length 2.5× length, cuticle appearing thick based on depth of staining with Chlorazol Black (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2000 m 2-VI- / 2006 lot 02 pyrethrin fog /
Allotype female (MNHN): same data as holotype.
Paratypes: Tahiti Nui; Pito Hiti, 2070 m el.,
This species name is derived from the type locality, Piti Hiti, resulting in an adjectival epithet – pitohitiensis – that is derived in parallel with that of the closely related species,
All specimens representing this species have been collected from 2000–2080 m elevation on Pito Hiti via pyrethrin fogging of moss-covered vegetation.
This species is easily diagnosed by the broadly ellipsoid elytra and small, little transverse pronotum, MPW/PL = 1.25 (n = 2) (
This species is known from 800–1100 m elevations on Mont Teatara, Tahiti Iti. One specimen was collected via pyrethrin fogging of mossy horizontal logs and mossy tree trunks.
Among species of the
The male holotype and only known specimen was collected between 900 and 1100 m elevation on Mont Marau.
This and the similarly sized
This species is known from 1080 m elevation on Mont Mauru, where the single known female specimen was collected by pyrethrin fogging tangles of dead fern fronds.
Among
This species is known to occur from 1200–1900 m elevation on Mont Aorai. All five known specimens were collected in association with moss-covered vegetation, either by finding specimens by hand in moss, or via application of pyrethrin insecticide fog to the vegetation.
This species is extremely similar to the preceding,
This species is known to live on Mont Aorai from 1265 m to 1900 m elevation. One of the four known specimens was collected by beating live and dead fern fronds over a beating sheet, indicating the use of an arboreal refuge during daytime.
Of the five
This species is known from Mont Marau at 1000–1400 m elevations. Associated ecological data are restricted to arboreal situations; in banks of living
Of the
This member of the
This species is known from Mont Marau at elevations between 1000 and 1400 m. All individuals with ecological data have been associated with arboreal microhabitats on a variety of plant substrates:
Among the
The four known specimens of
1 | Discal intervals on elytral base with shallow but distinct transverse microsculpture | 2 |
– | Discal intervals on elytral base glossy, microsculpture not visible (Marau) | 46. |
2 | Pronotal median base smoother mediobasally ( |
3 |
– | Pronotal median base densely punctured ( |
47. |
3 | Elytra with two setae at apex, the apical seta near apex of stria 2, and the subapical more basad in stria 7; pronotal lateral marginal depression narrow in apical half, concolorous with disc ( |
4 |
– | Elytra with a single apical seta near apex of stria 2, subapical seta absent; pronotal lateral marginal depression broad to front angle, rufoflavous, contrasted with rufopiceous pronotal disc ( |
48. |
4 | Elytra broadly ovate, MEW/MPW = 1.59 ( |
49. |
– | Elytra more narrowly ovate, MEW/MPW = 1.46 ( |
50. |
Among members of this group, this species is diagnosable by the very glossy upper body surface, with the vertex, pronotal disc, and discal elytral intervals extremely smooth such that tiny micropunctures – pore canals – are visible across the smooth surface. The pronotal hind angles are obtuse and only slightly protruded, the projection consisting of an expansion of the marginal bead surrounding the articulatory socket of the basal pronotal seta (
This species is known from the type series of five specimens collected at 1000 m elevation on Mont Marau.
Beetles of this species appear much like those
Unique among the
Male genitalia. Aedeagal median lobe only moderately curved, ventral surface of shaft straight apicad parameral articulations (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Iti / Mts. Teatara summit / 1195 m el. 17-IX-2006 lot 10 /
The patronymic species epithet curtisi honors the collector, Dr. Curtis P. Ewing, who collected extensively at the summit of Mont Teatara, one of several inaccessible French Polynesian peaks he has worked hard to explore.
The holotype and only known specimen was collected at 1195 m elevation at the summit of Mont Teatara, Tahiti Iti. The specimen came from a pyrethrin fog sample of a horizontal mossy log with an epiphytic
This species shares with
All that is known about this species’ distribution is the elevation of the type locality for the unique holotype: 1400 m elevation on Mont Marau.
This is the smallest-bodied species in the
The unique female holotype of this species was collected between 1000 and 1800 m elevation on Mont Aorai.
1 | Discal elytral striae lined with rounded punctures that laterally expand striae; elytral sutural interval and lateral two intervals contrastedly paler than brunneous intervals 2–6 ( |
51. |
– | Discal elytral striae indistinctly punctured, the punctures elongate, not laterally expanding striae; elytra distinctly paler laterally, but only the lateral marginal depression contrastedly paler than disc ( |
52. |
The pale rufoflavous head and pronotum, and pale-margined elytra with contrasting dark brunneous intervals 1–6 serve to diagnose this species at first glance (
Male aedeagal median lobe and associated parameres,
The known distribution of this species spans 1750–1900 m elevation on Mont Aorai.
This species can be diagnosed from
Male genitalia. Aedeagal median lobe bowed dorsally with straight ventral margin apicad parameral articulations (
Female reproductive tract. Bursa copulatrix narrowly extended from broader vagina, the bursal extension twice as long as broad (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2000 m 2-VI- / 2006 lot 02 pyrethrin fog /
Allotype female (MNHN) labeled as holotype.
