Collembola of Rapa Nui (Easter Island) with description of five endemic cave-restricted species

Abstract

Eight species of Collembola are reported from recent collections made in caves on the Polynesian island of Rapa Nui (Eas-ter Island). Five of these species are new to science and apparently endemic to the island: Coecobrya aitorererere n. sp., Cyphoderus manuneru n. sp., Entomobrya manuhoko n. sp., Pseudosinella hahoteana n. sp. and Seira manukio n. sp. The Hawaiian species Lepidocyrtus olena Christiansen & Bellinger and the cosmopolitan species Folsomia candida Wil-lem also were collected from one or more caves. Coecobrya kennethi Jordana & Baquero, recently described from Rapa Nui and identified as endemic, was collected in sympatric association with C. aitorererere n.sp. With the exception of F. candida, all species are endemic to Rapa Nui or greater Polynesia and appear to be restricted to the cave environment on Rapa Nui. A key is provided to separate Collembola species reported from Rapa Nui. We provide recommendations to aid in the conservation and management of these new Collembola, as well as the other presumed cave-restricted arthropods.

Full text

Accepted by P. Greenslade: 24 Mar. 2015; published: 24 Apr. 2015
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Collembola of Rapa Nui (Easter Island) with descriptions of five endemic
cave-restricted species
ERNEST C. BERNARD
1,4
, FELIPE N. SOTO-ADAMES
2
& J. JUDSON WYNNE
3
1
Entomology & Plant Pathology, University of Tennessee, 2505 E. J. Chapman Drive, 370 Plant Biotechnology, Knoxville, TN, USA
37996-4560. E-mail:
ebernard@utk.edu
2
Illinois Natural History Survey, University of Illinois, 1816 S Oak St., Champaign, IL 61820, E-mail: fsoto@illinois.edu
3
Department of Biological Sciences and Colourado Plateau Biodiversity Center, Northern Arizona University, Box 5640, Flagstaff,
AZ, USA 86011. URL: jutwynne.com
4
Corresponding author
Abstract
Eight species of Collembola are reported from recent collections made in caves on the Polynesian island of Rapa Nui (Eas-
ter Island). Five of these species are new to science and apparently endemic to the island: Coecobrya aitorererere n. sp.,
Cyphoderus manuneru n. sp., Entomobrya manuhoko n. sp., Pseudosinella hahoteana n. sp. and Seira manukio n. sp.
The Hawaiian species Lepidocyrtus olena Christiansen & Bellinger and the cosmopolitan species Folsomia candida Wil-
lem also were collected from one or more caves. Coecobrya kennethi Jordana & Baquero, recently described from Rapa
Nui and identified as endemic, was collected in sympatric association with C. aitorererere n.sp. With the exception of F.
candida, all species are endemic to Rapa Nui or greater Polynesia and appear to be restricted to the cave environment on
Rapa Nui. A key is provided to separate Collembola species reported from Rapa Nui. We provide recommendations to aid
in the conservation and management of these new Collembola, as well as the other presumed cave-restricted arthropods.
Key words: Chaetotaxy, taxonomy, key to species, Entomobryidae, Isotomidae, Paronellidae
Introduction
Due to a number of environmental factors including geographic isolation, island size and low topographic relief
(see Rolett & Diamond 2004), Rapa Nui (Easter Island) was predisposed to dramatic human-induced
environmental change. Following Polynesian colonization (800–1200 CE; Hunt & Lipo 2006, Shepardson et al.
2008) and sometime before European contact in 1722 (McCall 1990), a catastrophic ecological shift occurred
where the palm-dominated shrubland was ultimately replaced by grassland (Wynne et al. 2014). By the mid-
nineteenth century (several hundred years later), nearly the entire island was converted into a century-long sheep-
grazing operation (Fischer 2005). These human activities resulted in the loss of most stands of native vegetation
and the extinction of all terrestrial vertebrates.
The arthropod fauna of Rapa Nui was similarly impacted. Over the past four decades, entomologists have
commented on the impoverished native arthropod communities and the proliferation of nonnative invasive
arthropod species (Kuschel 1963, Campos & Peña 1973, Desender & Baert 1997). Prior to fieldwork conducted by
the third author, of the nearly 400 arthropod species known to the island, only 20 species (~5%) were identified as
either endemic or indigenous; the remaining arthropods were intentionally or accidentally introduced to the island
(J.J. Wynne, unpublished data).
There have been some efforts to assess Rapa Nui microarthropods (e.g., Hammer 1970, Mockford 1972), but
Collembola have scarcely been studied. Schött (1921) recorded a single specimen of Entomobrya multifasciata
(Tullberg, 1871). This record apparently was the source used by Kuschel (1963) to indicate a single species of
Collembola from Rapa Nui, but he did not provide a scientific name or attribution to Schött’s paper. The list of
arthropod species provided by Campos & Peña (1973) also included a single unnamed species, which attributed the

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report to Kuschel. Recently two additional springtail species were reported from Rapa Nui (Jordana & Baquero
2008): Arrhopalites caecus (Tullberg, 1871), a widespread Holarctic species also recorded from Australia, and
Coecobrya kennethi Jordana & Baquero, 2008, a species later identified as endemic to Rapa Nui (Wynne et al.
2014). The purpose of this paper is to describe recently collected cave Collembola fauna of Rapa Nui, including
five new species. A summary of Rapa Nui cave-restricted fauna has been published (Wynne et al. 2014).
Material and methods
Collection. This work was conducted on the Roiho lava flow, ~5 km north of the village of Hanga Roa over three
separate research trips in 2008, 2009 and 2011. The landscape surrounding the study area is characterized by gently
rolling hills (i.e., extinct scoria cones) with coastal cliff faces flanking the western-most boundary. Vegetation in
the study area was grassland and invasive guava (Psidium guajava L.) shrub. Within the collapsed pit and skylight
entrances of most caves, several invasive tree species occurred, including fig (Ficus sp.), avocado (Persea
americana Mill.), apple banana (Musa × paradisiaca L.), roseapple (Syzygium jambos (L.) Alston), P. guajava and
Eucalyptus spp.
Research teams used four methods (pitfall traps, time-constrained searches, opportunistic searches, timed
direct intuitive searches) to systematically sample 10 caves during the first two trips (16‒21 August 2008, 28
June‒17 July 2009). All specimens were placed in 95% ethanol as collected. For pitfall trapping and time-
constrained searches, sampling was performed at interval spacing along three transects, one at either wall and one
along the cave centerline. For each cave, sampling intervals were set at 10% of the cave’s length (e.g., for a 100 m
long cave, samples were taken every 10m). Three sampling stations were established at each sampling interval
where the passage width was ≥5m; otherwise, one trap was placed in a suitable location (e.g., friable sediment for
countersinking the trap). Pitfall trapping was set up for live-capture. Trap construction consisted of two 946-ml
stacked plastic containers (13.5 cm high, 10.8 cm diameter rim and 8.9 cm base). A teaspoon of peanut butter
placed in the bottom of the exterior container was used as bait. The bottom of the interior container had several
dozen holes to allow the bait to “breathe” to attract arthropods. When a pitfall trap could not be counter-sunk in the
cave sediment, ramps were built around each trap with local materials (e.g., rocks, cave sediment, wooden debris,
etc.) so invertebrates could access the trap and fall in. Rocks were placed around the edges of the trap and the trap
mouth was covered with a cap rock to discourage rodents. Traps were deployed for three to four days. Pitfall traps
were not deployed if the substrate was too hard for digging and/or lacked suitable ramp material.
Time-constrained searches involved estimating a one-meter radius around each pitfall trap sampling station
and then conducting a timed search. Searches were conducted for one to three minutes (one minute if no arthropods
were observed, three if arthropods were detected) before pitfall trap deployment and prior to trap removal.
Arthropods were collected as encountered (opportunistic collections), while deploying and removing pitfall
traps, and between timed search events. During these intervals personnel searched the ground, walls and ceilings as
they walked the length of each cave. In five caves where all the collecting methods were applied, we also
conducted timed direct intuitive searches (DIS) of fern-moss gardens by gently combing through the fern and moss
and looking beneath rocks for 40 search-minutes per garden (two observers × 20 minutes per observer), with up to
three gardens per cave. In four other caves searches were limited to fern-moss gardens only using DIS (40 minutes
total).
The deep zones of four of the caves were sampled 01‒07 August 2011 by means of bait sampling and timed
DIS. Three types of baits were placed directly on the ground and within cracks and fissures on cave walls, ceilings
and floors: sweet potato (Ipomoea batatas (L.) Lam.), chicken and fish entrails, and small branches from the local
shrubs hibiscus (Hibiscus rosa-sinensis L.) and Gaoho (Caesalpinia major (Medik.) Dandy & Exell). Two to three
stations of each bait type were deployed for four to five days within the deep zones of each cave. We also
conducted one DIS near the bait sampling arrays by searching the cave floor for 10 minutes within a 1 m2 quadrat
within the deep cave zones. Additional information on sampling and specimen numbers is available in
supplementary information attached to Wynne et al. (2014) at: http://www.bioscience.oxfordjournals.org/lookup/
suppl/doi:10.1093/biosci/biu090/-/DC1, or by contacting the third author, J. J. Wynne.
Cave Codes. The current standard practice for locality information is to provide geographical coordinates to
enable future accurate collecting and interpretation. Chilean park authorities requested that coordinates not be

