ArticlePDF Available

The terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island), with descriptions of two new species

Authors:

Abstract and Figures

Nine species of terrestrial isopods are reported for the Polynesian island of Rapa Nui (Easter Island) based upon museum materials and recent collections from field sampling. Most of these animals are non-native species, but two are new to science: Styloniscus manuvaka sp. n. and Hawaiioscia rapui sp. n. Of these, the former is believed to be a Polynesian endemic as it has been recorded from Rapa Iti, Austral Islands, while the latter is identified as a Rapa Nui island endemic. Both of these new species are considered ‘disturbance relicts’ and appear restricted to the cave environment on Rapa Nui. A short key to all the oniscidean species presently recorded from Rapa Nui is provided. We also offered conservation and management recommendations for the two new isopod species.
Content may be subject to copyright.
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 27
The terrestrial Isopoda (Crustacea, Oniscidea)
of Rapa Nui (Easter Island), with descriptions
of two new species
Stefano Taiti
1
, J. Judson Wynne
2
1 Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Via Madonna del Piano 10, 50019
Sesto Fiorentino (Florence), Italy 2 Department of Biological Sciences, Colorado Plateau Biodiversity Center,
Northern Arizona University, Box 5640, Flagsta, Arizona 86011-5614, USA
Corresponding author: Stefano Taiti (stefano.taiti@ise.cnr.it)
Academic editor: D. Bouchon|Received 28 February 2015|Accepted 18 May 2015|Published 30 July 2015
http://zoobank.org/56B35C30-E575-402C-8480-E73A7E463137
Citation: Taiti S, Wynne JJ (2015) e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island), with
descriptions of two new species. In: Taiti S, Hornung E, Štrus J, Bouchon D (Eds) Trends in Terrestrial Isopod Biology.
ZooKeys 515: 27–49. doi: 10.3897/zookeys.515.9477
Abstract
Nine species of terrestrial isopods are reported for the Polynesian island of Rapa Nui (Easter Island) based
upon museum materials and recent collections from eld sampling. Most of these animals are non-native
species, but two are new to science: Styloniscus manuvaka sp. n. and Hawaiioscia rapui sp. n. Of these, the
former is believed to be a Polynesian endemic as it has been recorded from Rapa Iti, Austral Islands, while
the latter is identied as a Rapa Nui island endemic. Both of these new species are considered ‘disturbance
relicts’ and appear restricted to the cave environment on Rapa Nui. A short key to all the oniscidean spe-
cies presently recorded from Rapa Nui is provided. We also oered conservation and management recom-
mendations for the two new isopod species.
Keywords
Crustacea, Isopoda, Oniscidea, new species, Rapa Nui, Easter Island, disturbance relicts, caves
ZooKeys 515: 27–49 (2015)
doi: 10.3897/zookeys.515.9477
http://zookeys.pensoft.net
Copyright Stefano Taiti, J. Judson Wynne. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE
Launched to accelerate biodiversity research
A peer-reviewed open-access journal
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
28
Introduction
Rapa Nui (Easter Island) is one of the most ecologically degraded islands in Polynesia.
A number of factors including geographic isolation, island size and low topographic
relief (Rolett and Diamond 2004) predisposed Rapa Nui to dramatic human-induced
environmental change. Between Polynesian colonization (800–1200 CE; Hunt and
Lipo 2006, Shepardson et al. 2008) and prior to European contact in 1722 (McCall
1990), a catastrophic ecological shift occurred where the palm-dominated shrubland
shifted to grassland (Wynne et al. 2014). By the mid-nineteenth century, most of the
island was converted into a century-long sheep-grazing operation (Fischer 2005).
Contemporarily, few native plant species remain and all terrestrial vertebrates have gone
extinct (Wynne et al. 2014). Researchers have described the arthropod communities of Rapa
Nui as being equally impoverished (Kuschel 1963, Campos and Peña 1973, Desender and
Baert 1997). Of the nearly 400 known arthropod species, only 30 species (~5%) have been
identied as either endemic or indigenous with the remaining species either intentionally or
accidentally introduced to the island (Wynne et al. 2014, Bernard et al. 2015).
rough eldwork led by the second author, at least eight island endemic and
two Polynesian endemic arthropod species have been recently identied (Wynne et
al. 2014). ese include one psocopteran (Mockford and Wynne 2013), six species
of collembolans (including ve new species and one Polynesian endemic; Bernard et
al. 2015), one recently described collembolan later identied as endemic (Jordana and
Baquero 2008; Wynne et al. 2014), and the two new species of terrestrial isopods de-
scribed in this paper. All of these animals are presumed to be cave-restricted and repre-
sent disturbance relicts – organisms now restricted to a fraction of their former range
due to extensive anthropogenic disturbance (Wynne et al. 2014). Given that one-third
of the island’s endemic arthropod fauna appear restricted to the cave environment, this
oers a unique opportunity for conservation and management.
With the exception of the Hawaiian islands (Taiti and Ferrara 1991, Taiti and
Howarth 1996, 1997, Taiti 1999, Rivera et al. 2002, Taiti et al. 2003, Santamaria
et al. 2013), terrestrial isopods from Polynesia are poorly known (see Jackson 1941
for a review). For Rapa Nui, only three species of terrestrial isopods were previously
recorded (Fuentes 1914): Ligia exotica Roux, 1828, Porcellio scaber Latreille, 1804,
and Armadillidium vulgare (Latreille, 1804). All of these isopods are non-native spe-
cies. e purpose of this paper is to identify the terrestrial isopod fauna of Rapa Nui,
including the descriptions of two new species.
Material and methods
Study area
Fieldwork was conducted on the Roiho lava ow, ~5 km north of the village of Hanga
Roa during three research trips in 2008, 2009 and 2011. e study area is characterized
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 29
by gently rolling hills (i.e., extinct scoria cones) with coastal cli faces anking the
western-most boundary. Vegetation was grassland and invasive guava (Psidium guajava)
shrub. Within the collapse pit and skylight entrances of most caves, several non-native
tree species occurred, including g (Ficus sp.), avocado (Persea americana), apple
banana (Musa × paradisiaca), roseapple (Syzygium jambos), guava (Psidium guajava)
and Eucalyptus spp.
e cave environment
Caves are zonal environments often consisting of four principle zones: (1) an entrance
(or light) zone representing a combination of both surface and cave climatic condi-
tions; (2) a twilight zone where light is diminished and surface climate conditions are
progressively dampened; (3) a transition zone characterized by complete darkness with
a further diminished inuence of surface climate conditions; and, (4) a deep zone (usu-
ally the deepest portion of the cave) where environmental conditions (e.g., complete
darkness, temperature, and air ow) remain relatively stable over time and the evapora-
tion rate is negligible (Howarth 1980, 1982). For each isopod detected within caves,
we provide a zone designation in the “type material examined” section.
Sampling
Cave and surface sampling was conducted. Research teams (led by the second author)
systematically sampled 10 caves during three research trips (16–21 August 2008; 28
June–17 July 2009; and 01–07 August 2011). Four methodologies (pitfall traps, time-
constrained searches, opportunistic collecting, and timed direct intuitive searches)
were applied to sample 10 caves during the rst two trips. Pitfall 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 exte-
rior container was used as bait. e bottom of the interior container had several dozen
holes to allow the bait to “breathe” to attract arthropods. Traps were deployed for three
to four days.
Time-constrained searches involved estimating a one-meter radius around each
pitfall trap sampling station and then conducting a timed search. Searches were con-
ducted 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.
Opportunistic collection involved collecting arthropods as encountered – while
deploying and removing pitfall traps, and between timed searches. During these in-
tervals, personnel searched the ground, walls and ceilings as they walked the length
of each cave. In ve caves (where all the collecting methodologies 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
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
30
per garden (two observers × 20 minutes per observer). In four additional caves, we
limited sampling to DIS within fern-moss gardens only (two observers × 20 minutes
per observer).
During the last research trip to the island, the deep zones of four of the caves were
sampled via bait sampling and DIS. ree types of baits were placed directly on the
ground and within cracks and ssures on cave walls, ceilings and oors: sweet potato
(Ipomoea batatas), chicken and sh entrails, and small branches from local hibiscus
(Hibiscus rosa-sinensis) and Gaoho (Caesalpinia major) shrubs. Two to three stations
of each bait type were deployed, for four to ve days, within the deep zone(s) of each
cave. At proximity to bait sampling arrays, we also conducted one DIS by searching
the cave oor for 10 minutes within a 1-m
2
area.