Paratypes: labeled as holotype (EMEC, 1); Tahiti Nui, Pito Hiti, 2070 m el.,
The species epithet compounds the Tahitian word for same, hō’ēā with the word for edge, hiti, signifying the similar coloration of pronotal and elytral margins in beetles of this species;
This species is known from pyrethrin fogging of moss-covered vegetation from 2000–2080 m elevation on Pito Hiti.
1 | Elytral microsculpture absent, surface glossy, coloration dark rufobrunneous to rufopiceous with aeneous reflection; elytral striae shallow and smooth, intervals slightly convex; pronotal lateral margin very narrow, edge beaded near lateral setae | 2 |
– | Elytral microsculpture evident, more or less transverse, coloration rufobrunneous without distinct metallic reflection; elytral intervals moderately convex, striae deep, punctate basally; pronotal margin narrow, edge upturned | 3 |
2 | Body dark, glossy piceous ( |
53. |
– | Body paler, rufopiceous ( |
54. |
3 | Elytral striae 2–6 continuously and uniformly impressed from disc to elytral basal groove ( |
4 |
– | Elytral striae 2–6 much deeper on disc than near elytral basal groove where striae may be very shallow and traceable or absent ( |
6 |
4 | Larger beetles, standardized body length 3.8–5.7 mm; pronotal and elytral margins narrowly paler, rufobrunneous to rufoflavous versus – depending on degree of melanization – correspondingly rufopiceous to rufobrunneous discs; one dorsal elytral seta in third interval | 5 |
– | Small beetles, standardized body length 3.3–3.4 mm; pronotal and elytral margins and suture broadly flavous, contrasted with rufobrunneous pronotal and elytral discs ( |
55. |
5 | Pronotal median base unformly depressed relative to disc, margined anteriorly at disc by dense longitudinal strigae ( |
56. |
– | Pronotal median base sloping anteriorly to meet disc, anterior margin nearly coplanar with disc and as punctured as remainder of median base ( |
57. |
6 | Third elytral interval with two dorsal setae, the setal articulatory sockets in evident, depressed punctures ( |
7 |
– | Third elytral interval glabrous, dorsal elytral setae absent ( |
58. |
7 | Elytral microsculpture consisting of dense transverse lines with little tendency to form a mesh | 8 |
– | Elytral microsculpture consisting of transverse mesh in regular transverse rows, crossconnections between transverse sculpticell margins obvious | 9 |
8 | Pronotal lateral margin distinctly sinuate before hind angle, laterobasal depression deep; pronotal median base covered with irregularly distributed, deep and distinct punctures; discal elytral striae convex, striae 1–7 distinctly punctate in basal half of length ( |
59. |
– | Pronotal lateral margin slightly sinuate before hind angle, laterobasal depression shallow; pronotal median base bearing 2–3 transverse rows of small, shallow, sparsely distributed punctures; discal elytral striae broadly, slightly convex, striae 1–5 minutely punctate in basal half of length ( |
60. |
9 | Pronotal hind angles protruded, right, pronotal lateral margin distinctly, angularly convergent immediately anterad hind angle; elytral striae distinctly punctate on disc, discal intervals convex ( |
61. |
– | Pronotal hind angles obtuse, denticulate, pronotal lateral margin subparallel for very short distance outside basal seta articulatory socket; elytral striae moderately shallow, punctate, but intervals only slightly convex ( |
62. |
Among the five species in the
Male genitalia. Aedeagal median lobe curved dorsally, apex broadened dorsoventrally with broad, flat apical face (
Female reproductive tract. Bursa copulatrix elongate, symmetrical and only slightly narrower than vagina in unstretched configuration (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2000 m 2-VI- / 2006 lot 02 pyrethrin fog /
Allotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2070 m 2-VI- / 2006 lot 01 pyrethrin fog /
Paratypes: 3 paratypes labeled as the holotype (CUIC, 2; EMEC, 1).
The Tahitian word nīnamu means blue as in the blue color of a lagoon, and signifies the metallic dorsal coloration of these beetles.
This species inhabits the summit area of Pito Hiti from 2000–2070 m elevation, with specimens collected from moss-covered vegetation through the application of pyrethrin fog.
Within the
This species is known only from near the summit of Mont Aorai, with all specimens collected between 1750 and 1900 m elevation.
This smallest-bodied Tahitian
Female reproductive tract. Bursa copulatrix relatively elongate, vagina of similar diameter and therefore not discernible (
Female reproductive tract dissections,
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2000 m 2-VI- / 2006 lot 02 pyrethrin fog /
Paratype: female specimen, reproductive tract dissected and left elytron removed (CUIC, 1).
The species epithet kokone is the Tahitian word for diminutive, and it signifies the very small body size of these beetles. Kokone is one of only a few Tahitian words (
This is another species known only from 2070 m elevation near the summit of Pito Hiti. As for the others, beetles of this species have been shown to inhabit arboreal moss on emergent vegetation through sampling using pyrethrin fog.
Among the smaller-bodied members of the
In 1977,
Among the
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2070 m 2-VI- / 2006 lot 01 pyrethrin fog /
The species epithet paahonu is taken from the Tahitian word for maroon, a rough approximation of the dorsal body coloration of the holotype.
Known from a pyrethrin fog sample of moss-covered vegetation taken 40 m elevation below the summit of Pito Hiti.