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included as the caves are an extremely vulnerable resource, are strictly protected, and are not being publicized. At
the national park’s request, we used cave codes rather than actual cave names. A copy of this paper, which includes
a table of cave names with associated cave codes, is on file with CONAF-Parque Nacional Rapa Nui headquarters
Hanga Roa, Easter Island, and CONAF, Jefe Departamento, Diversidad Biológica, Gerencia de Areas Protegidas y
Medio Ambiente, Santiago, Chile.
Preservation, mounting, observation. All material was preserved in 95% ethanol. Specimens were cleared in
Marc Andre I solution, then mounted in Hoyer’s medium. Slides were cured for three days in a 50ºC oven, after
which cover slips were ringed with sealant. Line drawings were made with the aid of a drawing tube or directly
from a computer monitor. Whole-specimen images were made with 5-megapixel Nikon Coolpix 5100 or 12-
megapixel Canon EOS T31 cameras mounted on a Zeiss Stemi stereo microscope. Characters were imaged with a
14-megapixel Q-camera on an Olympus BX63 differential interference microscope system. Full body images are
provided in corrected color; colors of detail images were chosen for clarity of structures and may not represent the
actual color of the feature presented. Most high-magnification images were contrast-enhanced with software in the
Olympus system.
Characterization of chaetotaxy. All species described in this contribution belong to superfamily
Entomobryoidea and share a similar chaetotaxy pattern. In general, the dorsal chaetotaxy of the head is described
following the AMP system of Jordana & Baquero (2005) and Soto-Adames (2008), while body chaetotaxy follows
Szeptycki (1979). Postlabial setae are identified as belonging to columns I, C, E, L and O following Soto-Adames
& Giordano (2011). The macroseta formula follows Soto-Adames & Taylor (2013) and represents all macrosetae
on Th. II (posterior to the collar), Th. III, Abd. I, Abd. II, Abd. III, Abd. IV (columns A and B + Column C +
Column T and all macrosetae external to T). The distribution of inner macrosetae on Abd. IV is described
according to the topographic zones defined by Jordana (2012). Chaetotaxic descriptions for two species were
supplemented by reference to recent papers: for Coecobrya aitorererere n. sp., Chen & Christiansen (1997),
Jordana & Baquero (2008) and Zhang et al. (2011); for Cyphoderus manuneru n. sp., Szeptycki (1979) and Jantarit
et al. (2014). Labial palp terminology follows Fjellberg (1998).
Terms and abbreviations. Abbreviations used in this paper are as follows: Ant. I, Ant. II, Ant. III, Ant. IV for
antennal segments; Th. II, Th. III for the mesothorax and metathorax; Abd. I‒VI for the abdominal segments. In the
figures open or filled circles with a slash represent pseudopores; open circles represent macrosetae; and filled dots
represent mesosetae unless described otherwise in the text.
Deposition of types. All type material and specimens listed in this paper will be deposited in Museo Nacional
de Historia Natural, Casilla 787, Santiago, Chile, with the exception of specimens preserved for future DNA
analysis. Two specimens each of Entomobrya manuhoko n. sp., Coecobrya aitorererere n. sp. and C. kennethi are
maintained in ethanol at -20°C in the Entomology & Plant Pathology Department, University of Tennessee,
Knoxville, TN, USA.
Taxonomy
Family Isotomidae
Folsomia candida (Willem, 1902)
A single female of this cosmopolitan species or species complex was collected 7 August 2011 in a bait trap within
the presumed deep zone (refer to Howarth 1982), of CHILE, Rapa Nui, Maunga Hiva Hiva region, Cave Q15-076/
078. This specimen possesses 10+10 dorsal manubrial setae, 16 ventral dental setae and 6 dorsal dental setae.
These characters fit well within the limits of the taxon as defined by Potapov & Yan (2012).
Family Entomobryidae
Entomobrya manuhoko Bernard, Soto-Adames & Wynne, n. sp.
Figures 1‒3

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Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, holotype female dissected and mounted on
three slides, 1 female paratype, 1 male paratype, 5 juvenile paratypes, Cave Q15-074, direct search of fern-moss
gardens, 13 July 2009, J. Wynne, coll. Additional paratypes: 1 male, 1 female, and 1 juvenile, same data as
holotype except collected at fig tree in entrance; 3 specimens, sex undetermined, from Cave Q15-076/078, direct
search of fern-moss gardens in entrance 3, 04 July 2009, J. Wynne, coll.; 2 juveniles from Cave Q15-127,
opportune collection within entrance, 02 July 2009, J. Wynne, coll.
Description. Males and females similar except where noted. Length up to 2.3 mm. macrosetae cylindrical,
rounded at tip (Fig. 3E). In ethanol background color light cream-yellow, markings primarily reddish to brownish
violet (Figs. 1A‒D). Ant. I pale, Ant. II pale proximally, red-violet distally, Ant. III pale at base otherwise
pigmented, Ant. IV completely violet. Head (Fig. 1B) with strong interantennal band, trident-shaped spot between
eyepatches, and lunate marking posteriorly; genae with prominent light violet patches extending nearly head length
(Fig. 1A). Prothorax with violet spots dorsally. Mesothorax, metathorax and Abd. I with three broken bands of
pigment, forming large spots. Abd. II and III with five broken longitudinal bands, including a lateral band on each
side. Median band of spots on mesonotum, metanotum and Abd. I‒III tergites connected by thin line of pigment.
Lateral pigment on Abd. IV consisting of anterior V-shaped marking and compact posterior spot. Dorsum of Abd.
IV with large clear central area surrounded by spots and posterior transverse band joined to form a five-armed
figure (Figs. 1B, C), medial stripe lacking; anterior spots on Abd. IV of male weakly connected. Abd. V with
arched transverse band, Abd. VI with posterior band. Femora and tibiotarsi with wide purple bands. Furcula not
pigmented. Juveniles with smaller, discontinuous spots on Abd. IV.
Apical bulb of antenna weakly bilobed or entire, pin seta entire; apex of Ant. IV with mix of smooth, pointed
setae, multiciliate setae, and sensilliform setae of various lengths (Fig. 2E). Prelabral setae ciliate, labral setae
smooth. Labral papillae with single points (Fig. 2A). Labial palp (Fig. 2B) with all guard setae except e4, e5 and
e7; guard setae a1 and b1 stout, arising well up on papilla B; lateral process of papilla E stout, rounded apically, not
reaching spine base; five proximal setae present. Labial triangle setae in female arranged as M1M2REL1L2
A1‒5,
seta M2 shorter than but similar to neighboring setae (Fig. 2C). Outer lobe of maxillary palp with three sublobal
hairs (Fig. 2D). Eight eyes in eye patch, eyes A and B slightly larger than others (Fig. 3A).
All tibiotarsi with prominent false joint at midpoint, marked by transverse cuticular grooves (Fig. 1G), and
with several enlarged ciliate setae among thinner, more coarsely ciliate setae (Figs. 1H, I). Inner edge of unguis
with four teeth; inner basal paired teeth arising at more than half the distance from ungual base; dorsal tooth and
lateral teeth present, lateral teeth not reaching paired inner teeth. Unguiculus lanceolate, with inner edge smoothly
curved, outer edge finely serrated. Tenent hair spatulate; all tibiotarsal setae ciliate except for stout, smooth,
slightly sinuate ventro-subapical seta on hind tibiotarsus (Figs. 1I, 2G). Trochanteral organ of hind leg with up to
18 straight, smooth microsetae of various lengths (Fig. 2F). Ventral tube anteriorly with 12+ 12 or 13+13 ciliate
setae, distal three setae on each side thickened and much longer than others; 21 posterior ciliate setae, distal four
setae thicker than others; each valve with 3 smooth and 10‒14 ciliate setae (Fig. 2H). Tenaculum with four teeth on
each arm and one anterior seta on the corpus. Manubrial plaque with 8+8 ciliate mesosetae arranged slightly
asymmetrically and 3+3 pseudopores arranged in a triangle; 10+10 subdistal ciliated manubrial mesosetae in one
row (Fig. 2J). Mucro with strong anteapical tooth meeting basal spine; uncrenulated part of dens about 1.7× mucro
length (Fig. 2K).
Cephalic seta A1 a mesoseta, A0‒A5 macrosetae; An0 and An1 mesosetae; M row with three macrosetae; S0
and S2 posterior to S1 setae, S5 and S5i macrosetae or large mesosetae. Macrosetal arrangement on posterior
region of head asymmetrical with two macrosetae in Ps row and varying numbers in other P-rows (Fig. 3A). H1
area with 4 setae, H2 with 1 seta (A5), H3 without setae, H4 with 2 setae, H4’ with 4 setae, H5 with 2 setae (Fig.
3A).
Thoracic chaetotaxy (Fig. 3B) asymmetrical. On mesonotum, anterior region (zone A) with up to 22
macrosetae exclusive of collar; medial (M) zone with pseudopore and 1‒4 macrosetae; lateral (L) zone, 3
macrosetae; Pm zone, up to 33 macrosetae; Pl zone, 5 or 6 macrosetae. On metanotum, M zone with pseudopore

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FIGURE 1. Entomobrya manuhoko n. sp. A) Lateral view. B) Dorsal view, intact specimen. C) Dorsal view, thoracic and
abdominal segments after clearing and mounting. D) Head, dorsal view. E) Mesosetae of various sizes on abdominal segment
V. F) Right submedial region of mesonotum showing range of setal socket sizes. G) Tibiotarsal pseudojoint. H) Variation in
tibiotarsal seta thickness and ciliation. I) Tibiotarsal setae and location of smooth subapical seta. Inset: closeup of smooth seta.
All scales are in μm.

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FIGURE 2. Entomobrya manuhoko n. sp. A) Anterior edge of labrum. B) Labial palpus. C) Setae along labial base. D)
Maxillary palpus. E) Apex of Ant. IV. F) Trochanteral organ. G) Hind foot. H) Chaetotaxy of ventral tube: an, anterior; la,
lateral; po, posterior. I) Male genital plate. J) Manubrial plate region; black circles with slash represent pseudopores, two
mesosetae drawn to show relative sizes. K) Mucro.
and 7‒10 macrosetae; L zone, 1‒3 macrosetae; Pm zone, 11‒12 macrosetae; Pl zone, 8 macrosetae. Mesonotum
with two distinct medial lines of mesosetae (Fig. 3B) and 3‒4 irregular rows of mesosetae along posterior margin
(Figs. 1D, 3B), sockets in this posterior field elevated in contrast to more anterior mesosetal sockets (Fig. 1D).
Metanotum with similar posterior field but without distinct medial mesosetal rows.
Mesosetae dense and greatly variable in length and diameter (Fig. 1E). Abdominal segments II‒V each with
single posterior row of short mesosetae; Abd. I with increased number of mesosetae at lateral ends of row (Figs.
3B‒D). Abd. I with 12+12 macrosetae and two small, lateral sensilla. On Abd. II setal field A1 with two
macrosetae and a sensillum, field A2 with 6‒7 macrosetae. Abd. III without macrosetae in field A3 (macroseta A1
absent), field A4 with 2 macrosetae and smooth sensillum; field A5 with one macroseta; 3 macrosetae, sensillum
d2 and a minute microseta in vicinity of bothriotrix m5. Adult Abd. IV (Fig. 3C) with asymmetrical arrangement of
14‒19 macrosetae in inner region: field A6 without macrosetae; A7 with 5 macrosetae; A8 with 3‒5 macrosetae;
A9 with 3 or 4 macrosetae and 1 or 2 smooth microsetae; A10 with 3‒5 macrosetae. Lateral region with 14‒17
macrosetae. Abd. V with 20 asymmetrically arranged macrosetae; Abd. VI without macrosetae. Juvenile Abd. IV
macrosetal numbers and arrangement also asymmetrical (Fig. 3D).
Male genital plate with 18 thick setae, with enlarged bases and narrowly rounded apices (Fig. 2I).