From 28 June through 08 July 2009 (total of 10 days), we deployed two 15 x 20
meter surface pitfall trapping grids. Surface Grid 1 (with trap numbers 1 - 20) was
established inland at the approximate center of our study area. Surface Grid 2 (with
trap numbers 21 - 40) was deployed at the western extent of the study area (~250 m
from the coastal cli face). All pitfall traps were countersunk to ground surface with
trap spacing at 5 m between each trap.
For additional information on sampling refer to Wynne et al. (2014) at: http://
www.bioscience.oxfordjournals.org/lookup/suppl/doi:10.1093/biosci/biu090/-/DC1
Cave codes
We recognize standard practice for locality information is to provide geographical
coordinates to facilitate future collecting and interpretation. However, Chilean park
ocials have requested that neither cave names nor coordinates be included due to
cultural and natural resource sensitivities of caves. In place of cave names, we used cave
codes supplied by CONAF – Parque Nacional Rapa Nui. A copy of this paper, which
includes a table of cave names with associated cave codes, is on le with CONAF –
Parque Nacional Rapa Nui headquarters Hanga Roa, Easter Island, and CONAF, Jefe
Departamento, Diversidad Biológica, Gerencia de Areas Protegidas y Medio Ambi-
ente, Santiago, Chile.
Preservation, mounting, observation
All material was preserved in 95% ethanol. Identications were based on morphologi-
cal characters with the use of micropreparations. Line drawings were made with the
aid of a camera lucida mounted on Wild M5 and M20 microscopes. Whole-specimen
images were captured using a 1.1 MP Canon 5D Mark II (with a 65 mm zoom lens)
mounted on a Visionary Digital BK Lab Plus camera mounting system. We used the
program Zerene Stacker to merge images into a composite image. Photoshop CS5 was
used for image post-processing.
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 31
Museum abbreviations
AMNH American Museum of Natural History, New York, USA;
BPBM Bernice P. Bishop Museum, Honolulu, Hawai‘i, USA;
MNHN Museo Nacional de Historia Natural, Santiago, Chile;
MZUF Museo di Storia Naturale, sezione di Zoologia, dell’Università di Firenze,
Florence, Italy;
YPM Peabody Museum of Natural History, Yale University, New Haven, Con-
necticut, USA.
Systematic account
Family Ligiidae
Genus Ligia Fabricius, 1798
Ligia exotica Roux, 1828
Ligyda exotica; Fuentes 1914: 315.
Remarks. e record of this species by Fuentes (1914) needs to be conrmed since this
littoral species has often been confused with other species in the past. Unfortunately,
no specimens of Ligia have been recently collected from Rapa Nui, most likely due to
lack of investigations along the littoral zones of the island.
Distribution. Pantropical.
Family Styloniscidae
Genus Styloniscus Dana, 1853
Styloniscus manuvaka sp. n.
http://zoobank.org/00706DDB-9E9C-4DEF-AEA5-E9289287B7AB
Figs 1A, 2–4
Styloniscus sp.; Wynne et al. 2014: 713, 714, g. 2b.
Type material examined. Chile, Rapa Nui: 1 holotype, 2 ♂♂, 2 ♀♀, 1 juv. para-
types (MNHN), Mahunga Hiva Hiva, Cave Q15-070, fern-moss garden (entrance
zone), direct intuitive search, 10.VII.2009, leg. J.J. Wynne; 2 ♂♂, 1 , 2 juvs. para-
types (MZUF), same location, 50 m from entrance, direct intuitive search (on de-
composing tree branches; twilight zone), 6.VIII.2011, leg. J.J. Wynne; 1 paratype
(MNHN), same data; 1 , 1 paratypes (BPBM), Mahunga Hiva Hiva, Cave Q15-
074, skylight entrance (1
st
entrance NE of main entrance; entrance zone), 3.VII.2009,
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
32
Figure 1. Styloniscus manuvaka sp. n.: A paratype in dorsal view. Hawaiioscia rapui sp. n.: B paratype
in dorsal view.
leg. J.J. Wynne; 1 paratype (BPBM), Mahunga Hiva Hiva, Cave Q15-119, timed
search at trap 4A, 5.VII.2009, leg. J.J. Wynne; 1 paratype (BPBM), same location,
Zone 2 (approx. cave deep zone), trap, sh entrails 1, 6.VIII.2011, leg. J.J. Wynne; 1
paratype (BPBM), Mahunga Hiva Hiva, Cave Q15-071, Zone 2 (approx. cave deep
zone), bait trap, sh entrails 1, 7.VIII.2011, leg. J.J. Wynne; 1 paratype (BPBM),
Cave Q15-067, fern-moss garden (entrance zone), direct intuitive search, 4.XII.2008,
leg J.J. Wynne.
Additional material examined. French Polynesia, Bass Islands (Austral Islands),
Rapa Iti Island: 4 ♂♂ (YPM), Pumarua-Maurua Ridge, Pumarua and some west, 500-
620 m, from dead leaves of the bird’s nest fern, Asplenium nidus, 9.I.1980, leg. G. Paulay.
Description. Maximum length: 4 mm, 4.2 mm. Dorsum brown with the
usual yellow muscle spots (Fig. 1A). Body ovoid with pleon narrower than pereon
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 33
Figure 2. Styloniscus manuvaka sp. n., paratype: A adult specimen in dorsal view B dorsal scale-seta
C cephalon in dorsal view D cephalon in frontal view E pleonite 5, telson and uropods F antennula
Gantenna.
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
34
Figure 3. Styloniscus manuvaka sp. n., paratype: A left mandible B right mandible C maxillula Dmaxilla
E maxilliped.
(Figs 1A, 2A). Vertex and pereon distinctly granulated with granulations arranged on
three rows on pereonite 1 and two rows on pereonites 2-7; pleon and telson smooth.
Dorsal surface with scale-setae as in Fig. 2B. Cephalon (Fig. 2C, D) with obtuse mid-
dle lobe slightly protruding frontwards compared with rounded lateral lobes; eye con-
sisting of three ommatidia in a triangle. Pleonites 3-5 reduced with small posterior
points. Telson (Fig. 2E) with concave sides and truncate apex. Antennula (Fig. 2F)
with second article shorter than rst and third; third article with 6 long aesthetascs at
apex. Antenna (Fig. 2G) with agellum as long as fth article of peduncle; agellum
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 35
cone-shaped, consisting of 5 articles with the second, third and fourth article bearing
two aesthetascs each. Left mandible (Fig. 3A) with 2 penicils; right mandible (Fig. 3B)
with 1 penicil. Maxillula (Fig. 3C) outer branch with 10 simple teeth and 2 long stalks;
inner branch with 3 penicils. Maxilla (Fig. 3D) apically bilobate, inner lobe wider than
outer lobe and bearing strong setae on its margin. Maxilliped (Fig. 3E) endite with a
stout apical penicil; basal article of the palp with 2 setae. Pereopods 6 and 7 with a
distinct water conducting system (Fig. 4B,C) on merus, carpus and propodus, and on
basis, ischium and merus, respectively.
Male. Pereopod 1 (Fig. 4A) merus and carpus with a line of short scales on sternal
margin. Pereopod 7 (Fig. 4C) ischium enlarged in the distal part, forming a at round-
ed lobe with two short and stout setae on tergal margin, sternal margin almost straight;
propodus with numerous long and thin setae on tergal margin. Genital papilla (Fig.
4D) with rounded and enlarged distal part. Pleopod 1 (Fig. 4D) exopodite triangular,
as wide as long, with rounded posterior margin; endopodite with agelliform distal
segment, about twice as long as basal one and slightly enlarged at apex. Pleopod 2
(Fig. 4E) exopodite very short, rectangular, about twice wider than long; endopodite
with distal segment about seven times longer than basal one, with tapering apical part
slightly bent outwards, acute apex.
Etymology. e species name is a combination of two Rapanui terms, manu and
vaka. Manu is “bug” and vaka is “canoe” or “boat”; when combined this translates to
“canoe bug.” Based upon the identication of this species, and a collembolan (Lepi-
docyrtus olena Christiansen & Bellinger, 1992) previously known from the Hawaiian
Islands only, Wynne et al. (2014) suggested both of these animals may have been
dispersed by the ancient Polynesians as they transported and transplanted cultivars
(called “canoe plants”), such as banana, taro and sugar cane, throughout the South
Pacic islands.