Of the
Male genitalia. Aedeagal median lobe distinctly dorsally curved, apex very short, the tip subacuminate (
Female reproductive tract. Bursa copulatrix and vagina separated by a distinct constriction (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1125 m / 4-IX-2006 lot 01 /
Paratypes: same data as holotype (CUIC, 2). The paratypes have been dissected for male genitalic and female reproductive tract description.
This species is named in honor of Dr. Kathleen Ball for her support for and active engagement in the carabidological community. It should be noted that
This species is known from the type series collected at 1125 m elevation on Mont Marau. The beetles were beaten from dead fern fronds that were associated with living plants growing on a wet rock face.
Among the
All that is known about the biology and distribution of this species is restricted to the 1976 collection of the eight-member type series from 1300–1400 m elevation on Mont Marau (
This species shares with
This species is so far known to be associated with low-stature
Among the
This species is known from the higher reaches of Mont Aorai, with the male holotype and female allotype collected near Fare Ata, 1900 m elevation.
This species shares with
Male genitalia. Aedeagal median lobe shaft broad, apex moderately extended beyond ostium and broad dorsoventrally (
Female reproductive tract. Bursa copulatrix ovately expanded apicad a slightly narrower vagina; bursal plus vaginal length twice as long as maximal breadth when compressed under cover slip (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hito el. 2070 m 2-VI- / 2006 lot 01 pyrethrin fog /
Allotype female (MNHN) labeled as holotype.
Paratypes: labeled as holotype (CUIC, 1; EMEC, 1); Pito Hiti, 2000 m el.,
The species epithet ehu is taken from the Tahitian word 'ehu, meaning the color red. The name signifies the reddish coloration of beetles of this species.
This species is known from 2000-2070 m elevation on Pito Hiti, and based on collections made using pyrethrin fog, spends at least part of its life within arboreal mossmats.
Male aedeagal median lobe and associated parameres,
1 | Elytral third interval with two dorsal setae, setal formula 2122 or 2121 | 2 |
– | Elytral third interval with one dorsal seta before midlength ( |
63. |
2 | Pronotal lateral margins distinctly subparallel or convergent anterad projected hind angles which are right to acute; body narrower, elytral basal margins sloping posteriorly immediately laterad humeral angle | 3 |
– | Pronotal lateral margins subparallel for very short distance anterad little projected hind angles which are slightly obtuse; body broad, elytral basal margins extended laterally outside humeral angle before anterior series of lateral elytral setae ( |
64. |
3 | Elytral lateral marginal depressions very broad, explanate and translucent from humeral angle posterad past anterior series of lateral elytral setae ( |
4 |
– | Elytral lateral marginal depression narrower just laterad humeral angle than posterad outside anterior series of lateral elytral setae ( |
65. |
4 | Eighth elytral interval broadly convex from middle of posterior series of lateral elytral setae to elytral apex, convexly raised laterally outside stria 7, never carinate; elytral lateral marginal depression of even breadth near bases of elytral striae 5–7 laterad humerus; elytra broadly ovate ( |
66. |
– | Eighth elytral interval raised above stria 7 from lateral elytral setae to elytral apex, the interval either flat laterally and sharply carinate, or more broadly convex and dorsally subcarinate; elytra narrowly ovate ( |
67. |
Beside the presence of only the anterior elytral seta, and therefore setal formula 2111, this species is set off from the other members of the
The unique holotype specimen was collected by Elwood Zimmerman between 1675 and 1900 m elevation on Mont Aorai. The specimen was beaten from moss on either a tree or a shrub.
This species shares the expanded pronotal and elytral lateral margins of other members of the
This Tahiti Iti representative of the species group is known from specimens collected 900–1100 m elevation on Mont Teatara. A teneral female specimen was obtained by beating
Among the five species currently known to comprise this group, this one has the least transverse pronotum (
Male genitalia. Aedeagal median lobe shaft narrow and of even diameter from parameral articulations to apex of ostium, barely narrower apically with an evenly proportioned dorsoventrally spatulate apex (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1125 m / 4-IX-2006 lot 01 /
The species epithet pahuhiti compounds the Tahitian words pāpū, meaning even, with hiti, meaning border or edge, the species name signifying the evenly explanate pronotal lateral margins.
The single known specimen was collected in a beating sample along with one specimen of
Most similar to
This species is known from lower elevations north of Mapura to Mapura summit, on the ridge that leads upward to Pihaaiateta and Pito Hiti. Specimens were collected during 1978 between 900 and 1200 m elevations by beating ferns (
From the most similarly appearing
This species is known from both Mont Marau and Mont Aorai; from 1125–1400 m elevation on the former, and 1210–1400 m on the latter. Various male specimens were dissected to verify conspecificity of populations on the two ridge systems, with observed variation precluding division into two species. There is variability in the dorsoventral expansion of the median lobe apex, with two specimens from Marau exhibiting a narrow apical extension and narrowly spatulate apex (
Specimens have been collected from the rotten cambial layer under the bark of a dead
This species is characterized by a narrow pronotum, MPW/PL = 1.14, pronotal lateral margins parallel for 0.10× pronotal length anterad the slightly obtuse hind angles, and the broadly based elytra, the humeri extended laterally, and the lateral elytral margins distinctly curved posteriorly outside the humeral angles (
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Pito Hiti el. 2070 m 2-VI- / 2006 lot 01 pyrethrin fog /
The Tahitian word tuea – square, level, fit together – well characterizes the tight fit of the prothorax to the elytra observed in this species.