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FIGURE 3. Entomobrya manuhoko n. sp. A) Head, adult dorsal chaetotaxy. B) Adult chaetotaxy of Th. II‒Abd. II. C) Adult
chaetotaxy of Abd. IIII‒VI. D) Late juvenile chaetotaxy of Abd. IV. E) Macroseta. Scales for A‒D = 200 μm.
Etymology. The species was named by combining two Rapanui terms, manu and hoko. Manu is Rapanui for
“insect” and used as a noun, while hoko is used as a modifier. Hoko is an ancient form of Rapanui dancing where
the dancer, usually male, jumps up and down (Fortin 2009).
Relationships. The body pattern of Entomobrya manuhoko n. sp. is quite similar to that of E. pulchra
Schäffer, 1897 (Schäffer 1897, Christiansen 1963) (Argentina, Bolivia) and the Brazilian E. pseudodecora
Rapoport, 1962 (Rapoport 1962), but differs in the shape of the dorsal pattern on Abd. IV. In E. manuhoko n. sp.

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adults have a five-armed figure without a medial band (Fig. 1B), fancifully resembling a squat penguin or a
swimming turtle. The pattern on Abd. IV of E. puchra and E. pseudodecora has disconnected spots or a weak to
strong medial band of pigment, which E. manuhoko n. sp. lacks. The mesonotum of E. manuhoko n. sp. has lateral
and medial spots, similar to that of E. pseudodecora, whereas E. pulchra has a U-shaped figure. The outer edge of
the unguiculus of E. manuhoko n. sp. is serrated, whereas the unguiculi of the other two species are illustrated as
smooth. The descriptions of E. pulchra and E. pseudodecora are brief and lack details of the chaetotaxy.
Coecobrya aitorererere Bernard, Soto-Adames & Wynne, n. sp.
Figures 4‒6, 7B
Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, holotype female and 7 paratypes (3 females, 2
males, two juveniles), all on slides, Cave Q15-071, hand collection in west passage, 12 July 2009, J. Wynne, coll.
Additional paratypes: 3 specimens, one female on slide, two of undetermined sex in ethanol, same data as holotype
except collected at fig tree in entrance; 3 specimens, one on slide, two in ethanol, sex undetermined, from Cave
Q15-076/078, direct search within fern-moss gardens in entrance 3, 04 July 2009, J. Wynne, coll.; 2 specimens, sex
undetermined, from Cave Q15-127, opportune collection within entrance, 02 July 2009, J. Wynne, coll.
Description. Length up to 1.4 mm. Body white (Fig. 7B), sometimes with sparse clusters of minute reddish
dots; all appendages white. Eyes absent. Macrosetae blunt or weakly truncate.
Antennal segment I dorsally with 3 small, smooth microsetae near the base, other setae of this segment ciliate
with three much smaller than the others (Fig. 4H); Ant. II (Figs. 4H, I) with a macroseta and several strong
mesosetae on outer side, about 12 smooth, pointed setae distributed mostly in distal half and 13 sensillum-like setae
of various lengths distributed primarily ventrally and laterally; ventromedially Ant. II with about 9 very small,
ciliate mesosetae. Ant. III (Figs. 4H, I) clothed mostly with ciliate mesosetae but also with about 23 sensilla and six
pointed, smooth setae: sensilla of sense organ paddle-like, exposed; 5 lunate sensilla along the lateral edges;
remaining 16 sensilla typical but of varying lengths. Apical bulb absent on Ant. IV (Fig. 4J); this segment with
several subapical to apical smooth, pointed setae without distinct sockets but with one or two ring-like expansions
near their bases, numerous arched sensilla that are minutely punctuate on their convex sides, typical sensilla with
rounded apices, and smooth, straight, socketed setae subapically and on the exterior face.
Prelabral setae smooth. Prelabrum and labrum (Fig. 4G) with 4-/5-5-4 setae, all setae smooth; bases of distal
two rows of setae elevated. Left mandible with four teeth, right mandible with five teeth. Outer lobe of maxillary
palp with three sublobal setae (Fig. 4F). Labial palp (Fig. 4D) with guard seta a1 stout, pointed, half the length of
b1; b2 on separate stalk, reaching tip of sensillum B; d4 shorter than other d-guard setae; lateral process stout,
rounded apically, reaching or slightly surpassing base of sensillum E; guard seta e1 short, tapering. Labial triangle
setae smooth, formula MrEL1L2A1‒5 (Fig. 4B). Six pairs of ventromedial setae along ventral groove, fifth pair
finely serrated, others smooth (Fig. 4C); arrangement of smooth and serrate postlabial setae varying between
specimens; setae x1 and x2 minute (Figs. 4B, C).
Dorsal head seta A1 absent (Fig. 4A); 3 macrosetae in zone H1, setae An1 a mesoseta; H2 with one macroseta
(A5); M-row with 3 macrosetae, M3 a mesoseta; S0 posterior to S1 setae; zone H4 with three macrosetae and a
mesoseta (S4i); H5 with two macrosetae. Group I with alternating meso- and macrosetae, Group II with 5
macrosetae. Cephalic bothriotricha short.
Hind trochanter (Fig. 5E) with 13 short, pointed microsetae in trochanteral organ and three longer setae, the
most proximal seta smooth and pointed. Hind tibiotarsus (Fig. 5F) with zig-zag row of seven smooth setae in distal
half, proximally with 1‒2 macrosetae and two long, stout mesosetae; one small ciliate seta basally and one minute
smooth microseta at level of macrosetae; other ciliate tibiotarsal setae of similar appearance. Foot complex (Fig.
5G) similar on all legs; tenent hair pointed; inner unguis edge with three teeth, paired basal teeth unequal, one tooth
large, wing-like, two small inner teeth about midway along edge; unguiculus weakly to clearly truncate (Figs.
6B‒D), with large outer wing-like tooth more than half the unguicular length.
Ventral tube (Fig. 5H) with 4+4 slender ciliate anterior setae, 3+3 short, smooth posterior setae, and 7+7 short,
smooth lateral setae. Tenaculum arms with 4 teeth each, corpus with one seta. Manubrium with two rows of smooth
setae (Fig. 5I), manubrial plaque with 2 pseudopores and two setae on each side. Dens with two smooth basal setae,
other setae ciliate. Spine of mucro (Fig. 5J) usually slightly bent, nearly reaching mucronal apex.

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FIGURE 4. Coecobrya aitorererere n. sp. A) Dorsal head chaetotaxy. B, C) Anterior ventral head chaetotaxy. D) Labial
palpus. E) Labial papilla E. F) Setae of maxillary palpus. G) Labrum. H) Ant. I–III, dorsal view, and enlargement of
crescentiform sensillum. I) Ant II–III, ventral view, and enlargement of sense rod in Ant. III apical organ. J) Apex of Ant. IV,
and enlargements of pointed seta with double basal ring and curved sensillum with punctate convex surface.

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FIGURE 5. Coecobrya aitorererere n. sp. A) Dorsal chaetotaxy: Th. II‒Abd. IV, left side, Abd. IV, right side without
mesosetal marks. B) Abd. V‒VI, left side, Question marks indicate mesosetae of uncertain affinity. C) Abd. II, bothriotrichal
region. D) Abd. III, bothriotrichal region. E) Hind trochanter. F) Hind tibiotarsus. G) Hind foot complex. H) Ventral tube; as,
anterior setae; ls, lateral setae; ps, posterior setae. I) Furcula, subdorsal view of left side, filled dots represent fringed mesosetae.
J) Mucro. Setal labels for Abd. IV and V after Zhang et al. (2011).

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FIGURE 6. Coecobrya aitorererere n. sp. A) Mesothorax, anterior lateral region; mi: microsensillum; ms: microseta. Dashed
arrow points anteriorly. B‒D) Images of unguiculi showing truncate distal region. Scales = 10 μm.
Macroseta formula 21, 20/5, 4, 3, 4+0+3, 6, 3 (Fig. 5A). Mesonotal macrosetae excluding collar setae arranged
as follows: Group I (corresponding with Zone T1) with 3 setae; Group II (Zone T2), 2; Group III, 3; Group IV, 2;
Group V, 9; Group VI, 2. One microseta dorsolaterally; laterally, relative positions of mesonotal microsensillum
and microseta (Fig. 6A) variable, microseta sometimes in more lateral position. Metanotal macroseta arrangement:
Group I, 8; Group II, 8; Group III, 2; Group IV, 2. Metanotum with two sensilla, one lateral to Group III, the other
in Group IV. Abd. I with five macrosetae and two sensilla; Abd. II with 3+1 macrosetae (m3, m3e, m3ep, m5) and
two microsetae (Figs. 5A, C); Abd. III with 1+2 macrosetae (m3, pm6, p6) and two microsetae (Figs. 5A, D). Abd.
IV (Fig. 5A) with four inner (I, M, A6, B5) and three outer macrosetae (E2‒4); approximately 15+15 sensilliform
setae, two outermost setae shorter than others; two transverse rows of four sensilliform setae anterior to I
macrosetae. Abd. V dorsally with 6+6 macrosetae (m2, m3, m5, p1, p3, p5), p4 a large, pointed mesoseta; 3+3
sensilla (as, ms, ls); extra mesosetae near m3 and a5. Abd. VI with 3+3 macrosetae (Fig. 5B).
Etymology. The species name is a combination of two Rapanui terms, aito and rererere. Aito is “jumper” and
rererere is translated as “skillful”; when combined this translates to “skillful jumper.”
Remarks. Coecobrya aitorererere n. sp. is similar to C. tenebricosa (Folsom, 1902) and will key to that
species in Chen & Christiansen (1997), Zhang et al. (2009) and Jordana (2012). The two species may be separated
by the arrangement and number of macrosetae and sensillum-like setae. The metanotum of C. aitorererere n. sp.
has eight macrosetae in Group I and eight in Group II, while C. tenebricosa has 9‒13 in Group I and 9‒10 in Group
II (Chen & Christiansen 1997, Zhang et al. 2009). On Abd. IV C. aitorererere n. sp. has two transverse rows of
four sensillum-like setae each anterior to the A3 macrosetae, whereas C. tenebricosa has a single row of six (Zhang
et al. 2009). The arrangement of Ant. III sensilla on C. aitorererere n. sp. also appears to differ markedly from that
of C. tenebricosa (Chen & Christiansen 1997) and other species figured by Zhang et al. (2009); however, these
sensilla can be difficult to see through the dense, ciliate mesosetae. Coecobrya aitorererere n. sp. differs from the
other Rapa Nui species, C. kennethi, in having two macrosetae in cephalic Group I (three in C. kennethi); two
macrosetae in mesothoracic Group IV rather than 3 as in C. kennethi; 8 macrosetae in metathoracic Group I rather
than 5; 3 outer macrosetae on Abd. IV rather than 6; ventral side of hind tibiotarsus with a zig-zag row of 7 smooth,
pointed setae (one distal smooth seta in C. kennethi).
Zhang et al. (2011) reported that the chaetotaxy of Abd. V was useful in separation of the Coecobrya spp. they
studied. In the current paper the designations of macro- or mesosetae on the abdomen were determined by
examination of specimens with setae still attached to their sockets; large, serrated, truncated setae (Fig. 5A) were
called macrosetae, and serrated, pointed setae were considered mesosetae. The m-macrosetae always originated
from large sockets, but in some specimens the sockets of the p-macrosetae (p1, p3, p5) were no larger than those of
the mesosetae, even though the attached setae clearly were macrosetae. Designations of relative setal size based on
socket size needs to be approached with caution.