Remarks. At present the genus Styloniscus includes about 45 species distributed in
the tropics and the southern hemisphere (Schmalfuss 2003; Nunomura 2007; Taiti
2014). e new species is characterized by the male pereopod 7 ischium enlarged in the
distal part with a at rounded lobe. A similar character is present also in a species from
Omaio, North Island, New Zealand, identied by Vandel (1952) as Styloniscus otakensis
(Chilton, 1901). e specimens redescribed and illustrated by Vandel certainly do not
belong to S. otakensis according to the redescription of this species provided by Green
(1971) on the basis of the type material studied by Chilton (1901) and on topotypic
material (Dunedin, South Island). In fact, the male pereopod 7 ischium does not show
any distinct lobe (compare g. 31 in Green 1971 with g. 37 in Vandel 1952), and the
shapes of the male pleopod 1 exopodite and pleopod 2 endopodite are signicantly dif-
ferent (compare gs 29 and 30 in Green 1971 with gs 38 and 39A in Vandel 1952).
us, the specimens from Omaio must belong to a distinct species yet to be named.
Styloniscus manuvaka sp. n. diers from S. otakensis sensu Vandel nec Chilton in having
6 instead of 5 aesthetascs at the apex of the antennula, 5 instead of 4 agellar articles of
the antenna, the male pereopod 7 ischium with two, instead of one, stout setae on the
tergal margin, and the male pleopod 2 endopodite with a thicker distal part.
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
36
Figure 4. Styloniscus manuvaka sp. n., paratype: A pereopod 1 B pereopod 6 C pereopod 7 D genital
papilla and pleopod 1 E pleopod 2.
On Rapa Nui, Styloniscus manuvaka sp. n. is presently restricted to the cave envi-
ronment, but is not troglomorphic (cave-adapted). is animal was detected within
the fern-moss gardens (entrance zone) of three caves, but also occurred within the
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 37
twilight and cave deep zones. is species was not detected during the surface sampling
work conducted in 2009, nor has it been identied during previous invertebrate in-
ventory work (e.g., Fuentes 1914, Olalquiaga 1946, Kuschel 1963, Campos and Peña
1973). e species also occurs on Rapa Iti, Bass Islands, where it is not restricted to
the cave environment. is species is considered a Polynesian endemic and it might be
present also on other Pacic islands.
Distribution. Presently known from Rapa Nui and Rapa Iti.
Family Philosciidae
Genus Hawaiioscia Schultz, 1973
Hawaiioscia rapui sp. n.
http://zoobank.org/56E14D72-3CF5-4E39-A655-15659F01B67A
Figs 1B, 5–7
Hawaiioscia sp.; Wynne et al. 2014: 714, 716, g. 2a.
Type material examined. Chile, Rapa Nui: 1 holotype, 2 ♀♀ paratypes (MNHN),
Mahunga Hiva Hiva, Cave Q15-034, pitfall trap 5A (twilight zone) 12.VII.2009, leg.
J.J. Wynne; 1 paratype (MZUF), 1 paratype (BPBM), same data, pitfall trap
7A (approx. deep zone); 1 Paratype (MZUF), Mahunga Hiva Hiva, Cave Q15-
076/078, pitfall trap 2C (light zone), 4.VII.2009, leg. J.J. Wynne.
Description. Maximum length: and 7.5 mm. Dorsum light brown with the
usual muscle spots (Fig. 1B). Body at, ovoidal, with pleon narrower than pereon, out-
line as in Fig. 5A. Dorsal body surface nely granulated with small triangular scale-setae
(Fig. 5B). Pereonites with no sulcus marginalis, gland pores absent. Noduli laterales (Fig.
5C, G) clearly visible, inserted on a small tubercle and disposed as follows: two on the
cephalic vertex, one per side on pereonites 1-6 with that on the fourth pereonite much
more distant from the lateral margin of the segment, and two per side on pereonite
7. Cephalon (Fig. 5D–F) with short triangular lateral lobes not protruding frontwards
compared with the obtuse middle lobe; frontal and supra-antennal lines absent; eyes
small, consisting of eight ommatidia. Pleon epimera reduced but with distinct poste-
rior points (Fig. 5A,H). Telson (Fig. 5H) triangular, about twice as wide as long, with
broadly rounded apex. Antennula (Fig. 5I) of 3 articles, second article slightly shorter
than rst and third; third article bearing two rows of 7 and 2 aesthetascs each, and 2 api-
cal aesthetascs. Antenna (Fig. 6A) long and thin, reaching back rear margin of pereonite
6; agellum as long as fth peduncular article, rst agellar article distinctly longer than
second and third, with two rows of 4 to 6 aesthetascs on each second and third article.
Mandibles (Fig. 6B,C) with molar penicil semidichotomized, i.e. consisting of 3-4 setae
on a common stem; left mandible with 2+1 and right mandible with 1+1 free penicils.
Maxillula (Fig. 6D) outer branch with 5+6 teeth, all simple; inner branch with two stout
subequal penicils. Maxilla (Fig. 6E) apically setose and bilobate with outer lobe wider
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
38
Figure 5. Hawaiioscia rapui sp. n., holotype: A adult specimen in dorsal view. paratype: B dorsal
scale-seta C co-ordinates of noduli laterales D cephalon in dorsal view E cephalon in frontal view Fcepha-
lon in lateral view G pereonites with noduli laterales H pleonite 5, telson and uropods I antennula.
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 39
Figure 6. Hawaiioscia rapui sp. n., paratype: A antenna B left mandible C right mandible D maxillula
E maxilla F maxilliped.
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
40
Figure 7. Hawaiioscia rapui sp. n., paratype: A pereopod 1 B pereopod 7 C genital papilla and pleo-
pod 1 D pleopod 2 E pleopod 3 exopodite F pleopod 4 exopodite G pleopod 5 exopodite.
than inner one. Maxilliped (Fig. 6F) endite apically setose and bearing a large penicil at
medial corner, proximal article of palp bearing 2 strong setae. Pereopods with elongated
articles and agelliform dactylar and ungual setae (Fig. 7A). Pleopodal exopodites with
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 41
no trace of respiratory structures. Uropod (Fig. 5H) protopod with a /\-shaped groove on
outer margin; insertion of endopodite slightly proximal to that of exopodite.
Male. Pereopod 1 carpus with a brush of trid spines on sternal margin (Fig. 7A).
Pereopod 7 (Fig. 7B) with no peculiar modications, ischium with sternal margin
straight. Pleopod 1 (Fig. 7C) exopodite cordiform, with a broadly rounded apex; en-
dopodite with thickset distal part, straight with rounded apex. Pleopod 2 (Fig. 7D)
with exopodite triangular, shorter than endopodite and bearing 5 setae on ouer mar-
gin. Pleopods 3-5 exopodite as in Fig. 7E–G.
Etymology. e new species is named after Sergio Rapu Haoa, a humanitarian
who has furthered cultural and archeological knowledge of Rapa Nui. Sergio was Rapa
Nui’s rst governor of Rapanui descent and the rst director and curator of Museo
Antropológico P. Sebastián Englert on Rapa Nui. He is also a world-renowned Rapa
Nui archaeologist and purveyor of Rapa Nui culture. He graciously provided logistical
support to the second author and his research teams while on Rapa Nui.
Remarks. Prior to discovering this new species, the genus Hawaiioscia consisted of
four troglomorphic species restricted to lava tube caves on the Hawaiian Islands (Schultz
1973; Taiti and Howarth 1997): H. parvituberculata Schultz, 1973 from Maui, H. mi-
crophthalma Taiti & Howarth, 1997 from O‘ahu, H. paeninsulae Taiti & Howarth,
1997 from Moloka‘i, and H. rotundata Taiti & Howarth, 1997 from Kaua‘i. No epigean
species in this genus were previously known. e new species shows all the characters of
the genus Hawaiioscia with the sole exception of the molar penicil of the mandible which
is semidichotomized instead of simple as in all the others species from Hawai‘i. Consid-
ering that all the most important characters (number and position of noduli laterales,
maxillular teeth, penicil on maxillipedal endite, uropod and shape of male pleopod 1) are
shared with all the other Hawaiioscia species, we include the new species in this genus.
Specimens from this new species were collected from both within the entrance
zone of one cave and the twilight zone of another cave. It is important to note, this
species does not have troglomorphic characteristics, such as body depigmentation or
eye reduction as do other congeners within Hawaiioscia. However, as with Styloniscus
manuvaka sp. n., this new species was not detected during the surface sampling eort,
nor has it been previously identied by earlier entomological surveys of the island.
us, we believe this animal to be restricted to cave environment on Rapa Nui.
Distribution. Presently endemic to Rapa Nui.
Family Platyarthridae
Genus Trichorhina Budde-Lund, 1908
Trichorhina tomentosa (Budde-Lund, 1893)
Material examined. Chile, Rapa Nui: 1 (BPBM), Mahunga Hiva, Cave Q15-074,
pitfall trap 1B (light zone), 30.VI.2009, leg. J.J. Wynne.