The single specimen was collected in a pyrethrin fog sample of moss-covered vegetation at 2070 m elevation on Pito Hiti.
1 | Pronotal lateral marginal depression evident, of equal breadth throughout length and not narrower at lateral seta, edge upturned; sutural stria 1 and stria 2 basally fused for 0.1× elytral length ( |
2 |
– | Pronotal lateral marginal depression obsolete, narrowest at position of lateral seta, wider at front angle, margin beaded or very narrowly upturned; sutural stria and stria 2 free at elytral base, though they may approach, or stria 2 may be obsolete | 3 |
2 | Dorsal microsculpture evident, pronotal disc with indistinct transverse mesh and transverse lines visible outside areas of reflected light, elytral intervals covered with distinct transverse microsculpture, the sculpticells ranging from twice as broad as long, to very elongate and so defining transverse lines (Marau) | 69. |
– | Dorsal microsculpture obsolete, pronotal disc glossy with only indistinct transverse-line microsculpture outside areas of reflected light, elytral intervals glossy, at most indistinct sculpticells occurring in depressed areas within striae (Aorai) | 70. |
3 | One supraorbital seta each side | 4 |
– | Two supraorbital setae each side | 12 |
4 | Elytral microsculpture indistinct or absent, integument glossy | 5 |
– | Elytral microsculpture distinct, transverse mesh to transverse lines | 7 |
5 | One or two dorsal elytral setae present in the basal half of third interval | 6 |
– | Third elytral interval glabrous on disc, dorsal elytral setae absent (Aorai) | 71. |
6 | One dorsal elytral seta present in the basal half of third interval (Teatara) | 72. |
– | Two dorsal elytral setae present in third interval, on near basal ⅓, the second beyond midlength (Teatara) | 73. |
7 | Elytral third interval with one dorsal seta before midlength or glabrous, no dorsal setae present (in rare instances of the common |
8 |
– | Elytral third intervals with two dorsal elytral seta bilaterally present | 74. M. vaifaufa Perrault |
8 | Dorsal elytral seta absent, third elytral interval glabrous | 9 |
– | One dorsal elytral seta present in third interval before midlength | 10 |
9 | Pronotal median base minutely punctate across width, surface glossy among the punctures, laterobasal depressions deep, margined laterally and basally by marginal bead, no median tubercle present ( |
75. |
– | Pronotal median base irregularly, longitudinally strigose along juncture with disc, surface with granulate isodiametric microsculpture among the large punctures and wrinkles, laterobasal depressions shallow with median tubercle ( |
76. |
10 | Elytral microsculpture consisting of elongate, more or less regularly aligned transverse mesh, elytral surface glossy but not iridescent; legs flavous, antennae rufoflavous to brunneous apically | 11 |
– | Elytral microsculpture consisting of fine transverse lines with little tendency to form a mesh, elytral surface subiridescent; legs rufoflavous, antennae darker apically, antennomeres 4–11 dark brunneous to piceous (Teatara) | 77. |
11 | Elytral striae shallow, deepest portions with elongate punctures that do not expand discal strial breadth, the punctures producing an undulated surface along the depth of the stria, elytral intervals broadly, slightly convex ( |
91. |
– | Elytral striae well developed, with rounded punctures that expand discal striae in basal half of elytral length, elytral intervals moderately convex ( |
78. |
12 | Discal elytral striae, especially striae 3–5, very shallow, interrupted or scarcely indicated between some of the strial punctures | 13 |
– | Discal elytral discal striae continuous, shallow to deeply incised | 15 |
13 | Two or one dorsal elytral setae present, setal formula 2121 or 2111 | 14 |
– | Third elytral interval glabous, dorsal elytral seta absent, setal formula 2101 ( |
79. |
14 | Third elytral interval with two dorsal elytral setae, one near basal ¼ and the second just posterad elytral midlength ( |
80. |
– | Third elytral interval with one dorsal elytral seta at ~ 0.30× elytral length ( |
81. |
15 | Third elytral interval glabrous on disc, dorsal elytral setae absent | 16 |
– | One or two dorsal elytral setae present in third interval | 18 |
16 | Elytra ovoid, body robust, MEW/MHW = 2.25–2.26 ( |
17 |
– | Elytra narrowly ovoid, body gracile, MEW/MHW = 2.03 ( |
82. |
17 | Discal elytral striae irregular, wavering, elongate punctulae in depths of striae, striae 2–5 shallow to absent basally, not deeply joined to basal groove, sutural stria 1 and stria 2 free basally ( |
83. |
– | Discal elytral striae smooth, deep, striae 1–5 deep and continuous basally, joined to basal groove, sutural stria 1 and stria 2 fused basally ( |
84. |
18 | Elytral microsculpture indistinct, discal intervals entirely glossy or indistinct sculpticells visible only in small patches | 19 |
– | Elytra with distinct microsculpture, either a transverse mesh or transverse lines that cross entire breadth of elytral intervals | 21 |