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Coecobrya kennethi Jordana & Baquero, 2008
Figure 7A
Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, one specimen, Cave Q15-076/078, pitfall trap,
04 July 2009, J. Wynne, coll.; 4 specimens, Cave Q15-070, hand collection in cave, sympatric with C. aitorererere
n. sp., 10 July 2009, J. Wynne, coll.
Coecobrya kennethi is a white species (Fig. 7A) easily separated from C. aitorererere n. sp. by characters
described under that species.
FIGURE 7. Lateral views of whole specimens. A) Coecobrya kennethi Jordana & Baquero. B) Coecobrya aitorererere n. sp.
C) Pseudosinella hahoteana n. sp. D) Seira manukio n. sp. E) Lepidocyrtus olena Christiansen & Bellinger. All scales 200 μm.
Images 7A, C and D were previously published in BioScience, 64, 711‒718 (2014).
Seira manukio Soto-Adames, Bernard & Wynne, n. sp.
Figures 7D, 8, 9; Table 1
Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, holotype female, Cave Q15-056, opportune
collection near entrance, 08 July 2009, J. Wynne, coll.; one paratype juvenile, Cave Q15-076/078, direct search
within fern-moss gardens in entrance 3, sympatric with E. manuhoko n. sp., 04 July 2009, J. Wynne, coll.
Description. Size to 1.6 mm. Background color creamy white (Fig. 7D), with black eye patch, purple
antennae, a wide light purple transverse band covering posterior margin of Abd. I, all of Abd. II–III, and the
anterior margin of Abd. IV; another band covering the posterior half of Abd. IV and posterior margin of Abd. V.
Legs with a tint of purple, more intense on distal end of femora. Scales present on Ant. I–II and base of Ant. III, all
leg segments, both faces of ventral tube, head, body, both faces of manubrium and ventral face of dens.

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FIGURE 8. Seira manukio n. sp. A) Dorsal macroseta on dens base. B) Metathoracic claw complex. C) Distal end of dens and
mucro. D) Detail of eye patch. E) Dorsal chaetotaxy of head. Open circles represent macrosetae, small filled circles microsetae.
F) Chaetotaxy of labium and postlabium. Open circles represent ciliate setae, filled circles are smooth setae, and ‘V’ represents
scales. G) Ventral face of manubrium. H) Mesothorax chaetotaxy. Inset displays alternate arrangement of setae in paratype. I)
Metathorax chaetotaxy. J) Chaetotaxy of first abdominal segment.

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FIGURE 9. Seira manukio n. sp., dorsal chaetotaxy. A) Second abdominal segment. B) Third abdominal segment C) Fourth
abdominal segment.
Ant. IV apical bulb bilobed; subapical sense organ not seen. Sense organ of Ant. III not seen. Ant. II with 3
inner and 1 outer bothriotricha. Ant. I with 1 inner bothriotrix. Eye patch with 8 eyes (Figs. 8D, E); eye H smaller
than others. Eye patch valley with 4‒5 setae (q sometimes absent). Base of antennae with 5 macrosetae in series
An. Head dorsally with 12 anterior, 8 posterior and 6‒7 cervical macrosetae (Fig. 8E). Head microsetae acuminate

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and smooth. Prelabral and all labral setae smooth. Distal margin of labrum with inner papillae unisetose, outer
papillae with 4‒5 microspines. Outer maxillary lobe with basal and distal setae smooth, basal seta longer than distal
seta; sublobal plate with 3 setae. Lateral process of labial papilla E acuminate, seta-like, bent anteriorly and
surpassing tip of papilla. Proximal labial setae smooth. Labial triangle (Fig. 8F) setae as M1M2
rEL1L2A1‒5; all
posterior setae ciliate, except r, which is smooth and conic. Postlabium with very few scales, all setae ciliate (Fig.
8F), without modified setae.
Mesothorax not produced anteriorly; with 6 anterior, 9 medial, and 16 posterior macrosetae (Fig. 8H).
Metathorax with 14 macrosetae (Fig. 8I). First abdominal segment (Fig.8J) with a6 and 4 macrosetae. Second
abdominal segment (Fig. 9A) with macrosetae a2, m3, m3e and m5; supplementary setae associated with
bothriotricha m2 and a5 apically ciliate; a3 anterior to and not reaching sensillum as. Third abdominal segment
(Fig. 9B) with 1 inner (m3) and 4 lateral macrosetae (am6, pm6, p6, p7i), p7 an enlarged mesoseta; all
supplementary setae distally ciliate, seta a3 not reaching as; lateral sensilla d2 and se present. Fourth abdominal
segment (Fig. 9C) with 8 inner macrosetae: 3 in zone 7 (A4, A3, C1) 2 in zone 9 (B4, B5) and 3 in zone 10 (A5,
A6, B6); number of lateral macrosetae 10‒13. All supplementary setae distally ciliate. Posterior setae 4+4.
Trochanteral organ with at least 21 poorly organized setae. Tenent hair spatulate. Unguis (Fig. 8B) with 4 inner
teeth inserted at 53%, 53%, 74%, and 90% of inner edge; both unpaired teeth longer than paired teeth; external
teeth arising near middle of outer edge of unguis. Unguiculus lanceolate, appearing at some angles to be finely
serrate distally on all legs. Anterior face of ventral tube with 1+1 distal macrosetae, proximal setae not seen clearly;
lateral valves with 4 basal and 7‒8 distal setae; posterior face with 1+1 distal smooth setae. Manubrium with four
distal ventral setae forming a single row (Fig. 8G). Manubrial plate with 4 outer and 3 inner flexible macrosetae
separated by 2 pseudopores (Fig. 8A). Dens base with flexible macrosetae as on manubrial plate. Dens tubercle
absent. Mucro (Fig. 8C) typical for genus, falcate and without mucronal spine.
Remarks. Seira manukio n. sp. is unique among members of the genus with 4 macrosetae on Abd. I in having
12 inner mesothoracic macrosetae, 7 metathoracic macrosetae in group PA, and head macrosetae Ps2 and Pa4.
Table 1 provides a list of additional characters that distinguish the new species from other Seira with 4 Abd. I
macrosetae. The color pattern of S. manukio n. sp. resembles the pattern in S. gobalezi Christiansen & Bellinger,
1992 from Hawaiʻi; however, the head of the Hawaiian species carries only one macroseta in the M1/S1 complex,
lacks macroseta M2, while Abd. 1 has 5 macrosetae.
Etymology. This species was named by combining two Rapanui words, manu and kio. Manu is a general term
for “insect” or “bug,” while kio is the verb “to hide”, thus “hiding bug.” During times of war and conflict on the
island, the Rapanui people would seek refuge in caves (Routledge 2007). These caves are known as ana kionga or
"caves for hiding."
TABLE 1. Diagnostic table for species of Seira with four macrosetae on the first abdominal segment and S. gobalezi.
Seira gobalezi is included because its color pattern resembles that of the new species.
......
continued on the next page
Species/Character
a
Head M1/
S1
Head M2 Head S0 Head Ps2 Head Pa4 Head Pp5 Th. II setae in
complex m1-m2
manukio n. sp. 2+2 + + + + + 5‒6
bipunctata 1+1++– – – 4
delamarei 1+1 – + – – – 4
dowlingi 1+1 – + – – + 4
fulva 2+2++– – – 3
nicoya 1+1+++ – – 4
mataraquensis 2+2 + – + – – 3
socotrae 1+1 – + – – + 4
terrestris 1+1 – + ? ? ? 4‒5
gobalezi 1+1 – + ? ? ? 5

