Collembola of Rapa Nui (Easter Island) with

Zootaxa 3949 (2): 239–267 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press

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http://dx.doi.org/10.11646/zootaxa.3949.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:F678EF65-50F2-4B30-BDFD-A1DF3295D144

Collembola of Rapa Nui (Easter Island) with descriptions of five endemic cave-restricted species ERNEST C. BERNARD1,4, FELIPE N. SOTO-ADAMES2 & J. JUDSON WYNNE3 1

Entomology & Plant Pathology, University of Tennessee, 2505 E. J. Chapman Drive, 370 Plant Biotechnology, Knoxville, TN, USA 37996-4560. E-mail: [email protected] 2 Illinois Natural History Survey, University of Illinois, 1816 S Oak St., Champaign, IL 61820, E-mail: [email protected] 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 (Easter 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 Willem 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 midnineteenth century (several hundred years later), nearly the entire island was converted into a century-long sheepgrazing 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

Accepted by P. Greenslade: 24 Mar. 2015; published: 24 Apr. 2015

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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 timeconstrained 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 12megapixel 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 M1M2REL1L2A1‒5, 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.



251

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 M1M2rEL1L2A1‒5; 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. Species/Charactera

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

......continued on the next page



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TABLE 1. (Continued) Species/Charactera

Th. II setae in complexes PA, PB, PC

Th. III inner Abd. I setae 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

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 thinwalled 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 M1M2rEL1L2A1-5: 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


BERNARD ET AL.

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.



255

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. Speciesa

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

?

?

?

?

continued. Speciesa

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

?

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



257

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 M1rE1L1L2A1-5: 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). COLLEMBOLA OF RAPA NUI


259

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 COLLEMBOLA OF RAPA NUI


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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. 2. 3. 4. 5. 6. 7. 8. 9. -

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 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 Scales on head and body absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Scales present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Eyes and pigment present; mucro bidentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Eyes and pigment absent; mucro falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Body pigment arranged as five longitudinal bands arranged dorsally and laterally; pigment on dorsum of fourth abdominal segment 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 Hind tibiotarsus with ventral row of seven smooth pointed setae. . . . . . . . . . . . . . . . . . . . . . . . . . . .Coecobrya aitorerere n. sp. Hind tibiotarsus with one ventral smooth, pointed seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coecobrya kennethi 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 Mucro falcate; body with transverse bands of pigment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seira manukio n. sp. Mucro bidentate, anteapical tooth present; pigment not in bands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .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



265

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.


BERNARD ET AL.

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.



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