19 | Eyes little convex, posterior margin of ocular lobe meeting gena at > 135° angle, ocular ratio 1.31–1.41 ( |
20 |
– | Eyes convex, posterior margin of ocular lobe distinctly projected, meeting gena at < 135° angle, ocular ratio 1.47–1.50 ( |
85. |
20 | Eyes very flat, frons relatively broad, ocular ratio = 1.31; pronotum densely, longitudinally strigose anterad transverse impression, median base distinctly depressed relative to disc, the surface shagreened by isodiametric sculpticells among deep, distinct punctures (Marau) | 86. |
– | Eyes slightly projected, ocular ratio = 1.41; pronotal anteromedial surface glossy, anterior transverse impression obsolete, median base sparsely lined with small punctures, the surface among punctures glossy (Teatara) | 87. |
21 | One dorsal elytral seta present in the basal half of third elytral interval (or in rare instances of the abundant |
22 |
– | Two dorsal elytral setae bilaterally present in third interval | 26 |
22 | Elytra with distinct microsculpture consisting of dense transverse lines without formation of a mesh | 23 |
– | Elytra with evident transverse microsculpture arranged in a mesh that is more or less transverse | 25 |
23 | Pronotal lateral margins anterad hind angles convergent to the level of the anterolateral portion of the median base, then divergent, hind angles projected ( |
24 |
– | Pronotal lateral margins convergent for only a very short distance anterad hind angles, divergent for much of distance laterad median base, hind angles denticulate ( |
88. |
24 | Elytra subquadrate, broad basally, MEW/HuW = 1.87; elytral striae 5 and 6 obsolete basally, the striae not joined to elytral basal groove at humeral angle ( |
89. |
–. | Elytra subovate, humeri rounded, MEW/HuW = 2.0; elytral striae 5 and 6 continuous basally though shallow, joined to elytral basal groove at humeral angle ( |
90. |
25 | Male aedeagal median lobe short and broad ( |
91. |
– | Male aedeagal median lobe long and narrow ( |
92. |
26 | Vertex of head and pronotal disc glossy, lacking distinct, raised, mesh-like microsculpture ( |
27 |
– | Vertex of head and pronotal disc covered with raised isodiametric to transverse sculpticells, the surface with cupreous reflection ( |
93. |
27 | Elytral disc with well-developed transverse microsculpture | 28 |
– | Elytral disc glossy, microsculpture indistinct | 32 |
28 | Discal elytral striae deep, smooth to finely punctate, the punctures not expanding strial breadth; elytra dark with aeneous metallic reflection or pallid, brunneous | 29 |
– | Discal elytral striae distinctly punctate basally, punctures expanding strial breadth; elytra rufopiceous (Marau) | 94. |
29 | Pronotal hind angles distinct, right to acute, pronotal lateral margin distinctly sinuate anterad hind angle ( |
30 |
– | Pronotal hind angles obtuse-rounded, little projected, pronotal lateral margin moderately sinuate anterad hind angle ( |
95. |
30 | Pronotum more transverse, MPW/PL = 1.25–1.26 ( |
31 |
– | Pronotum less transverse, MPW/PL = 1.16–1.20 ( |
96. |
31 | Elytral striae 3–5 obsolete on elytral base, elytral humeral region smooth ( |
97. |
– | Elytral striae 3–5 continuous and traceable to juncture with elytral basal groove ( |
98. |
32 | Elytra narrowly ovoid, not so broad relative to pronotum, MEW/MPW = 1.52–1.62 ( |
33 |
– | Elytra broad, orbicular, pronotum narrow, MEW/MPW = 1.65–1.76 ( |
99. |
33 | Pronotal base broader, MPW/BPW = 1.50–1.53; lateral elytral striae 6–7 as deep or deeper than discal striae 1–5 | 34 |
– | Pronotal base narrower, MPW/BPW = 1.56–1.58; discal elytral striae moderately deep, lateral stria 6–7 much shallower, 7th indistinct (Teatara) | 100. |
34 | Dorsal body surface glossy, rufobrunneous, but without bronzed metallic reflection; elytra symmetrically ellipsoid, lateral margins nearly parallel at midlength ( |
101. |
– | Dorsal body surface glossy, rufobrunneous, but with bluish metallic reflection on pronotal disc; elytra slightly obovate, lateral margins more narrowly rounded near humeri, margins convex at midlength ( |
72. |
Male aedeagal median lobe and associated parameres,
Male aedeagal median lobe and associated parameres,
Female reproductive tract dissections,
Male aedeagal median lobe and associated parameres,
This species and
Male genitalia. In keeping with the isolated external anatomy of this species within Tahitian
Female reproductive tract. Bursa copulatrix broad, slightly expanded apically, length 1.6× maximal breadth compressed under microslide cover slip (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1165 m / 4-IX-2006 lot 03 /
Allotype female (MNHN) labeled as the holotype.
Paratypes: labeled as the holotype (CUIC, 6; EMEC, 1; NMNH, 1).
The species epithet compounds the Tahitian words tā’ati, meaning unite or join together, and tore, which means, among other things, streaked, or military stripes. The epithet signifies the fused bases of elytral striae 1 + 2.