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TABLE 1. (Continued)
a
Sources: Christiansen & Bellinger 2000 [(S. bipunctata (Packard, 1873), S. delamarei Jacquemart, 1980, S. fulva
(Schött, 1896)]; Soto-Adames 2008 (S. dowlingi (Wray, 1953)); Christiansen & Bellinger 1998 (S. nicoya Christiansen &
Bellinger, 1998); Bellini & Zeppelini 2008 (S. mataraquensis Bellini & Zeppelini, 2008); Barra 2004 (S. socotrae Barra
2004); Christiansen & Bellinger 1992 (S. terrestris Folsom, 1932; S. gobalezi Christiansen & Bellinger, 1992).
Lepidocyrtus olena Christiansen & Bellinger, 1992
Figs. 7E, 10‒11; Table 2
Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, Cave Q15-074, baited pitfall trap 2A,
3.VII.2009, J. Wynne, 1 female on slide and 1 female in alcohol; 3 females, same locality data but in fern-moss
garden, 1 on slide and 2 in ethanol.
Description. Size to 1.3 mm. Background color creamy white, with purple pigment limited to eye patch and
antennae (Fig. 7E). Scales distributed on head, body and ventral face of furcula.
Apical bulb of Ant. IV absent; subapical sense organ not seen. Sense organ of Ant. III with 2 normal thin-
walled rods. Eye patch with eyes G and H subequal and smaller than others. Eye patch valley setae not seen. Dorsal
head macrosetae (Fig. 10A) limited to A0, Pa5, and 8‒9 in series An. Anterior head microsetae fusiform and
ciliate, posterior microsetae acuminate and smooth. Prelabral and all labral setae smooth. Distal margin of labrum
smooth. Outer maxillary lobe with basal and distal setae smooth and subequal; sublobal plate with 4 setae. Lateral
appendage of labial papilla E slender, bent anteriorly and reaching tip of papilla. Proximal labial setae smooth.
Labial triangle (Fig. 10B, C) setae as M1
M2rEL1L2A1-5: M2 larger than M1, which is larger than r. Distribution
of postlabial setae as in Fig. 10B: setae on anterior row smooth, setae on posterior rows ciliate; 2 lateral microsetae
reduced and smooth.
Body with dorsal macrosetal formula 00/0223+1+7. Mesothoracic hood produced anteriorly, blunt (Fig. 7E);
posterior row with 6 setae (Fig. 10D). Chaetotaxy of metathorax complete (cf. Szeptycki 1979). First abdominal
segment with a6 and 10 posterior setae. Second abdominal segment (Fig. 10G) with macrosetae m3 and m5;
supplementary setae associated with bothriotricha m2 and a5 fan-shaped; m4i and p5p absent; a2 ciliate; a2p and
all other normal setae present and smooth; a3 anterior to sensillum as. Abd. III (Fig. 10H) setae mi, ml, and a2
broad fans; a3 anterior and barely reaching sensillum as; sensillum as thicker than usual and about half the length
of m3; setae p3, p4, p5 and d2 normal; Li, Lm, Ll, a6, im, em, and am6 broad fans; a7 smooth, detached from
bothriotrichal complex. Fourth abdominal segment (Fig. 11) with 4 inner (B4, B5, B6, C1) and 9 latero-anterior
(D2, D3, E1, E2, E3, E4, F1, F2, F3) macrosetae, E1 and E4 smaller than others; supplementary seta s present, all
supplementary setae fan-shaped, D1 longest; T3 and D1p almost in row, closer to bothriotrix T2 than to T4;
macroseta F2 in arch with D2 and E2; posterior setae 12‒13.
Trochanteral organ with at least 12 setae. Metatibiotarsi without outstanding posterior setae. Tenent hair
spatulate. Unguis (Fig. 10F) with 4 inner teeth, basal pair slightly sequential rather than strictly paired, teeth
inserted at 39%, 42%, 67% and 86% of inner edge. Unguiculus truncate on all legs, posterior edge smooth. On
ventral tube, distal row of anterior face with 2 macrosetae; lateral flaps and posterior face not seen. Manubrial plate
Species/Character
a
Th. II setae in
complexes PA, PB, PC
Th. III inner
setae
Abd. I
setae
Abd. IV zone 6–7
setae
Unguiculus shape
manukio n. sp. 7, 3, 6 12 4 0 lanceolate
bipunctata 5, 3, 5–6 8 4 0 lanceolate
delamarei 5, 3, 4–5 8 4 0 lanceolate
dowlingi 5, 3, 5 8 4 0 lanceolate
fulva 5, 3, 8 8 4 0 truncate
nicoya 5, 4, 4 7 4 5 truncate
mataraquensis 5, 3, 3 7 4 0 lanceolate
socotrae 5, 3, 5 8 4 0 lanceolate
terrestris 5, 3, 4 8‒9 2, 4 0? lanceolate
gobalezi 5, 3, 5 11 5 0? lanceolate

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FIGURE 10. Lepidocyrtus olena. A) Dorsal chaetotaxy of head. B) Chaetotaxy of labium and postlabium. Open and filled
circles are coarsely ciliate and smooth setae, respectively. C) Detail of labial triangle chaetotaxy. D) Mesothorax chaetotaxy. E)
Mucro and detail of mucronal spine. F) Hind foot complex. G) Second abdominal segment chaetotaxy. H) Third abdominal
segment chaetotaxy, showing alternative condition of lateral setae. Open and filled circles represent macro- and microsetae,
respectively, unless otherwise indicated.

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FIGURE 11. Lepidocyrtus olena. Chaetotaxy of fourth abdominal segment. Open and filled circles represent macro- and
microsetae, respectively. Microsetae anterior to bothriotricha T2 and T4 are fan-shaped, as shown enlarged for setae s and D1.
of furcula with 3 inner and 4‒5 outer setae. Dens tubercle absent. Mucro (Fig. 10E) with apical tooth longer than
subapical; basal spine denticulate.
Remarks. Lepidocyrtus olena belongs to the L. nigrosetosus Folsom, 1927 species group, characterized by a
blunt, anteriorly produced mesothoracic hood, no head macroseta between A0 and Pa5, anterior row of postlabial
setae smooth, 4 inner macrosetae on Abd. IV and truncate prothoracic unguiculus. Lepidocyrtus olena can be
separated from the reasonably well-described members of this species group by the presence of ciliate labial
triangle setae M1 and r, presence of Abd. II seta m3e, and absence of dental tubercle. Of the the small sized (less
than 2 mm long) members of the L. nigrosetosus group, L. olena is most similar, and may be a junior synonym of L.
gaeyi Denis, 1924 from French Guiana or L. schmidti Handschin, 1927 from Costa Rica; the three species differ
only in that L. gaeyi and L. schmidti have three inner ungual teeth. These three species are otherwise identical in all
characters described. Of the large size (more than 2.5 mm long) members of the L. nigrosetosus group, the
individuals from Rapa Nui are most similar to L. nigrosetosus, L. immaculatus Folsom, 1932 and L. leleupi
Jacquemart, 1976, but the characters listed above distinguish the species. Christiansen and Bellinger (1992)
suggested that L. olena might refer to morphologically distinct juveniles of L. immaculatus and thus, not a valid

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species. However, one of the individuals of L. olena examined is an adult female and the differences with L.
immaculatus remain; L. immaculatus has a dental appendage and L. olena does not. In species with dental
appendage, the structure is present even in very young juveniles.
Table 2 summarises the diagnostic differences between all species listed above. Most species belonging to the
L. nigrosetosus group are poorly described and putative diagnostic characters are uncertain. The small species
described by Denis (1924) and Handschin (1927) are so poorly characterised by current standards that no detailed
comparison is possible. The larger members of the group also are difficult to distinguish, and in fact L. leleupi may
be a junior synonym of L. nigrosetosus, differing consistently only in colour pattern, whereas the only character
separating L. immaculatus from L. nigrosetosus is the number of setae along the ventral head groove (Table 2).
The presence in Rapa Nui caves of a species originally described from Hawaiʻi may seem unexpected.
However, members of the L. nigrosetosus species group appear to be tropical generalists that are easily transported
as a result of human economic activities. It is not even certain that L. olena is native to Hawaiʻi. As mentioned
above, other members of the L. nigrosetosus group occur in Central and South America and the Caribbean region.
It is possible that colonization of both Rapa Nui and Hawaiʻi by L. olena might have been mediated by ancient
Polynesians transporting plants (e.g., banana, sugarcane, taro, etc.) as they colonized eastward across the south
Pacific (Wynne et al. 2014). Alternatively, colonization may have occurred more recently within shipments of
ornamental or agricultural plants from the South American continent.
TABLE 2. Diagnostic table for New World and Hawaiian species of Lepidocyrtus without dorsal head macrochaetae
posterior to A0 but with smooth prelabral and anterior postlabial setae and truncate prothoracic unguiculus.
continued.
a
Sources: Christiansen & Bellinger 1992 (L. olena); Mari Mutt 1986 (L. nigrosetosus); Jacquemart 1976 (L. leleupi);
Denis 1924 (L. gaeyi); Handschin 1927 (L. schmidti).
Pseudosinella hahoteana Soto-Adames, Bernard & Wynne, n. sp.
Figures 7C, 12
Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, holotype (juvenile), Cave Q15-074, direct
search in fern-moss garden, sympatric with Lepidocyrtus olena, 13 July 2009, J. Wynne, coll.
Description. Size to 1.1 mm. Body uniformly light blue with darker streaking on Abd. IV (Fig. 7C). Scales
present on head, body and ventral face of furcula.
Apical bulb of Ant. IV large, unlobed (Fig. 12B); subapical sense organ acuminate, as large as guard sensillum;
chaetotaxy dominated by thin-walled, apically blunt sensilla instead of normal acuminate, thick walled setae. Sense
Species
a
Leg scales Labial seta m1 Labial seta r Setae along ventral groove
olena coxa only ciliate ciliate 4
immaculatus femur smooth reduced conic 4
nigrosetosus femur smooth or ciliate reduced conic 5–6
leleupi femur ciliate reduced conic 5–6
gaeyi ?? ? ?
schmidti ?? ? ?
Species
a
Abd. II seta m3e Inner teeth of hind unguis Hind Unguiculus Dental tubercle
olena present 4 truncate absent
immaculatus absent 4 truncate present
nigrosetosus absent 4 lanceolate to
weakly truncate
present
leleupi absent 4 truncate present
gaeyi ? 3 truncate ?
schmidti ? 3 truncate ?

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FIGURE 12. Pseudosinella hahoteana n. sp. A) Dorsal chaetotaxy of head. B) Fourth antennal segment, apical papilla. C)
Labial triangle chaetotaxy. D) Second abdominal segment chaetotaxy. E) Third abdominal segment chaetotaxy. F) Fourth
abdominal segment chaetotaxy. G) Metathoracic foot complex. H) Mucro and distal section of dens showing transition from
crenulate to smooth surface. Open and closed circles represent macro- and microsetae, respectively.