Distribution. Pantropical. Introduced to greenhouses worldwide.
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
42
Family Porcellionidae
Genus Porcellionides Miers, 1877
Porcellionides pruinosus (Brandt, 1833)
Material examined. Chile, Rapa Nui: 2 ♂♂, 3 ♀♀ (AMNH 18360), Cannibal Cave,
21.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez;
1 (BPBM), Mahunga Hiva Hiva, Cave, Q15-074, time search at 1A (light zone),
30.VI.2009, leg. J.J. Wynne; 1 , 3 ♀♀ (BPBM), same location, leaf litter beneath
skylight, (3
rd
entrance NW of main entrance; entrance zone), direct intuitive search,
2.VIII.2011, leg. J.J. Wynne; 1 (BPBM), Mahunga Hiva Hiva, Cave Q15-067, fern-
moss garden (entrance zone), direct intuitive search, 10.VII.2009, leg. J.J. Wynne; 1
(BPBM), Mahunga Hiva Hiva, Cave Q15-070, fern-moss garden (entrance zone),
direct intuitive search, 10.VII.2009, leg. J.J. Wynne.
Distribution. Cosmopolitan species of Mediterranean origin.
Genus Porcellio Latreille, 1804
Porcellio scaber Latreille, 1804
Porcellio scaber; Fuentes 1914: 315; Wynne et al. 2014: 716.
Material examined. Chile, Rapa Nui: 1 (AMNH 18362), VIII.1999, leg. C.
Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez; 2 ♂♂, 1 (AMNH
18363), Maunga Tangaroa, 20.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reani-
er, H. Tonnemacher and S. Lopez; 2 ♂♂, 3 ♀♀ (AMNH 18365), Ana te Pahu,
21.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S.
Lopez; 2 ♂♂, 13 ♀♀ (AMNH 18364), Poike region, 25.VIII.1999, leg. C. Boyko,
J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez; 1 (BPBM), Mahunga
Hiva Hiva, surface in front of Cave Q15-038, opportunistic collection (eastern-most
collapse pit, southern extent near cave entrance), 18.VIII.2008, J.J. Wynne; 1
(BPBM), Mahunga Hiva Hiva, surface grid 2, 27°06'41.3"S, 109°25'09.2"W, pitfall
trap 21, 10.VII.2009, leg. J.J. Wynne; 1 juv. (BPBM), Mahunga Hiva Hiva, surface
in front of Cave Q15-038, timed search at 1C (eastern-most collapse pit on south-
ern extent near cave entrance), 20.VIII.2008, leg. J.J. Wynne; 3 ♂♂ (BPBM), Cave
Q15-038, fern-moss garden (entrance zone), direct intuitive search, 4.XII.2008,
leg. J.J. Wynne; 1 , 2 ♀♀ (BPBM), Mahunga Hiva Hiva, Cave Q15-076/078,
opportunistic collection, 4.VII.2009, leg. J.J. Wynne; 1 , 2 ♀♀ (BPBM), Ma-
hunga Hiva Hiva, Cave Q15-070, fern-moss garden (entrance zone), direct intuitive
search, 13.VII.2009, leg. J.J. Wynne; 2 ♂♂, 1 (BPBM), Mahunga Hiva Hiva,
Cave Q15-074, skylight entrance (1
st
entrance NW of main entrance; entrance
zone), opportunistic collection, 3.VII.2009, leg. J.J. Wynne; 1 juv. (BPBM), Ma-
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 43
hunga Hiva Hiva, Cave Q15-067, fern-moss garden (entrance zone), direct intui-
tive search, 10.VII.2009, leg. J.J. Wynne; 1 (BPBM), Mahunga Hiva Hiva, Cave
Q15-127, entrance zone, pitfall trap 1A, 5.VII.2009, leg. J.J. Wynne; 1 (BPBM),
same data, pitfall traps 1B; 1 , 1 (BPBM), Mahunga Hiva Hiva, surface grid 1,
27°06'53.1"S, 109°24'20.3"W, pitfall trap 3, 10.VII.2009, leg. J.J. Wynne; 1 , 1
(BPBM), same location, pitfall trap 10, 10.VII.2009, leg. J.J. Wynne; 2 ♂♂, 5 ♀♀
(BPBM), same location, pitfall trap 12, 10.VII.2009, leg. J.J. Wynne.
Distribution. Cosmopolitan species of western European origin.
Porcellio laevis Latreille, 1804
Material examined. Chile, Rapa Nui: 2 ♂♂ (AMNH 18362), VIII.1999, leg. C.
Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez; 2 ♂♂, 1 juv. (AMNH
18363), Maunga Tangaroa, 20.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reanier,
H. Tonnemacher and S. Lopez; 1 , 1 (AMNH 18365), Cave Q15-074, loca-
tion within cave not reported, 21.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reani-
er, H. Tonnemacher and S. Lopez; 2 ♂♂, 7 ♀♀ (AMNH 18361), La Pérouse Bay,
21.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez.
Distribution. Cosmopolitan species of Mediterranean origin.
Family Armadillidiidae
Genus Armadillidium Brandt, 1831
Armadillidium vulgare (Latreille, 1804)
Armadillidium vulgare; Fuentes, 1914: 315.
Material examined. Chile, Rapa Nui: 1 (AMNH 18362), VIII.1999, leg. C.
Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez; 6 ♂♂, 5 ♀♀
(AMNH 18365), Cave Q15-074, location within cave not reported, 21.VIII.1999,
leg. C. Boyko, J. Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez; 3 ♂♂, 9
♀♀ (AMNH 18366), Hotel Hanga Roa, Hanga Roa, 21.VIII.1999, leg. C. Boyko, J.
Tanacredi, S. Reanier, H. Tonnemacher and S. Lopez; 2 ♂♂, 2 ♀♀ (AMNH 18364),
Poike region, 25.VIII.1999, leg. C. Boyko, J. Tanacredi, S. Reanier, H. Tonnemach-
er and S. Lopez; 1 , 1 (BPBM), Mahunga Hiva Hiva, surface in front of Cave
Q15-038, timed search at 1B (eastern-most collapse pit, southern extent near cave
entrance), 20.VIII.2008, leg. J.J. Wynne; 1 (BPBM), Cave Q15-038, fern-moss
garden (entrance zone), direct intuitive search, 21.XIII.2008, leg. J.J. Wynne; 2 ♀♀
(BPBM), Mahunga Hiva Hiva, surface grid 2, 27°06'41.3"S, 109°25'09.2"W, pitfall
trap 23, 10.VII.2009, leg. J.J. Wynne; 1 (BPBM), surface grid 1, 27°06'53.1"S,
109°24'20.3"W, pitfall trap 17, 10.VII.2009, leg. J.J. Wynne; 3 ♀♀ (BPBM), same
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
44
location, pitfall trap 19, 10.VII.2009, leg. J.J. Wynne; 1 , 3 ♀♀ (BPBM), same loca-
tion, pitfall trap 4, 10.VII.2009, leg. J.J. Wynne.
Distribution. Cosmopolitan species of Mediterranean origin.
Family Armadillidae
Genus Venezillo Verhoe, 1928
Venezillo parvus (Budde-Lund, 1885)
Material examined. Chile, Rapa Nui: 2 ♂♂ (BPBM), Mahunga Hiva Hiva, surface
grid 2, 27°06'41.3"S, 109°25'09.2"W, pitfall trap 39, 10.VII.2009, leg. J.J. Wynne; 1
(BPBM), same location, pitfall trap 32, 10.VII.2009, leg. J.J. Wynne.
Distribution. Widespread in tropical and subtropical regions. It has been intro-
duced to European greenhouses. For diagnostic gures of this species see Schmidt
(2003).
Key to species of terrestrial isopods from Rapa Nui
1 Antennal agellum with >10 articles; eye with >100 ommatidia ...Ligia exotica
Antennal agellum with <6 articles, eye with <30 ommatidia .....................2
2 Antennal agellum of 5 articles .................................Styloniscus manuvaka
Antennal agellum of 3 or 2 articles ...........................................................3
3 Antennal agellum of 3 articles .......................................Hawaiioscia rapui
Antennal agellum of 2 articles ...................................................................4
4 Body depigmented; eye consisting of a single ommatidium ..........................
................................................................................Trichorhina tomentosa
Body pigmented; eye consisting of several ommatidia ................................. 5
5 Body slightly convex, unable to roll up into a ball .......................................6
Body strongly convex, able to roll up into a perfect ball .............................. 8
6 Cephalon with a V-shaped suprantennal line; pereonite 1 with posterior mar-
gin straight and posterior corners rounded ............ Porcellionides pruinosus
Cephalon without suprantennal line; pereonite 1 with posterior margin more
or less concave at sides and posterior corners right-angled or acute .............7
7 Dorsal body surface smooth ..................................................Porcellio laevis
Dorsal body surface distinctly granulated .............................Porcellio scaber
8 Cephalon with a triangular frontal scutellum; telson trapezoidal; uropod exo-
podite attened, lling the gap between telson and pleonite 5 ......................