These beetles were discovered by pyrethrin fogging a wet rock face above a small stream. The rock face lay on the sidewall of a small gulch that was shaded by trees, and was about 6 m in height and about 10 m across, the fractured face lying at a 75° angle to the horizontal. The surface was covered with mosses and liverworts, and the beetles crawled out of the cracks due to the action of the pyrethrin. One adult and four larvae of
Sharing the basally fused elytral striae 1 + 2 and narrow pronotum with
This species’ known distribution is Mont Aorai from 1000–1750 m elevation. Two of the three known specimens were collected in a vinegar pitfall trap, demonstrating that this species shares geophilic habits with its sister,
This is the smallest-bodied species in the
Holotype female (MNHN) labeled: Aorai - Tahiti / piege 1100/1400 m / 20/21.V.1978 (pink paper label // Museum Paris / 1994 / Coll. G. Perrault (pink paper) // ε (hand written on white label) // HOLOTYPE / Mecyclothorax / konemata / J.K. Liebherr 2013 (black-bordered red label).
The species epithet compounds the Tahitian word kone, meaning small, with mata, meaning eye; konemata signifying the small eyes characterizing this species.
This specimen was collected by Georges Perrault on Mont Aorai, 1000–1400 m elevation in 1978, and placed a the end of his collection in a small section of undescribed material. The epsilon – i.e., “ε” – identifier label suggests that he understood this specimen to represent something distinctive, but he did not have the opportunity to describe it.
Among the eight
This species is known from Mont Teatara, Tahiti Iti, from 900–1100 m elevation. One specimen was obtained through the application of pyrethrin fog to a horizontal mossy log
This species is distinguished by the narrowly ovoid elytra with very deep elytral striae, the lateral striae 7 and 8 actually more deeply incised than the discal striae. All striae are deeply impressed to the elytral basal groove, and striae 1 + 2 are basally fused from the basal groove to a level halfway along the parascutellar striole (
This species is known from the type series of three specimens collected on Mont Teatara, 900–1100 m elevation, plus a fourth specimen collected in a 2006 pyrethrin fog sample of a moss-covered
This and
The unique type specimen was collected at 1000 m elevation at a location labeled Parc Vaifaufa, Tahiti Iti (
These beetles are the largest in the
Male genitalia. Aedeagal median lobe gracile, shaft moderately narrow and slightly curved (
Female reproductive tract. Bursa copulatrix basally as broad as vagina, apically narrowed and parallel sided when compressed under microslide cover slip (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1275 m / 10-IX-2006 lot 02 /
Allotype female (MNHN) labeled as holotype.
Paratypes: labeled as the holotype (CUIC, 2); Mt. Marau, 1280 m el., 6-xi-1999, Polhemus, pyr. fog
The species epithet arboricola, tree loving, is of parallel derivation to that of the anatomically similar species
This species is known to occupy
Of the three Tahitian species in the
This species is the geographically most widespread Tahitian
Among the
Individuals of this species have been found from 900–1200 m elevation on Mont Teatara, Tahiti Iti. Beetles have been found inhabiting arboreal mosses, and among the appressed leaves of the clumped
Along with the previous species,
The holotype and paratype were collected between 900–1100 m elevation on Mont Teatara, Tahiti Iti.
Among the Tahitian
Male genitalia. Aedeagal median lobe very small, about half the length relative to adult body length when compared to aedeagi of species with similarly sized males (
Holotype male (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau summit el. 1445 m / 4-IX-2006 lot 04 /
The species epithet compounds the Tahitian words rahi, or big, and mata, or face, the name signifying the disproportionately large head in beetles of this species.
The single specimen was found by beating vegetation on the summit ridge of Mont Marau, 1445 m elevation. The vegetation included low stature, 3–4 m tall
This is one of only two
This species is known to inhabit Mont Mauru from 960–1110 m elevation. It is associated with arboreal microhabitats based on its discovery through beating vegetation, including
The second of the
This species inhabits Mont Marau from 900–1400 m elevation. It has been found by in dead
Of the four Tahitian
Holotype female (MNHN) labeled: French Polynesia: Tahiti Iti / Mts. Teatara NW ridge / 1146 m el. 19-IX-2006 lot 09 /
The species epithet is the Tahitian word oaoa, meaning narrow, the name signifying the narrowly ovate elytra characterizing this species.
The type specimen was found on Mont Teatara, Tahiti Iti in
This is the broadest-bodied species (
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1315 m / 10-IX-2006 lot 05 /
The species epithet compounds the Tahitian words mānina, or smooth, with popoti – i.e., roach or cockroach – signifying the smooth elytral striae of this beetle.
This is a species of Mont Marau, known from 1315 m elevation. The lone specimen was included in a pyrethrin fog sample from moss-covered
Quite similar in appearance to
Holotype female (MNHN) labeled: French Polynesia: Tahiti Nui / Mt. Marau road el. 1185 m / 4-IX-2006 lot 02 /
The species epithet compounds the Tahitian words huna, meaning secret or hidden, with popoti, cockroach, signifying the similarity of this species to
This species is known only from a pyrethrin fog sample taken at 1185 m on Mont Marau. The vegetation fogged was unremarkable, consisting of a
Of the eight Tahitian species in the
Male genitalia. Aedeagal median lobe robust, shaft broad with expanded ventral margin at ostium (
Female reproductive tract. Bursa copulatrix columnar, length nearly 3× maximal breadth when pressed under microslide cover slip (
Holotype male (MNHN) labeled: SOCIETY IS: Tahiti / Tahiti Nui Mont Marau / 1280 m el. 6-XI-1999 / D.A. Polhemus pyr. fog / sta. 1
Allotype female (MNHN) labeled as holotype.