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organ of Ant. III with 2 normal rods and at least 6 additional short, blunt sensilla. Eye patch with 4 eyes (Fig. 12A).
Dorsal head macrosetae (Fig. 12A) with A0, A2, A3, Pa5 present in addition to 6+6 in series An. All head
microsetae acuminate, weakly ciliate. Prelabral setae weakly ciliate, labral setae smooth. Distal margin of labrum
smooth. Outer maxillary lobe with basal and distal setae smooth and subequal; sublobal plate with 4 setae. Lateral
appendage of labial papilla E bent anteriorly and not reaching tip of papilla. Proximal labial setae smooth. Labial
triangle setae (Fig. 12C) as M1
rE1L1L2A1-5: A1 serrate, other elements in series A smooth; r short, smooth and
conic, all other posterior setae coarsely ciliate. All postlabial setae ciliate, modified setae absent.
Dorsal macrosetae formula 00/0222+0+6. Mesothoracic hood not developed; macrosetae absent; polychaetosis
absent. Metathorax chaetotaxy complete (cf. Szeptycki 1979); macrosetae absent. On Abd. I seta a6 absent, all
posterior setae present. Second abdominal segment (Fig. 12D) with macrosetae m3 and m5 present; supplementary
setae associated with bothriotricha m2 and a5 smooth or weakly ciliate; all other microsetae smooth; setae a2p, m4i
and p5p absent; seta a3 anterior to and reaching sensillum as. Supplementary setae on Abd. III weakly ciliate (Fig.
12E), all other microsetae smooth; seta a3 anterior and reaching sensillum as; as less than half as long as m3;
sensillum d2 absent; seta a7 anterior to im and em, reaching am6. Fourth abdominal segment (Fig. 12F) with 2
inner (B5, B6) and 6 lateral macrosetae (T6, D2, E1, E2, E3, F1), E1 smaller than other macrosetae; 1‒2 additional
latero-posterior macrosetae of uncertain homology present; all supplementary setae ciliate or weakly ciliate, setae s
and a absent; seta T3 clearly anterior and not reaching D1p; microseta F2 in row with D2 and E2; posterior setae
absent.
Trochanteral organ with 6 setae, 5 arranged in 'V' formation and 1 detached. Metatibiotarsi without outstanding
posterior setae. Tenent hair acuminate. Unguis (Fig. 12G) with 3 inner teeth inserted at 34%, 46%, 56% of inner
edge; distal paired tooth minute; lateral and outer teeth ending near basal third of claw. Unguiculus lanceolate on all
legs, posterior lamella with 2‒3 small teeth.
On ventral tube distal row of anterior face with 2 macrosetae. Lateral flaps and posterior face not seen. Dens
tubercle absent. Mucro (Fig. 12H) with apical tooth slightly longer than subapical, basal spine smooth.
Remarks. Pseudosinella hahoteana n. sp. is the only member of the genus with four eyes, apical bulb on Ant.
IV and acuminate tenent hair. The new species is most similar to the East Asian P. tridentifera Rusek, 1971, from
which it differs by having an apical bulb on Ant. IV, labial seta r smooth and conic, macroseta m3 on Abd. II,
macroseta B6 on Abd. IV and inner unpaired ungual tooth inserted distally (42% in P. tridentifera vs. 56% in P.
hahoteana). Two other members of the genus, P. canariensis Gama, 1974 and P. fonsa Christiansen & Bellinger,
1992, have four eyes and only two inner macrosetae on the fourth abdominal segment, but both species carry seta
a2p on Abd. II and lack the apical bulb on Ant. IV.
Etymology. This animal was named using a combination of two Rapanui terms that when translated to English
mean “mouth of the cave.” Hahote is Rapanui for “mouth” and ana means “cave.” Currently, this species is known
only from fern-moss gardens within the cave entrance.
Family Paronellidae
Cyphoderus manuneru Bernard, Soto-Adames & Wynne, n. sp.
Figures 7B, 13‒15
Material examined. CHILE, Rapa Nui, Maunga Hiva Hiva region, holotype male, Cave Q15-070, opportune
collection on decomposing wood 50 m from cave entrance, 06 April 2011, J. Wynne, coll.
Description. White. Eyes absent. Length 1.01 mm, head diagonal 0.26 mm. Antennal segments III and IV
missing. Head and body scaled, ventral side of head with 2+2 scales (Fig. 13B); Antennal segments I and II with a
few scales (Figs. 13E, G); most scales oblong, rounded apically, with dense, evenly spaced barbs, without outer
fringe; some scales on head and antennae apically bilobed (Figs. 13G, 14B, C).
Antennal segments III and IV broken off. Ant. I dorsally with 3 small, smooth microsetae and 8 ciliate
mesosetae (Fig. 13E), ventrally with 17 smooth microsetae of various lengths and 4 ciliate mesosetae (Fig. 13F).
Dorsal side with 3 scale scars, ventral side without evidence of scales. Ant. II dorsally (Fig. 13G) with 5 scales in
basal half, minute proximo-basal microseta, one lateral and 4 distal microsetae, 3 apical sensilla, and 13 ciliate
mesosetae. Ventrally, Ant. II without scales, with 10 microsetae of which two are longer than the others, one
microseta adjacent to a smooth, pointed seta of similar length and 15 ciliated setae, several of them shorter than the
others (Fig. 15H).

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FIGURE 13. Cyphoderus manuneru n. sp. A) Dorsal chaetotaxy of head and body. B) Ventral cephalic chaetotaxy. C)
Mesochaetae. D) Lateral edge of mesonotum. E) Labrum, anterior region of head, and dorsal view of Ant. I. F) Ventral view of
Ant. I. G) Dorsal view of Ant. II. H) Ventral view of Ant. II. I) Labial palpus. J) Labial papillae E (left) and B (right) showing
guard seta bases. K) Head of maxilla.

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FIGURE 14. Cyphoderus manuneru n. sp. A) Labial palpus, arrow indicates lateral process on sensillum E. B) Oval head
scale. C) Bilobed head scales. D) Pore on Abd. III with associated grooves. E) Ciliated mesoseta A4 and associated smooth
microseta of Abd. IV. F‒H) Sublateral, dorsal, and lateral views of mucrones, respectively. Scales are in μm.
Labrum with 4/5-5-4 smooth setae (Fig. 13E), anterior margin not visible. Outer lobe of maxilla with one
sublobal hair. Head of maxilla somewhat obscured but with six lamellae (Fig. 13K), lamella 1 tapering and
extending past capitulum teeth, lamella 2 similar but shorter, lamella 3 short, lamella 4 extending horizontally,
lamellae 5 and 6 small, narrow. Left mandible with 4 teeth, right mandible with 5 teeth. Labial palp (Figs. 13I, J)
with sensilla A–E, sensilla B, D and E extending past apices of longest guard setae. Thirteen guard setae present.
Guard a1 small, oval, set in weak depression; guards b1, b2, d1, d2 and e2 long and relatively stout, most with
distinct bases (Fig. 13J); lateral process (lp) extending past base of sensillum E, slightly enlarged apically, pointed
at tip. Four proximal setae. Base of labium with smooth setae M, E, L1 and A1‒5, seta L2 minute (Fig. 13B).
Most head setae consisting of short, smooth, acuminate microsetae (Figs. 13A, B, E); body segments mostly with
small mesosetae. Prelabral setae smooth and acuminate, other frontal setae ciliate; frontal seta F0 reduced to
minute projection. Each cephalic bothriotrix with associated ciliate microseta. Lateral cephalic mesosetae long,
strongly ciliated (Fig. 13C). Head with one median seta corresponding to S0. Ventral side of head with four pairs of
medial setae, the posterior pair ciliated (Fig. 13B).
Macrosetal formula 00/00020. Mesonotum dorsally with collar of short, pointed setae, 3+3 anterior setae and
3+3 posterior setae (Fig. 13A); right side with m-seta, lacking on left side. Sensilla S1 and S2 present, S1 near
antero-lateral margin, S2 mid-lateral (Fig. 13D). Metanotum with single row of 4+4 setae, Abd. I with 3+3 setae,
Abd. II and III without setae between pseudopores. Second abdominal segment bothriotrichal region with 5 ciliate
microsetae, smooth setae absent (Fig. 15A). Outer bothriotrichal field of Abd. III with 5 ciliate and 2 smooth
microsetae, inner field with 3 ciliate and one smooth microsetae (Figs. 15B, C). Fourth abdominal segment with
inner macroseta B4 and outer macroseta E3. Mesosetae A4, A5 and T7 ciliate, their associated microsetae smooth

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and narrowly rounded apically (Figs. 13A, 14E). Seta C2 absent. Seta De3 half the length of neighboring
bothriotrix, smooth. Anterior bothriotrichal field with 5 ciliated microsetae (Fig. 15D), posterior bothriotrix
associated with macroseta E3 and with 4 ciliated microsetae, one of which is minute (Fig. 15E). Fourth abdominal
segment with 7+8 posterior setae. Fifth abdominal segment with 5+5 dorsal mesosetae.
FIGURE 15. Cyphoderus manuneru n. sp. A‒E) Abdominal sensory fields, left side, refer to Fig. 13A. A) Abd. II. B) Abd. III,
outer field. C) Abd. III, inner field. D) Abd. IV, anterior field. E) Abd. IV, posterior field. F) Trochanteral organ of hind leg. G)
Forefoot complex. H) Hind foot complex. I) Dens and mucro, dorsal view, with enlargement of smaller terminal scale-like seta.
J) Dens and mucro, ventral view.
Legs without scales; tenent hair pointed, smooth. Setae of legs consisting of stout, spinulate or strongly ciliate
mesosetae, smooth setae absent. Fore and middle ungues (Fig. 15G) with lateral and outer teeth forming a weak
tunica; inner edge with 3 teeth, basal pair with one tooth much larger, wing-like, basal paired tooth minute; distal
unpaired tooth larger than small basal tooth; unguiculus truncate distally, outer wing present. Hind foot (Fig. 15H)
similar but claws larger; unguis with doubled lateral teeth, one inner tooth and large inner wing extending past
unguiculus, which is truncate and winged. Trochanteral organ consisting of 10 small, smooth, stout pointed setae
arranged in two irregularly horizontal rows of five and a proximal seta (Fig. 15F). Ventral tube not clearly seen.
Tenaculum arms each with four teeth, corpus with one seta. Manubrial plaque with 2+2 pseudopores. Dorsal
surface of dens (Fig. 15I) with about 7 ciliate setae and at least 7 broad, densely denticulate or ciliated scales; apex
of dens with 2 long, robust scales, one about half length of mucro, other nearly as long as mucro, surfaces densely
covered with ciliate denticles that also form a fringe; each dens with two finely ciliate interior basal setae and one
smooth distal exterior seta. Ventral surface of dens (Fig. 15J) covered with slender, smooth, hyaline scales. Mucro
straight, nearly half as long as dens, with strong apical and anteapical teeth and minute, acutely pointed external
tooth almost level with anteapical tooth; fine membranes ending at external and anteapical teeth (Figs. 14F‒H, 15I,
J). Genital plate round with linear medial aperture, without associated setae.