............................................................................... Armadillidium vulgare
Cephalon with no frontal scutellum; telson hour-glass shaped; uropod protopo-
dite attened, lling the gap between telson and pleonite 5 ......Venezillo parvus
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 45
Discussion
Nine species of terrestrial isopods are known from Rapa Nui: Ligia exotica, Styloniscus
manuvaka sp. n., Hawaiioscia rapui sp. n., Trichorhina tomentosa, Porcellionides pruino-
sus, Porcellio laevis, P. scaber, Armadillidium vulgare, and Venezillo parvus.
Only one species (Ligia exotica) is littoral, halophilic, and widely distributed along
coastal habitats in the tropics. We have not examined any specimens belonging to
this species and its identication needs to be conrmed. Littoral habitats have not
been adequately sampled on Rapa Nui and other littoral species may also be present
on the island. Two species (Trichorhina tomentosa and Venezillo parvus) have a wide
distribution in the tropics, and four species of European or Mediterranean origin (Por-
cellionides pruinosus, Porcellio laevis, P. scaber, and Armadillidium vulgare) are now cos-
mopolitan. All of these species were introduced to Rapa Nui due to human activities.
Styloniscus manuvaka sp. n. and Hawaiioscia rapui sp. n. are Polynesian and Rapa Nui
endemics, respectively.
Given that few native arthropod species remain on Rapa Nui (Wynne et al. 2014),
the two new isopod species are a signicant contribution to the island’s natural history.
Together with the other eight endemics described by Bernard et al. (2015), Mockford
and Wynne (2013) and Jordana and Baquero (2008), these disturbance relicts have
persisted despite several hundred years of extreme environmental change and interac-
tions with non-native species (Wynne et al. 2014).
Despite their persistence, these endemic species are considered imperiled (Wynne
et al. 2014). S. manuvaka and H. rapui may be operating under extinction debts (Tri-
antis et al. 2010). is may occur once a population has become isolated following a
signicant environmental perturbation, such as habitat loss or fragmentation (Tilman
et al. 1994). Habitat loss has occurred dramatically and at an island-wide scale on Rapa
Nui. Both S. manuvaka and H. rapui were detected in low numbers. Neither of these
species were detected during earlier inventory work (see Fuentes 1914, Olalquiaga
1946, Kuschel 1963, Campos and Peña 1973) or our surface sampling eort.
Further, the combined eects of global climate change and interactions with non-
native species may further threaten the persistence of these endemic isopods. Com-
petition with non-native species has been identied as threatening the persistence of
surface-dwelling endemic arthropods on other island ecosystems (see Chown et al.
2007, Fordham and Brook 2010, Vitousek et al. 1997). Increased drought conditions
are predicted for the sub-tropics (IPCC 2013) and other Polynesian islands (Chu et
al. 2010). We also know non-native species represent the majority of known arthro-
pods on Rapa Nui. For example, of the seven known non-native isopod species, the
cosmopolitan P. scaber was detected in the greatest numbers in both surface sampling
and within caves (Wynne et al. 2014). Additionally, P. scaber is a well-established non-
native species being rst detected by Fuentes (1914). In Hawai‘i, P. scaber is consid-
ered to be an invasive species and one of the most damaging non-native arthropods in
the native ecosystems (Howarth et al. 2001).
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
46
Conservation and management of these endemic terrestrial isopods (as well as the
other endemic species) and their habitats should be a high priority for the Rapanui
community, policy makers and resource managers. Given the concerns associated with
global climate change and non-native invasive species, a captive breeding program of
these new species is recommended. Captive breeding of isopods is relatively easy and
inexpensive (Sutton 1972). Such a program may be developed in collaboration with
CONAF, Museo Antropológico P. Sebastián Englert de Rapa Nui and potentially
secondary school classrooms on the island. By captively breeding these animals in a
variety of locations, their long-term persistence may be somewhat safeguarded, and
will facilitate the establishment of viable populations for future reintroduction eorts.
Additionally, this will provide an opportunity for researchers to obtain information
associated with the life history characteristics of these endemic species. Also, once large
captive populations are established, experiments examining competition with the non-
native isopods may be performed.
Northup and Welbourn (1997) proposed that moss garden habitats in New Mex-
ico lava tube caves may serve as a source habitat for arthropods colonizing cave deep
zones. Fern-moss gardens within Rapa Nui caves may provide this same function.
All known congeners of H. rapui sp. n. are cave-adapted isopods from the Hawaiian
Islands. H. rapui sp. n. was detected within both entrance and twilight zones. If this
species persists, it is possible parapatric speciation may occur as has been suggested for
other Hawaiioscia species from the Hawaiian Islands (Rivera et al. 2002).
Finally, we know little concerning the distributions of these endemic isopods. We
recommend additional surveys be conducted in other caves on the island, as well as in
other habitats likely to support terrestrial isopods (and endemic arthropods, in gener-
al). is nal step will provide resource managers with the ability to better characterize
endemic isopod habitat, and to further 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 eld. e Explorers Club and the National
Speleological Society partially funded the eld research. Color images of S. manuvaka
sp. n. and H. rapui sp. n. were provided by Caitlin Chapman and Neil Cobb, Colorado
Plateau Museum of Arthropod Biodiversity (CPMAB), Northern Arizona University.
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 47
Jacob Higgins formerly of CPMAB assisted with identications of known isopod spe-
cies. Julianna Rapu provided suggestions on etymology and conrmations of Rapa Nui
place names. We are also grateful to Dr. Christopher Boyko and Dr. Eric A. Lazo-
Wasem for the loan of the material deposited in AMNH and YPM, respectively.
References
Bernard EC, Soto-Adames FN, Wynne JJ (2015) Collembola of Rapa Nui (Easter Island) with
descriptions of ve endemic cave-restricted species. Zootaxa 3949: 239–267.
Brandt JF (1831) Isopoda. In: Brandt JF, Ratzeburg JTC (Eds) Medizinische Zoologie oder
getreue Darstellung und Beschreibung der iere, die in der Arzneimittellehre in Betract
kommen, in systematischer Folge herausgegeben. Trowitzsch und Sohn, Berlin 2: 70–84,
pls 12–13.
Budde-Lund G (1885) Crustacea Isopoda terrestria per familias et genera et species descripta.
Nielsen and Lydiche, Hauniae [Copenhagen], 319 pp.
Budde-Lund G (1893) Landisopoder fra Venezuela, insamlede of Dr. F. Meinert. Entomologiske
Meddelelser 4: 111–129.
Budde-Lund G (1908) Isopoda von Madagaskar und Ostafrika. Mit Diagnosen verwandter
Arten. In: Voeltzkow A (Ed.) Reise in Ostafrika in den Jahren 1903-1905. Wissenschaftliche
Ergebnisse 2: 265–308, pls 12–18.
Campos SL, Peña GLE (1973) Los insectos de isla de Pascua (Resultados de une prospeccion
entomologica). Revista Chilena de Entomología 7: 217–229.
Chilton C (1901) e terrestrial Isopoda of New Zealand. Transactions of the Linnean Society
of London, Zoology 8: 99–152, pls 11–16. doi: 10.1111/j.1096-3642.1901.tb00502.x
Chown SL, Slabber S, McGeoch MA, Janion C, Leinaas HP (2007) Phenotypic plasticity me-
diates climate change responses among invasive and indigenous arthropods. Proceedings of
the Royal Society B 274: 2531–2537. doi: 10.1098/rspb.2007.0772
Christiansen K, Bellinger P (1992) Collembola. Insects of Hawaii 15: 1–445.
Chu P-S, Chen YR, Schroeder TA (2010) Changes in precipitation extremes in the Ha-
waiian Islands in a warming climate. Journal of Climate 23: 4881–4900. doi:
10.1175/2010JCLI3484.1
Dana JD (1853) Crustacea. Part 2. Isopoda. United States exploring Expedition during the
years 1838, 1839, 1840, 1841, 1842 under the Command of Charles Wilkes, U.S.N.
13(2): 696–805, pls 46–53.
Desender K, Baert L (1997) Conservation of terrestrial arthropods on Easter Island as exempli-
ed by the beetle fauna. Conservation Biology 11: 836–838.