Paratypes labeled as holotype (CUIC, 2; NMNH, 1).
The species epithet compounds the Tahitian words fefe, or curved, and mata, meaning eyes or face. The name refers to the convex eyes that differentiate this species from the one below,
This species is known from 1280 m elevation on Mont Marau. The type series comprised a portion of pyrethrin fog samples from moss-covered
Among species of the
Female reproductive tract. Bursa copulatrix elongate, narrowly rounded apically, length 2.6× width when compressed under microslide cover slip (
Holotype female (MNHN) labeled: Mt. Marau, Tahiti / 1200–1400 m / XII-1977 / J. Gourvès (on reverse) (pink paper) // Museum Paris / 1994 / Coll. G. Perrault (pink paper) // Ƶ (white paper) // HOLOTYPE / Mecyclothorax maninamata / J.K. Liebherr 2013 (black-bordered red label).
Etymology. The species epithet compounds the Tahitian words manina, in this usage meaning flat, and mata, herein meaning eye. The name therefore refers to the flat eyes diagnostic of this species.
The holotype specimen was placed at the end of the Georges Perrault collection, with the tertiary label “Ƶ” indicating his recognition of it as distinct, though undescribed. It was collected between 1200 and 1400 m elevation on Mont Marau, though the situation within which Professeur Gourvès collected it was not recorded.
Along with abundant
This is the most commonly encountered species on Mont Teatara, Tahiti Iti.
This comprises one of the three species within the
Holotype female (MNHN) labeled: French Polynesia: Tahiti Iti / Mts. Teatara NW ridge 1080– / 1100 m el. 17-IX-2006 lot 07 /
The species epithet niho is the Tahitian word for tooth, the name referring to the denticulate pronotal hind angles.
This species has been recorded from near 1100 m elevation, Mont Teatara, Tahiti Iti. The pyrethrin fog sample from a moss-covered
Among the three
This species is known from 1080–1100 m elevation on Mont Mauru. The specimen was collected from dense tangles of dead fern fronds (
Very much like the preceding two species,
The two type and only known specimens of this species were collected near the summit of Mont Aorai at 1900m elevation.
Among the
Variation. Limited setal variation occurs among individuals of this abundant, geographically widespread species. A large series of 77 individuals – Mt. Marau, 1335 m el., 15-ix-2006 lot 01 (CUIC) – includes 2 individuals with just the posterior supraorbital seta, 1 with two supraorbital setae on the right side versus one on the left, and 74 with the usual 2111setal formula.
This species is broadly distributed on Monts Marau and Aorai, with recorded localities from 1100–1380 m elevation on the former, and 1067–1900 m elevation on the latter. Based on pyrethrin fog samples, these beetles occupy moss-covered
This species (
The aedeagal configuration reported here (
This species is known from the Pito Hiti massif from 1000–1200 m elevation above the
Instantly recognizable among
This species is distributed from 1200–1900 m elevation on Mont Aorai. Most of the specimens have been collected in pitfall traps indicating activity in the litter or near the soil surface.
Among
This species is recorded from 900–1380 m elevation on Mont Marau. It has been collected using pyrethrin fog in association with ferns growing along the slopes of a small gulch.
This
This species is recorded from 880–1110 m elevation on Mont Mauru. Most of the specimens have been collected in association with ferns, either by beating or through application of pyrethrin fog. Only 2 of the 22 known specimens were collected by beating flowering plants; a mixed beating sample of
Among
This species is recorded from 900–1200 m elevation on the lower reaches of the western ridge leading to Pito Hiti; above
Beetles of this species are most similar to those of
The single adult specimen was collected in moss on a low rockface along the Faatautia River as the river exits from a large, uncollapsed lava tube at 705 m elevation on Mont Mauru (see fig. 2A,
Sharing the basally constricted, moderately transverse pronotum with the prevous two species, though the pronotum is more constricted basally in this species; MPW/BPW = 1.58–1.70 (n = 2). The eyes are much more convex in this species (
Along with
This species is known from over 100 specimens – mostly collected in pitfall traps – at localities ranging 800–1400 m elevation on Mont Marau.
Beetles of this species bear superficial similarity to
This species has been recorded from 900–1100 m elevation on Mont Teatara, Tahiti Iti. All specimens with ecological data have been associated with moss-covered trunks or horizontal logs, with collections made by beating small trunks or through application of pyrethrin fog to downed logs.