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Labral, antennal and ventral tube pseudopores not seen. Mesonotum, metanotum and Abd. I‒IV each with one
pair of dorsal pseudopores, those of Abd. II displaced laterally; body pseudopores usually with narrow grooves in
obliquely longitudinal orientation (Fig. 14D). Posterior margin of Abd. IV with 1+2 pseudopores in posterior setal
field (Fig. 13A). Manubrial pseudopores not seen.
Etymology. The species was named by combining two Rapanui terms, manu and neru. Manu is Rapanui for
“bug” or “insect.” Neru is the name used for Rapanui girls who, historically, were secluded in caves so they would
acquire a fair complexion (Fischer 1994), thus, “pale insect”. Because all of these Collembola are believed to be
restricted to caves (i.e, isolated from surface-dwelling arthropod communities), naming one after the Rapanui
youth who remained in caves to attain a fair complexion is appropriate.
Remarks. Cyphoderus manuneru n. sp. is a member of the bidenticulati-group of Cyphoderus, possessing two
well-developed teeth on the mucro (Delamare Deboutteville 1948). The problems with identification of
Cyphoderus in this group were summarized succinctly by Jantarit et al. (2014), revolving partly around the
contradictory redescriptions of the generotype, C. albinus Nicolet, 1842. Szeptycki (1979) illustrated the
developmental chaetotaxy of C. albinus collected from ant nests in Poland. Jantarit et al. (2014) described two new
Thai species (C. songkhlaensis, C. khaochakanus), introduced several new morphological characters, and clearly
illustrated the chaetotaxic pattern of all segments. These two species must be considered the first adequately
described species in the bidenticulati-group; lack of such details for previously described species is a serious
impediment to identification of members of this genus described earlier. The labral setae of C. manuneru n. sp. are
smooth while those of the two Thai species are ciliate. Tenent hairs are pointed on C. manuneru n. sp., whereas
those of the Thai species are spatulate. Setae A4 and A5 of Abd. IV on C. manuneru n. sp. are ciliate and their
accessory microsetae are smooth; the Thai species have A4 and A5 smooth and their accessory microsetae ciliate.
Key to Collembola species reported from Rapa Nui
1. Body globular, first four abdominal segments fused, antennae elbowed between segments 3 and 4. . . . . . . Arrhopalites caecus
- Body elongated, first four abdominal segments separated, antennae straight, not elbowed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Third and fourth abdominal segments of approximately equal length; eyes absent; color white . . . . . . . . . . . Folsomia candida
- Fourth abdominal segment much longer than third segment; eyes present or absent; color variable. . . . . . . . . . . . . . . . . . . . . . 3
3. Scales on head and body absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Scales present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
4. Eyes and pigment present; mucro bidentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Eyes and pigment absent; mucro falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5. Body pigment arranged as five longitudinal bands arranged dorsally and laterally; pigment on dorsum of fourth abdominal seg-
ment arranged as a five-armed figure in adults, in juveniles as disconnected spots . . . . . . . . . . . . Entomobrya manuhoko n. sp.
- Body pigment arranged as transverse bands on posterior body segments; pigment of dorsum of fourth abdominal segment
appearing as a transverse zig-zag band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Entomobrya multifasciata
6. Hind tibiotarsus with ventral row of seven smooth pointed setae. . . . . . . . . . . . . . . . . . . . . . . . . . . .Coecobrya aitorerere n. sp.
- Hind tibiotarsus with one ventral smooth, pointed seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coecobrya kennethi
7. Mucro elongated, about half length of dens, with two strong teeth and minute third tooth; mucronal spine absent; dens with
large scale-like setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cyphoderus manuneru n. sp.
- Mucro short, less than one third length of dens, falcate or bidentate with subapical spine, dens only with small ventral scales
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Mucro falcate; body with transverse bands of pigment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seira manukio n. sp.
- Mucro bidentate, anteapical tooth present; pigment not in bands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
9. Eyes 8+8, mesonotum projecting forward over thorax; body brown or orange . . . . . . . . . . . . . . . . . . . . . . . Lepidocyrtus olena
- Eyes 4+4, mesonotum without forward projection; body blue . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudosinella hahoteana n. sp.
Discussion
We describe here five new Collembola species presumed endemic to Rapa Nui, one newly recorded species
believed to be endemic to greater Polynesia, and one cosmopolitan species. These findings are a significant
contribution to the island’s natural history, as this work increases the total number of endemic arthropod species by
nearly one third, from 20 to 26 species. These animals also represent over one half (six of 10) of the newly recorded
cave-restricted endemic arthropods of Rapa Nui. Wynne et al. (2014) gives a summary of these cave-restricted
animals.

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Two of the new species, P. hahoteana and C. manuneru, are described from single individuals; the former is a
juvenile and the latter is a male. In most cases the authors would not consider describing species based on single
specimens, and recognize that single-specimen descriptions should be avoided if possible. In the present case the
acquisition of additional specimens is not feasible without extraordinary expense. In our opinion it is unwise and
scientifically short-sighted to suppress two distinct taxa, which would bias our understanding of Rapa Nui
biodiversity. Further, we would be doing a disservice to the understanding of the island's natural history by omitting
these two species from formal description.
We know of only two previously described microarthropod species from Rapa Nui caves: the springtail
Coecobrya kennethi Jordana and Baquero (2008) (Collembola), suggested by R. Jordana (in litt.) as endemic to
Rapa Nui, and a presumed endemic psocopteran, Cyptophania pakaratii Mockford and Wynne (Mockford &
Wynne, 2013). Both are currently known only from caves. A widespread Holarctic species, Arrhopalites caecus
(Tullberg, 1871), also was reported from a cave by Jordana & Baquero (2008). A surface pitfall trapping effort in
2009 did not detect any springtails (Wynne et al. 2014). Currently, the five new species described in this paper
appear to be endemic, and along with L. olena, are also cave-restricted. Of these six Collembola species, only three
species were detected beyond the fern-moss gardens. We suggest these areas may serve as source habitats for these
animals. Northup & Welbourn (1997) proposed that moss garden habitats in New Mexico lava tube caves could
serve as source habitats for arthropods colonizing cave deep zones. Fern-moss gardens within Rapa Nui caves may
provide this same function.
Conservation and management of these species and their habitats should be a high priority for the Rapanui
community, policy makers and resource managers. Native terrestrial ecosystems on Rapa Nui are depauperate
faunistically (Desender & Baert 1997, Wynne et al. 2014). Despite intensive sampling, three of five new species
we detected had two or fewer individuals (i.e. one species with two specimens, two species with one specimen
each), and all six endemic species are believed to be restricted to caves. Thus, it is possible some of these species
are operating under extinction debts. This process is often associated with populations that have been isolated
following a significant environmental perturbation, such as habitat loss or fragmentation (Tilman et al. 1994), as is
the case for Rapa Nui. We suggest that the combined effects of anthropogenic climate change and species
interactions with non-native invasive species may further threaten the persistence of these endemic Collembola.
This combined effect challenges the persistence of surface-dwelling endemic arthropods within other island
ecosystems (Chown et al. 2007, Fordham & Brook 2010, Vitousek et al. 1997). The predicted impacts associated
with sub-tropical environments and other Polynesian islands has been discussed (e.g., IPCC 2013, Chu et. al.
2010), and we know alien species represent the majority of known arthropods on Rapa Nui.
We recommend all fern-moss garden habitats within cave entrances and beneath cave skylights be closed to
recreational use. We also recommend roping off these areas to further deter visitor access, as well as providing
signage describing the sensitivity of these areas. Fencing with wooden posts and rock cairns to support ropes
should be sufficient to deter ingress (see Wynne (2013) for additional details). Efforts should be made to place
fence posts in areas where there are no fern and moss present. Signage should be placed at trailheads and posted in
areas at proximity to where fern-moss gardens occur. Finally, additional surveys should be conducted in other
caves on the island, as well as in other habitats likely to support Collembola populations. This final step will
provide resource managers with the ability to better characterize endemic Collembola habitat, and to improve our
understanding of the distribution of these animals on Rapa Nui.
Acknowledgements
JJW wishes to thank Ninoska Cuadros Hucke, Susana Nahoe, and Enrique Tucky of CONAF-Parque Nacíonal
Rapa Nui and Consejo de Monumentos, Rapa Nui, for administrative and logistical support. Cristian Tambley of
Campo Alto Operaciones and Sergio Rapu provided additional logistical assistance. Javier Les of the Sociedad de
Ciencias Espeleológicas and Andrzej Ciszewski of the Polish Expedition team provided cave maps. Kyle Voyles
co-developed the cave-dwelling arthropod sampling protocol. Christina Colpitts, Lynn Hicks, Bruce Higgins,
Alicia Ika, Talina Konotchick, Scott Nicolay, Knutt Petersen, Lázero Pakarati, Victoria Pakarati Hotus, Pete
Polsgrove, Dan Ruby, and Liz Ruther were invaluable in the field. Juliana, Sergio, and Mata’u Rapu graciously
worked with us to assign Rapanui names for the new species. The Explorers Club and the National Speleological