Fabricius JC (1798) Supplementum Entomologiae Systematicae. Proft & Storch, Hafniae [=
Copenhagen], 296–306.
Fischer SR (2005) Island at the end of the world: e turbulent history of Easter Island. Reak-
tion Books, London, 304 pp.
Fordham DA, Brook BW (2010) Why tropical island endemics are acutely susceptible to global
change. Biodiversity and Conservation 19: 329–342. doi: 10.1007/s10531-008-9529-7
Stefano Taiti & J. Judson Wynne / ZooKeys 515: 27–49 (2015)
48
Fuentes F (1914) Contribucion al estudio de la fauna de la Isla de Pascua. Boletin del Museo
Nacional de Chile 7(1): 285–318, 1 map.
Green AJA (1971) Styloniscidae (Isopoda, Oniscoidea) from Tasmania and New Zealand. Papers
and Proceedings of the Royal Society of Tasmania 105: 59–74.
Howarth FG (1980) e zoogeography of specialized cave animals: a bioclimatic model. Evolution
34: 394–406. doi: 10.2307/2407402
Howarth FG (1982) Bioclimatic and geological factors governing the evolution and distribution
of Hawaiian cave insects. Entomologia Generalis 8: 17–26.
Howarth FG, Nishida GM, Evenhuis NL (2001) Insects and other terrestrial arthropods. In:
Staples GW, Cowie RH (Eds) Hawai’i’s invasive species: a guide to invasive plants and
animals in the Hawaiian Islands. Mutual Publishing & Bishop Museum Press, Honolulu,
41–62.
Hunt T, Lipo C (2006) Late colonization of Easter Island. Science 311: 1603‒1606. doi:
10.1126/science.1121879
[IPCC] Intergovernmental Panel on Climate Change (2013) Climate Change 2013: e physical
science basis. http://www.ipcc.ch/report/ar5/wg1/ [accessed 10 February 2015]
Jackson H (1941) Check-list of the terrestrial and fresh-water Isopoda of Oceania. Smithsonian
miscellaneous Collections 99: 1–35.
Jordana R, Baquero E (2008) Coecobrya kennethi sp. n. (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 Fer-
nandez, Desventuradas, and Galapágos Islands. California Academy of Sciences, Occasional
Papers 44: 79–95.
Latreille P (1804) Histoire naturelle, générale et particulière, des crustacés et des insectes.
Cloportides 7: 25–49.
McCall G (1990) Rapanui and outsiders: e early days. In: Illius B, Barthel TS (Eds)
Circumpacica; Festschrift für omas S. Barthel. Lang, Frankfurt am Main, 165–225.
Miers E (1877) On a collection of Crustacea, Decapoda and Isopoda, chiey from South
America, with descriptions of new genera and species. Proceedings of the Zoological Society
of London 1877: 653–679, pls 66–69.
Mockford EL, Wynne JJ (2013) Genus Cyptophania Banks (Psocodea: Lepidopsocidae):
Unique features, augmented description of the generotype, and descriptions of three new
species. Zootaxa 3702: 437–449. doi: 10.11646/zootaxa.3702.5.3
Northup DE, Welbourn WC (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.
Nunomura N (2007) Terrestrial isopod crustaceans from Hachijo Island, middle Japan. Bulletin
of the Toyama Science Museum 30: 17–36.
Olalquiaga FG (1946) Anotaciones entomológicas: Insectos y otros artrópodos colectados en
Isla de Pascua. Agricultura Técnica 7: 231–233.
Rolett B, Diamond J (2004) Environmental predictors of pre-European deforestation on Pacic
Islands. Nature 431: 443–446. doi: 10.1038/nature02801
e terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island)... 49
Rivera MAJ, Howarth FG, Taiti S, Roderick GK (2002) Evolution in Hawaiian cave-adapted
isopods (Oniscidea: Philosciidae): vicariant speciation or adaptive shifts? Molecular Phylo-
genetics and Evolution 25: 1–9. doi: 10.1016/S1055-7903(02)00353-6
Roux P (1828) Crustacés de la Méditerranée et de son littoral. Imprimerie d’Achard, Marseilles,
174 pp., pls 1–45. doi: 10.5962/bhl.title.8729
Santamaria CA, Mateos M, Taiti S, De Witt TJ, Hurtado LA (2013) A complex evolution-
ary history in a remote archipelago: phylogeography and morphometrics of the Hawaiian
endemic Ligia isopods. PLoS ONE 8(12): e85199. doi: 10.1371/journal.pone.0085199
Schmalfuss H (2003) World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter
Beiträge zur Naturkunde (A) 654: 1–341.
Schmidt C (2003) Contribution to the phylogenetic system of the Crinocheta (Crustacea,
Isopoda). Part 2. (Oniscoidea to Armadillidiidae). Mitteilungen aus dem Museum für
Naturkunde in Berlin, Zoologische Reihe 79: 3–179. doi: 10.1002/mmnz.20030790102
Shepardson B, Shepardson D, Shepardson F, Chui S, Graves M (2008) Re-examining the evi-
dence for late colonization on Easter Island. Rapa Nui Journal 22: 97–101.
Sutton S (1972) Woodlice. Ginn & Company Limited, London, 144 pp.
Taiti S (1999) Terrestrial isopods from Midway Atoll (Crustacea: Oniscidea). Bishop Museum
Occasional Papers 59: 37–38.
Taiti S (2014) e terrestrial Isopoda (Crustacea, Oniscidea) of the Maldives. Tropical Zoology
27: 9–33. doi: 10.1080/03946975.2014.894397
Taiti S, Arnedo MA, Lew SE, Roderick GK (2003) Evolution of terrestriality in Hawaiian
species of the genus Ligia (Crustacea, Oniscidea). Crustaceana Monographs 2: 85–102.
Taiti S, Ferrara F (1991) Terrestrial Isopods (Crustacea) from the Hawaiian Islands. Bishop
Museum Occasional Papers 31: 202–227.
Taiti S, Howarth FG (1996) Terrestrial isopods from the Hawaiian Islands (Isopoda: Oniscidea).
Bishop Museum Occasional Papers 45: 59–71.
Taiti S, Howarth FG (1997) Terrestrial isopods (Crustacea, Oniscidea) from Hawaiian caves.
Mémoires de Biospéologie 24: 97–118.
Tilman D, May RM, Lehman CL, Nowak MA (1994) Habitat destruction and the extinction
debt. Nature 371: 65–66. doi: 10.1038/371065a0
Triantis KA, Borges PAV, Ladle RJ, Hortal J, Cardoso P, Gaspar C, Dinis F, Mendonça E, Silveira
LMA, Gabriel R, Melo C, Santos AMC, Amorim IR, Ribeiro SP, Serrano ARM, Quartau
JA, Whittaker RJ (2010) Extinction debt on oceanic islands. Ecography 33: 285–294. doi:
10.1111/j.1600-0587.2010.06203.x
Vandel A (1952) Les trichoniscides (Crustacés - Isopodes) de l’hémisphère austral. Leur place
systématique leur intérêt biogéographique. Mémoires du Muséum National d’Histoire
Naturelle (A) 6: 1–116.
Verhoe KW (1928) Über einige Isopoden der Zoologischen Staatssammlung in München.
38. Isopoden. Aufsatz. Zoologischer Anzeiger 76: 25–36, 113–123.
Vitousek PM, D’Antonio CM, Loope LL, Rejmánek M, Westbrooks R. (1997) Introduced
species: A signicant component of human-caused global change. New Zealand Journal
of Ecology 21: 1–16.
Wynne JJ, Bernard EC, Howarth FG, Sommer S, Soto-Adames FN, Taiti S, Mockford EL,
Horrocks M, Pakarati L, Pakarati-Hotus V (2014) Disturbance relicts in a rapidly chang-
ing world: the Rapa Nui (Easter Island) factor. BioScience 64(8): 711–718. doi: 10.1093/
biosci/biu090
... Each was a short-range endemic species with each species detected within an individual cave; these four species were described from coastal Kauai, Maui, Molokai, and Oahu. Nearly two decades later, Hawaiioscia rapui Taiti & Wynne, 2015 was described from two caves on Rapa Nui (Easter Island). Initially propounded as an island endemic and disturbance relict (i.e., a species with a relictual distribution due to anthropogenic activities), this terrestrial isopod was believed to be restricted to caves due to extensive surface disturbance (Taiti & Wynne 2015;Wynne et al. 2014). ...