As one of the dorsally glossy members of the
This species is known from three specimens collected 1000–1400 m elevation on Mont Aorai, one collected in a vinegar pitfall trap (
The Tahitian
The number of specimens collected, known species diversity, and percent endemism for
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Tahiti Nui/Marau | 947 | 7 | 31(28) | 90% |
Tahiti Nui/Aorai | 461 | 9 | 30(25) | 83% |
Tahiti Nui/Pito Hiti | 157 | 9 | 21(18) | 86% |
Tahiti Nui/Mauru | 40 | 4 | 7(7) | 100% |
Tahiti Iti | 387 | 5 | 17(17) | 100% |
Moorea/Tohiea | 90 | 4 | 7(7) | 100% |
Chorographic depiction of closely related representative species from areas of endemism within Tahiti and Moorea; modified from
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The two massifs with the greatest elevational disparity – Aorai and Pito Hiti – also exhibit the most disparate fauna, with members of 9 of the 10 species groups found on each mountain. Though the species groups are taxonomic constucts based on key characters, and therefore unlikely to represent monophyletic groups in all cases, the various groups represent a diversity of body forms. Thus a broader representation of species groups is likely to signify broader representation of heretofore unidentified clades across the Tahitian
Comparisons of island or lake size, species diversity, percent endemism, and areal extent of species for several diverse and geographically restricted animal radiations. The criterion “km2/species” represents the average range size per species within a restricted area (island or lake) assuming all species are completely allopatric (i.e., parapatric). For Maui
|
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|
|
|
---|---|---|---|---|---|
Tahiti |
|
1,045 | 101 | 100 | 10.3 |
Tahiti Nui |
|
788 | 84 | 100 | 9.4 |
Tahiti Iti |
|
257 | 17 | 100 | 15.1 |
Moorea |
|
134 | 7 | 100 | 19.1 |
Hawaiian Islands |
|
14,899 |
239 |
100 | 62.3 |
Maui |
|
1,887 | 142 |
100 | 13.3 |
Haleakala |
|
1,440 | 115 |
100 | 12.5 |
West Maui |
|
447 | 27 |
100 | 16.6 |
Hawaiian Islands |
|
16,329 |
415 |
100 | 39.3 |
Maui |
|
1,887 | 123 |
53 | 15.3(29.0) |
Haleakala |
|
1,440 | 103 |
36 | 14.0(38.9) |
West Maui |
|
447 | 58 |
21 | 7.7(37.2) |
Hawaii Island |
|
10,430 | 6 |
100 | 1738.3 |
Hawaii Island | Cave |
10,430 | 7 |
100 | 1490.0 |
Lake Tanganyika |
|
32,900 | 250 |
95-99 | 131.6 |
Lake Victoria |
|
68,800 | ~700 |
95-99 | 98.3 |
Lake Malawi |
|
29,600 | ~700 |
95-99 | 42.3 |
† Summed areas of islands occupied by taxon.
References:
1
2
3
4
5
6
One would predict that populations of widespread species occupying several massifs should be found in proximate elevational zones of habitat on those neighboring massifs. Based on recorded localities, this is so. The very large-bodied
All other species occupying these three mountains are precinctive to only one, demonstrating that the narrow ridges and cols have not maintained sufficient dispersal avenues to preclude geographic isolation and subsequent speciation. Such isolation has occurred even though the beetles may currently be found in the same elevational zones as the widespread species. For example, the members of the species triplet
A slightly more complicated pattern affirms the existence of ecological factors beyond mere landscape physiognomy as critical to speciation. The four species exhibiting lamellate male parameres –
Numerous sets of representative species were proposed by
Analogous additions of recently discovered species to complexes hypothesized by Perrault include the addition of Moorea’s
Sympatric most-similar species are also recorded in this radiation, most often from Mont Teatara, Tahiti Iti (
The
The Hawaiian Islands support the second great radiation within the genus
Comparisons of diversity, ages of origin, and resultant calculated net diversification interval (N.D.I.) (
|
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|
|
---|---|---|---|---|
Society Islands |
|
108 | 1.4 |
0.30 |
Hawaiian Islands |
|
239 | 1.8 |
0.33 |
Hawaiian Islands |
|
415 | 5.33, 26–30 |
0.88, 4.3–5.0 |
Hawaii Island |
|
6 | 0.78 |
0.44 |
Hawaii Island | Cave |
7 | 0.13 |
0.07, 0.40 |
Lake Tanganyika |
|
250 | 9.0–12.0 |
1.6–2.2 |
Lake Victoria |
|
~700 | 0.19–0.27, 5.0 |
0.03–0.04, 0.76 |
Lake Malawi |
|
~700 | 2.4–4.6 |
0.37–0.70 |
References:
1 See
2
3
4
5
6
7
Among Pacific insects, the Hawaiian
Worldwide, the adaptive radiations of cichlid fishes in the East African Rift lakes serve as textbook examples of adaptive radiation (
And so we return to the question posed at the beginning of this article; why so many species of
Sexual selection has been invoked as a factor facilitating speciation in Hawaiian
The generalized predatory life cycle of
Nearly all of the taxa newly described above were collected during a biotic survey of Tahiti and Moorea conducted June–September 2006. Many of the new species described herein were collected by Dr. Elin Claridge on or near the summit of Pito Hiti, and I thank her for providing this material for description. Biotic survey activities were supported by N.S.F. award DEB–0451971, Survey of terrestrial arthropods of French Polynesia; Prof. R.G. Gillespie principal investigator. The Délégation à la Recherche de Polynésie française, Dr. Priscille Frogier directeur, provided permission to collect. Dr. Jean-Yves Meyer provided much appreciated logistical support and advice. The Richard B. Gump South Pacific Research Station provided vehicles, and research and living space during the expedition. I thank Drs. C.P. Ewing and D.A. Polhemus for their part in Bugstrafe 2006, and for cooperatively sharing their taxonomic discoveries. I also thank Mons. Thibault Ramage for providing the specimens described as