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Society partially funded the field research. ECB thanks Juan Luis Jurat-Fuentes for translation of the description of
E. pseudodecora. Finally, the authors are grateful to the manuscript reviewers, whose comments greatly improved
this paper.
Literature cited
Barra, J.-A. (2004) Le genre Seira (Collembola, Entomobryidae) du Yémen continental. Zoosystema, 26, 291–306.
Bellini, B.C. & Zeppelini, D. (2008) Three new species of Seira Lubbock (Collembola, Entomobryidae) from Mataraca,
Paraíba, Brazil. Zootaxa, 1773, 44‒54.
Campos, S.L. & Peña, G.E.L. (1973) Los insectos de Isla de Pascua. Revista Chileana de Entomología, 7, 217‒229.
Chen, J.-X. & Christiansen, K. (1997) Subgenus Coecobrya of the genus Sinella (Collembola: Entomobryidae) with special
reference to the species of China. Annals of the Entomological Society of America, 90, 1‒19.
http://dx.doi.org/10.1093/aesa/90.1.1
Chown, S.L., Slabber, S., McGeoch, M.A., Janion, C. & Leinaas, H.P. (2007) Phenotypic plasticity mediates climate change
responses among invasive and indigenous arthropods. Proceedings of the Royal Society B, 274, 2531–2537.
http://dx.doi.org/10.1098/rspb.2007.0772
Christiansen, K. (1963) Preliminary notes on the genus Entomobrya in South America with special reference to Patagonia.
Biologie de l’Amérique Australe, 2, 149‒159.
Christiansen, K. & Bellinger, P. (1992) Collembola. Insects of Hawaii, 15, 1‒445.
Christiansen, K. & Bellinger, P. (1996) Cave Pseudosinella and Oncopodura new to science. Journal of Cave and Karst
Studies, 58, 38–53.
Christiansen, K. & Bellinger, P. (1998) Marine littoral Collembola of North and Central America. Bulletin of Marine Science,
42, 215‒245.
Christiansen, K. & Bellinger, P. (2000) A survey of the genus Seira (Hexapoda: Collembola: Entomobryidae) in the Americas.
Caribbean Journal of Science, 36, 39‒75.
Chu, P-S., Chen, Y.R. & Schroeder, T.A. (2010) Changes in precipitation extremes in the Hawaiian Islands in a warming
climate. Journal of Climate, 23, 4881–4900.
http://dx.doi.org/10.1175/2010JCLI3484.1
Delamare Deboutteville, C. (1948) Recherches sur les Collemboles termitophiles et myrmécophiles. Archives de Zoologie
Expérimentale et Générale, 85, 261‒425.
Denis, J.R. (1924) Sur les Collemboles du Muséum de Paris (1er Partie). Annales de la Société Entomologique de France, 93,
211‒260.
Desender, K. & Baert, L. (1997) Conservation of terrestrial arthropods on Easter Island as exemplified by the beetle fauna.
Conservation Biology, 11, 836‒838.
Fischer, S.R. (1994) Rapanui’s “great old words”: E timo te akoako. The Journal of the Polynesian Society, 103, 413‒443.
Fischer, S.R. (2005) Island at the end of the world: The turbulent history of Easter Island. Reaktion Books, London, 304 pp.
Fjellberg, A. (1998) The labial palp in Collembola. Zoologische Anzeiger, 237, 309–330.
Folsom, J.W. (1927) Insects of the Subclass Apterygota from Central America and the West Indies. Proceedings of the U. S.
Natural History Museum, 72, 1–16.
http://dx.doi.org/10.5479/si.00963801.72-2702.1
Folsom, J.W. (1932) Hawaiian Collembola. Proceedings of the Hawaiian Entomological Society, 8, 51‒80.
Fordham, D.A. & Brook, B.W. (2010) Why tropical island endemics are acutely susceptible to global change. Biodiversity and
Conservation, 19, 329–342.
http://dx.doi.org/10.1007/s10531-008-9529-7
Fortin, M. (2009) The development of theatre in Easter Island. Hakararama I Te A'amu O Rapa Nui. Unpublished M.A. thesis,
University of Otago, Dunedin, 211 pp. [New Zealand]
Gama, M.M. (1974) Systématique évolutive des
Pseudosinella. X. Espéces provenant de Yougoslavie, de Bulgarie at des Iles
Canaries (Insecta: Collembola), Revue Suisse de Zoologie, 81, 551‒559.
Hammer, M. (1970) A few oribatid mites from Easter Island. Pacific Insects, 12, 279‒289.
Handschin, E. (1927) Collemboles aus Costa Rica. Entomologische Mitteilungen, 16, 110–118.
Howarth, F.G. (1982) Bioclimatic and geologic factors governing the evolution and distribution of Hawaiian cave insects.
Entomologia Generalis, 8, 17–26.
Hunt, T. & Lipo, C. (2006) Late colonisation of Easter Island. Science, 311, 1603‒1606.
http://dx.doi.org/10.1126/science.1121879
[IPCC] Intergovernmental Panel on Climate Change (2013) Climate Change 2013: The physical science basis. Available from:
http://www.ipcc.ch/report/ar5/wg1/ (accessed 31 January 2015)
Jacquemart, S. (1976) XXII.- Collemboles noveaux des Iles Galapagos. Mission zoologique belge aus iles Galapagos et en
Ecuador (N. et J. Leleup, 1964-1965), 3, 137‒157.
Jacquemart, S. (1980) Collemboles entomobryens noveaux d’Afrique central. Bulletin de l'Institut Royal des Sciences

BERNARD ET AL.
266
·
Zootaxa 3949 (2) © 2015 Magnolia Press
Naturelles de Belgique Entomologie, 52, 1‒15.
Jantarit, S., Satasook, C. & Deharveng, L. (2014) Cyphoderus (Cyphoderidae) as a major component of collembolan cave
fauna in Thailand, with description of two new species. ZooKeys, 368, 1‒21.
http://dx.doi.org/10.3897/zookeys.368.6393
Jordana, R. (2012) Synopses on Palaearctic Collembola: Capbryinae & Entomobryini. Soil Organisms, 84, 1‒390.
Jordana, R. & Baquero, E. (2005) A proposal of characters for taxonomic identification of Entomobrya species (Collembola,
Entomobryomorpha), with description of a new species. Abhandlungen und Berichte des Naturkundemuseums, Görlitz,
76, 117‒134.
Jordana, R. & Baquero, E. (2008) Coecobrya kennethi n. sp. (Collembola, Entomobryomorpha) and presence of Arrhopalites
caecus (Tullberg, 1871) from Ana Roiho cave (Maunga Hiva Hiva), Rapa Nui-Easter Island. Euryale, 2, 68‒75.
Kuschel, G. (1963) Composition and relationship of the terrestrial faunas of Easter, Juan Fernandez, Desventuradas, and
Galapágos Islands. California Academy of Sciences, Occasional Papers, 44, 79‒95.
McCall, G. (1990) Rapanui and outsiders: The early days. In: Illius, B. & Barthel, T.S (Eds.), Circumpacifica; Festschrift für
Thomas S. Barthel. Lang, Frankfurt am Main, pp. 165‒225. [Germany]
Mari Mutt, J.A. (1986) Puerto Rican species of Lepidocyrtus and Pseudosinella (Collembola: Entomobryidae). Caribbean
Journal of Science, 22, 1‒48.
Mockford, E.L. (1972) Psocoptera records from Easter Island. Proceedings of the Entomological Society of Washington, 74,
327‒329.
Mockford, E.L. & Wynne, J.J. (2013) Genus Cyptophania Banks (Psocodea: Lepidopsocidae): Unique features, augmented
description of the generotype, and descriptions of three new species. Zootaxa, 3702 (5), 437‒449.
http://dx.doi.org/10.11646/zootaxa.3702.5.3
Northup, D.E. & Welbourn, W.C. (1997) Life in the twilight zone---lava tube ecology, natural history of El Malpais National
Monument. New Mexico Bureau of Mines and Mineral Resources Bulletin, 156, 69–82.
Packard, A.S. (1873) Synopsis of the Thysanura of Essex County, Mass., with description of a few extralimital forms. Report of
the Peabody Academy, 5, 23‒51.
Potapov, M. & Yan, G. (2012) Folsomia of China I ‒ fimetaria group (Collembola: Isotomidae). Annales de la Société
Entomologique de France, Nouvelle Série, 48, 51‒56.
http://dx.doi.org/10.1080/00379271.2012.10697750
Rapoport, E.H. (1962) Colembolos de Bahia Blanca (Argentina) III. Publicaciones del Instituto de Edafología e Hidrología,
No. 2, 3‒24.
Rolett, B. & Diamond, J. (2004) Environmental predictors of pre-European deforestation on Pacific Islands. Nature, 431, 443–
446.
http://dx.doi.org/10.1038/nature02801
Routledge, K. (2007) The mysteries of Easter Island: the story of an expedition (Reprint of original 1919 edition). Cosimo,
New York, 568 pp.
Rusek, J. (1971) Zweiter beitrag zur kenntnis der Collembola (Apterygota) Chinas. Acta Entomologica Bohemoslovaca, 68,
108‒137.
Schäffer, C. (1897) Hamburger magalhänische Sammelreise, Apterygoten, 8, 1‒48.
Schött, H. (1896) North American Apterygogenea. Proceeding of the California Academy of Science, 6, 169–196.
Schött, H. (1921) Collembola aus den Juan Fernadez-Inseln und der Osterinsel. In
: Skottsberg, C. (Ed.), The natural history of
Juan Fernandez and Easter Island, III, pp. 33‒39.
Shepardson, B., Shepardson, D., Shepardson, F., Chui, S. & Graves, M. (2008) Re-examining the evidence for late colonisation
on Easter Island. Rapa Nui Journal, 22, 97–101.
Soto-Adames, F.N. (2008) Postembryonic development of the dorsal chaetotaxy in Seira dowlingi (Collembola,
Entomobryidae); with an analysis of the diagnostic and phylogenetic significance of primary chaetotaxy in Seira. Zootaxa,
1683, 1–31.
Soto-Adames, F.N & Giordano, R. (2011) New species of springtails in the Proisotoma genus complex from Vermont and New
York, USA with descriptive notes on Ballistura alpa Christiansen & Bellinger 1980 (Hexapoda, Collembola, Isotomidae).
Zookeys, 147, 19–37.
http://dx.doi.org/10.3897/zookeys.147.2093
Soto-Adames F.N. & Taylor, J.T. (2013) Chaetotaxy and character evolution in Trogolaphysa (Collembola, Entomobryoidea,
Paronellidae), with descriptions of two new species from caves in Belize. Zookeys, 323, 35‒74.
http://dx.doi.org/10.3897/zookeys.323.4950
Szeptycki, A. (1979) Chaetotaxy of the Entomobryidae and its phylogenetical significance. Morpho-systematic studies of
Collembola, IV. Polska Akademia Nauk, Zaklad Zoologii Systematycznej Doswiadczalnej, 219 pp.
Tilman, D., May, R.M., Lehman, C.L. & Nowak, M.A. (1994) Habitat destruction and the extinction debt. Nature, 371, 65–66.
http://dx.doi.org/10.1038/371065a0
Tullberg, T. (1871) Förteckning öfver svenska Podurider. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 28,
143‒155.
Vitousek, P.M., D’Antonio, C.M., Loope, L.L., Rejmánek, M. & Westbrooks, R. (1997) Introduced species: A significant
component of human-caused global change. New Zealand Journal of Ecology, 21, 1–16.

Zootaxa 3949 (2) © 2015 Magnolia Press
·
267
COLLEMBOLA OF RAPA NUI
Wray, D.L. 1953. Some new species of springtail insects (Collembola). Occasional Papers, Nature Notes, Raleigh North
Carolina, 1, 1‒7.
Wynne, J.J. (2013) Inventory, conservation and management of lava tube caves at El Malpais National Monument. Park
Science, 34, 45‒55.
Wynne, J.J., Bernard, E.C., Howarth, F.G., Sommer, S., Soto-Adames, F.N., Taiti, S., Mockford, E.L., Horrocks, M., Pakarati,
L. & Pakarati-Hotus, V. (2014) Disturbance relicts in a rapidly changing world: The Rapa Nui (Easter Island) factor.
Bioscience, 64, 711‒718.
http://dx.doi.org/10.1093/biosci/biu090
Zhang, F., Deharveng, L. & Chen, J.-X. (2009) New species and rediagnosis of Coecobrya (Collembola: Entomobryidae), with
a key to the species of the genus. Journal of Natural History, 43, 2597‒2615.
http://dx.doi.org/10.1080/00222930903243970
Zhang, F., Yu, D. & Xu, G. (2011) Transformational homology of the tergal setae during postembryonic development in the
Sinella-Coecobrya group (Collembola: Entomobryidae). Contributions to Zoology, 80, 213–230.