... Nearly two decades later, Hawaiioscia rapui Taiti & Wynne, 2015 was described from two caves on Rapa Nui (Easter Island). Initially propounded as an island endemic and disturbance relict (i.e., a species with a relictual distribution due to anthropogenic activities), this terrestrial isopod was believed to be restricted to caves due to extensive surface disturbance (Taiti & Wynne 2015;Wynne et al. 2014). As this species was not subterranean-adapted, the authors posited it may have had an island-wide distribution prior to the arrival of the ancient Polynesians to Rapa Nui (Wynne et al. 2014(Wynne et al. , 2016. ...
... In most cases, important taxonomic characters cannot be sufficiently resolved resulting in questionable identifications. All specimens were identified using the species description for H. rapui and the taxonomic key provided in Taiti & Wynne (2015). As the only character(s) requiring measurements was the length of the habitus, data for individuals from the MMH and Rapa Nui populations ( Fig. 2) are provided. ...
Article
Full-text available
Hawaiioscia rapui Taiti & Wynne, 2015 was first described from two caves on Rapa Nui and considered a potential island endemic and disturbance relict (i.e., an organism that becomes a relict species due to anthropogenic activities). As this species was not subterranean-adapted, it may have had an island-wide distribution prior to the arrival of the ancient Polynesians to Rapa Nui. We report new records for Hawaiioscia rapui beyond its type locality. These findings extend this animal’s range to the closest neighboring island, Motu Motiro Hiva (MMH), 414 km east by northeast of Rapa Nui. We also report information on this animal’s natural history, discuss potential dispersal mechanisms, identify research needs, and provide strategies for management. Our discovery further underscores that MMH likely harbors a unique and highly adapted halophilic endemic arthropod community. Conservation policies will be required to prevent alien species introductions; additionally, an inventory and monitoring program should be considered to develop science-based strategies to manage the island’s ecosystem and species most effectively.
... Species were identi ed using dichotomous keys and scienti c articles (Noël & Séchet, 2007;Taiti & Judson Wynne, 2015;Vandel, 1960Vandel, , 1962). ...
Preprint
Full-text available
Several studies report the high capacity of bioindication of Isopoda (Crustacea, Oniscidea), related to their important ability to accumulate contaminants, useful in soil ecotoxicology and in bioindication activities. Any change in the Isopods population, diversity, and life cycle can indicate relevant pollution levels. The analysis of target tissues, as hepatopancreas, is another emerging approach (from a cytologic/histologic level) to detect contaminant accumulation from different sources. In this study, tissue disaggregation procedures were optimised on hepatopancreas, and Flow Cytometry (FC) was applied to detect cell viability and several cell functions. After disaggregation, two hepatopancreatic cell types, Small (S) and Big (B), were still recognizable: they differ in morphology and behaviour. The analyses were conducted for the first time on Isopods from sites at different conditions of ecological disturbance through a cytometric re-interpretation of ecological-environmental parameters. Significant differences in cell functional parameters were found, highlighting that Isopod hepatopancreatic cells can be efficiently analysed by FC and represent standardisable, early biologic indicators, tracing environmental-induced stress through cytologic/histologic analyses.
... either occurred or still occurs in similar habitats on the surface, the importance of relict plant species restricted to cave entrances has been discussed for southern China (Monro et al. 2018). Additionally, several arthropod species globally are restricted to cave entrances in Polynesia (Mockford and Wynne 2013, Bernard et al. 2015, Taiti and Wynne 2015 and North America (Benedict 1979, Wynne andShear 2016) due to either extensive surface disturbance and glacial interglacial cycles, respectively. Thus, it is possible this species is a 'disturbance relict' restricted to the entrance of Shangshuiyan Cave and potentially other area cave entrances with similar vegetation. ...
Article
Full-text available
We synthesized the current knowledge of cave-dwelling millipede diversity from Guangxi Zhuang Autonomous Region (Guangxi), South China Karst, China and described six new millipede species from four caves from the Guilin area, northeastern Guangxi. Fifty-two cave-dwelling millipedes are known for the region consisting of 38 troglobionts and 14 troglophiles. Of the troglobionts, 24 are presently considered single-cave endemics. New species described here include Hyleoglomerisrukouqu sp. nov. and Hyleoglomerisxuxiakei sp. nov. (Family Glomeridae), Hylomusyuani sp. nov. (Family Paradoxosomatidae), Eutrichodesmusjianjia sp. nov. (Family Haplodesmidae), Trichopeltisliangfengdong sp. nov. (Family Cryptodesmidae), and Glyphiulusmaocun sp. nov. (Family Cambalopsidae). Our work also resulted in range expansions of Pacidesmustrifidus Golovatch & Geoffroy, 2014, Blingulussinicus Zhang & Li, 1981 and Glyphiulusmelanoporus Mauriès & Nguyen Duy-Jacquemin, 1997. As with many hypogean animals in Southeast Asia, intensive human activities threaten the persistence of both cave habitats and species. We provide both assessments on the newly described species’ distributions and recommendations for future research and conservation efforts.
... Two sampling methodologies were used. Direct Intuitive Searches (Taiti & Wynne 2015), implemented at all sites, consisted of hand-collection of specimens during searches in the sand and decomposing organic matter, roots and bark of trees, fallen logs and under rocks. The estimated time of the searches was about 20 minutes per observer (three observers) at each sampling site. ...
Article
Full-text available
In this study, we report the results of a taxonomic survey of terrestrial isopods from Isla Grande, a ca 2 km2 island located in the north of Cartagena de Indias, Colombia. We found a total of 17 species belonging to nine families and 10 genera. Eight of these species have been reported only from the Caribbean region, nine are recorded for the first time in Colombia, and three are new to science and described here: Tylos negroi López-Orozco, Carpio-Díaz & Campos-Filho sp. nov., Stenoniscus nestori López-Orozco, Taiti & Campos-Filho sp. nov. and Armadilloniscus luisi Carpio-Díaz, Taiti & Campos-Filho sp. nov. Our revision also determined that the genus Microphiloscia is a junior synonym of Halophiloscia; and moreover, Halophiloscia trichoniscoides comb. nov. is redescribed. We also provide illustrations for Armadilloniscus caraibicus and Armadilloniscus ninae. Most (16) of the species were found in coastal habitats (i.e., beaches, coastal lagoons and mangroves), whereas the tropical dry forest harbored only two species. Molecular phylogenetic inferences supported the presence of three species of Tylos in Isla Grande (i.e., one new species and a new lineage within each of two known species). Our work on Tylos highlights the importance of combining taxonomic and molecular analyses to support taxonomic decisions and uncover cryptic diversity. Due to the multiple threats to Caribbean coastal habitats, taxonomic and molecular genetic research are urgently needed to understand biodiversity patterns of oniscideans in the Caribbean, which will inform strategies for their protection. Such studies will also contribute to our knowledge of their evolution, ecology, and potential uses, as well as the factors that have shaped the remarkable Caribbean biodiversity.
... These systems often support troglomorphic (subterranean-adapted) species with narrow geographic ranges (i.e., occurring within a single cave or watershed [6][7][8][9][10][11][12][13][14][15]) and are often represented by small populations [16,17]. Cave entrances have also been identified as important habitats for relict arthropod species from the last glaciation [18][19][20][21][22] and extensive surface disturbance [23][24][25]. While some areas have been identified as hotspots for endemism and diversity [7,26], cave communities in most regions globally remain largely unknown. ...
Article
Full-text available
Since the initial experiments nearly 50 years ago, techniques for detecting caves using airborne and spacecraft acquired thermal imagery have improved markedly. These advances are largely due to a combination of higher instrument sensitivity, modern computing systems, and processor intensive analytical techniques. Through applying these advancements, our goals were to: (1) Determine the efficacy of methods designed for terrain analysis and applied to thermal imagery; (2) evaluate the usefulness of predawn and midday imagery for detecting caves; and (3) ascertain which imagery type (predawn, midday, or the difference between those two times) was most informative. Using forward stepwise logistic (FSL) and Least Absolute Shrinkage and Selection Operator (LASSO) regression analyses for model selection, and a thermal imagery dataset acquired from the Mojave Desert, California, we examined the efficacy of three well-known terrain descriptors (i.e., slope, topographic position index (TPI), and curvature) on thermal imagery for cave detection. We also included the actual, untransformed thermal DN values (hereafter "unenhanced thermal") as a fourth dataset. Thereafter, we compared the thermal signatures of known cave entrances to all non-cave surface locations. We determined these terrain-based analytical methods, which described the "shape" of the thermal landscape hold significant promise for cave detection. All imagery types produced similar results. Down-selected covariates per imagery type, based upon the FSL models, were: Predawn-slope, TPI, curvature at 0 m from cave entrance, as well as slope at 1 m from cave entrance; midday-slope, TPI, and unenhanced thermal at 0 m from cave entrance; and difference-TPI and slope at 0 m from cave entrance, as well as unenhanced thermal and TPI at 3.5 m from cave entrance. Finally, we provide recommendations for future research directions in terrestrial and planetary cave detection using thermal imagery.
... Eight woodlice species have been reported on Rapa Nui (Taiti and Wynne 2015). Styloniscus manuvaka and Hawaiioscia rapui are endemic, and the rest are cosmopolitan. ...
Article
Full-text available
Rongorongo is a non-deciphered writing system from Rapa Nui (Easter Island). Because the island was isolated from the outside world until relatively recently, rongorongo has the potential of being one of only a few instances in human history of an independent invention of writing. However, no scientific consensus exists regarding the time span for when rongorongo was used. Its cessation in the 1860s is well-known but its origins are not. Here, we report on detailed analysis of one of the 23 existing rongorongo artifacts-the Berlin Tablet-including botanical wood identification, radiocarbon dating, and photogrammetric study. The wood used to create the tablet was identified as Pacific rosewood, Thespesia populnea, a species that once grew on Rapa Nui, which counters previous theories that the tablet was made from salvaged driftwood. The radiocarbon date, adjusted in accordance to the ethnographic data, suggests that the tablet was made some time between ca. AD 1830 and 1870. Prior to its collection, the tablet had spent a significant amount of time within a cave context that destroyed around 90% of its content. The text is estimated to have been over 5000 signs long, more than double the length of the next longest rongorongo text.
... Especie cosmopolita, originaria del área mediterránea (Schmalfuss 2003). En Chile es frecuente en hábitats intervenidos; sin embargo, solo ha sido registrada formalmente en Rapa Nui (Taiti y Wynne 2015). El material aquí examinado corresponde al primer registro en Chile continental. ...
Article
Full-text available
Terrestrial isopods (Crustacea: Isopoda: Oniscidea) from Bosque Fray Jorge National Park, Coquimbo Region, Chile. The Bosque Fray Jorge National Park, located in the coastal mountain range of the Coquimbo region, occupies an area of arid mediterranean climate with marine influence, dominated by vegetative formations of desert scrub and with presence of forested relics associated with peaks exposed to coastal fogs. This work describes the terrestrial isopod fauna of the Bosque Fray Jorge National Park based on material obtained during two campaigns carried out in 2017. The surveys were successful only in damp microhabitats, such as the rocky supratidal, mist forests, water bodies and springs associated with human settlements. A total of 12 species, distributed in 11 genera and 9 families were registered, which constitutes an increase of 92% in the local oniscofauna diversity. A half of the isopods species collected are exotic, while among native species, at least two of them have not been described.The exotic species Haplophthalmus danicus is the first representative of the family Trichoniscidae known in Chile, while Platyarthrus aiasensis and Trichorhina tomentosa are the first myrmecophilous species. On the other hand, although Porcellionides pruinosus and Armadillidium vulgare had already been recorded in the country, these are the first known locations in continental Chile. Finally, the exotic species Niambia capensis is cited for the first time in areas far from marine influence. Among the native isopods, only Chileoniscus marmoratus had previously been registered in the park. However, the presence of Benthanoides sp., Ligia novizelandiae and Tylos chilensis are confirmed, because these species have been previously reported in locations northern and southern the study area. The record of the family Dubioniscidae requires confirmation, as it is based on incomplete material. This work has allowed to add five species to the Chilean oniscofauna, so the diversity of the group would reach a total of 15 families, 27 genera and 50 species in the country.
Article
Full-text available
Alien species are a significant threat to natural ecosystems and human economies. Despite global efforts to address this challenge, the documented number of alien species is rapidly increasing worldwide. However, the magnitude of the impact of alien species may vary significantly across habitats. For example, some habitats are naturally less prone to biological invasions due to stringent abiotic and biotic characteristics, selecting for a limited number of introduced species possessing traits closely related to the native organisms. Subterranean ecosystems are quintessential examples of habitats with strong environmental filters (e.g. lack of light and scarcity of food), driving convergent adaptations in species that have successfully adapted to life in darkness. Despite these stringent environmental constraints, the number of records of alien species in subterranean ecosystems has increased in recent decades, but the relevant literature remains largely fragmented and mostly anecdotal. Therefore, even though caves are generally considered very fragile ecosystems, their susceptibility to impacts by alien species remains untested other than for some very specific cases. We provide the first systematic literature survey to synthesise available knowledge on alien species in subterranean ecosystems globally. This review is supported by a database summarising the available literature, aiming to identify gaps in the distribution and spread of alien invertebrate species in subterranean habitats, and laying the foundations for future management practices and interventions. First, we quantitatively assessed the current knowledge of alien species in subterranean ecosystems to shed light on broader questions about taxonomic biases, geographical patterns, modes of dispersal, pathways for introductions and potential impacts. Secondly, we collected species-specific traits for each recorded alien species and tested whether subterranean habitats act as ecological filters for their establishment, favouring organisms with pre-adaptive traits suitable for subterranean life. We found information on the presence of 246 subterranean alien species belonging to 18 different classes. The dominant alien species were invertebrates, especially insects and arachnids. Most species were reported in terrestrial subterranean habitats from all continents except Antarctica. Palaearctic and Nearctic biogeographic regions represented the main source of alien species. The main routes of introductions into the recipient country are linked to commercial activities (84.3% of cases for which there was information available). Negative impacts have been documented for a small number of case studies (22.7%), mostly related to increased competition with native species. For a limited number of case studies (6.1%), management strategies were reported but the effectiveness of these interventions has rarely been quantified. Accordingly, information on costs is very limited. Approximately half of the species in our database can be considered established in subterranean habitats. According to our results, the presence of suitable traits grants access to the stringent environmental filter posed by subterranean environments, facilitating establishment in the new habitat. We recommend that future studies deepen the understanding of invasiveness into subterranean habitats, raising public and scientific community awareness of preserving these fragile ecosystems.
Article
Full-text available
Subterranean ecosystems are among the most widespread environments on Earth, yet we still have poor knowledge of their biodiversity. To raise awareness of subterranean ecosystems, the essential services they provide, and their unique conservation challenges, 2021 and 2022 were designated International Years of Caves and Karst. As these ecosystems have traditionally been overlooked in global conservation agendas and multilateral agreements, a quantitative assessment of solution-based approaches to safeguard subterranean biota and associated habitats is timely. This assessment allows researchers and practitioners to understand the progress made and research needs in subterranean ecology and management. We conducted a systematic review of peer-reviewed and grey literature focused on subterranean ecosystems globally (terrestrial, freshwater, and saltwater systems), to quantify the available evidence-base for the effectiveness of conservation interventions. We selected 708 publications from the years 1964 to 2021 that discussed, recommended, or implemented 1,954 conservation interventions in subterranean ecosystems. We noted a steep increase in the number of studies from the 2000s while, surprisingly, the proportion of studies quantifying the impact of conservation interventions has steadily and significantly decreased in recent years. The effectiveness of 31% of conservation interventions has been tested statistically. We further highlight that 64% of the reported research occurred in the Palearctic and Nearctic biogeographic regions. Assessments of the effectiveness of conservation interventions were heavily biased towards indirect measures (monitoring and risk assessment), a limited sample of organisms (mostly arthropods and bats), and more accessible systems (terrestrial caves). Our results indicate that most conservation science in the field of subterranean biology does not apply a rigorous quantitative approach, resulting in sparse evidence for the effectiveness of interventions. This raises the important question of how to make conservation efforts more feasible to implement, cost-effective, and long-lasting. Although there is no single remedy, we propose a suite of potential solutions to focus our efforts better towards increasing statistical testing and stress the importance of standardising study reporting to facilitate meta-analytical exercises. We also provide a database summarising the available literature, which will help to build quantitative knowledge about interventions likely to yield the greatest impacts depending upon the subterranean species and habitats of interest. We view this as a starting point to shift away from the widespread tendency of recommending conservation interventions based on anecdotal and expert-based information rather than scientific evidence, without quantitatively testing their effectiveness.
Article
Full-text available
304 individual of isopods were collected for the period from June 2019 to September 2019 from different areas of Najaf Governorate. The results showed recording of three species of Isopoda: Porcellio Leavis, Porcellio scaber and Porcellio spinicornis for the first time in Najaf. The highest percentage of appearance of Porcellio Leavis was (40.8%), while the lowest percentage of P. spinicornis was (22.7%), due to the high temperature in the summer in which the samples were collected, as well as the different environments for collecting samples in terms of organic matter available in the soil.