Terrestrial Arthropod Surveys
on
Pagan Island, Northern Marianas
Neal L. Evenhuis, Lucius G. Eldredge, Keith T. Arakaki,
Darcy Oishi, Janis N. Garcia & William P. Haines
Pacific Biological Survey, Bishop Museum,
Honolulu, Hawaii 96817
Final Report
November 2010
Prepared for:
U.S. Fish and Wildlife Service,
Pacific Islands Fish & Wildlife Office
Honolulu, Hawaii
Evenhuis et al. — Pagan Island Arthropod Survey
BISHOP MUSEUM
The State Museum of Natural and Cultural History
1525 Bernice Street
Honolulu, Hawai’i 96817–2704, USA
Copyright© 2010 Bishop Museum
All Rights Reserved
Printed in the United States of America
Contribution No. 2010‐015 to the Pacific Biological Survey
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TABLE OF CONTENTS
Executive Summary ......................................................................................................... 5
Background ..................................................................................................................... 7
General History .............................................................................................................. 10
Previous Expeditions to Pagan Surveying Terrestrial Arthropods ................................ 12
Current Survey and List of Collecting Sites .................................................................. 18
Sampling Methods ......................................................................................................... 25
Survey Results ............................................................................................................... 26
Taxon Accounts ............................................................................................................. 28
Acarina ............................................................................................................... 28
Araneae .............................................................................................................. 28
Blattaria .............................................................................................................. 28
Chilopoda .......................................................................................................... 29
Coleoptera ........................................................................................................ 29
Collembola ........................................................................................................ 30
Dermaptera ....................................................................................................... 31
Diplopoda .......................................................................................................... 32
Diptera ............................................................................................................... 32
Embiida .............................................................................................................. 33
Hemiptera .......................................................................................................... 33
Homoptera ......................................................................................................... 34
Hymenoptera ...................................................................................................... 34
Isopoda .............................................................................................................. 36
Isoptera .............................................................................................................. 37
Lepidoptera ........................................................................................................ 37
Mantodea .......................................................................................................... 39
Neuroptera ........................................................................................................ 40
Odonata ............................................................................................................. 40
Orthoptera .......................................................................................................... 40
Phasmatodea ...................................................................................................... 41
Pseudoscorpionida ............................................................................................. 42
Psocoptera ......................................................................................................... 42
Scorpionida ....................................................................................................... 42
Siphonaptera ..................................................................................................... 43
Thysanoptera ..................................................................................................... 43
Thysanura ........................................................................................................... 43
Significant Findings ...................................................................................................... 44
Threats to Fruit Bats or the Megapode Fauna ................................................................. 46
Acknowledgements ....................................................................................................... 46
Literature Cited ............................................................................................................. 46
Appendices .................................................................................................................... 49
Appendix I. Bibliography of Pagan Terrestrial Arthropods ............................... 50
Appendix II. Checklist of Pagan Arthropods ..................................................... 55
Appendix III. Cory Campora Field Notes ......................................................... 66
Evenhuis et al. — Pagan Island Arthropod Survey
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EXECUTIVE SUMMARY
This report is part of the “Marianas Expedition Wildlife Survey 2010” (MEWS 2010), a
U.S. Fish & Wildlife Service (USFWS) project funded by the Department of Defense U.S. Marines and is tasked to gather natural resource information on fish and wildlife in
the Mariana Islands. This information is required by federal regulations to properly
determine the potential impacts that will occur due to the shifting of significant military
resources from Okinawa to the Territory of Guam and the Commonwealth of the
Northern Mariana Islands. As part of this military build-up, Pagan is under consideration
as a live fire training area.
This report gives the results of a terrestrial arthropod survey conducted by USFWS and
NAVPAC personnel on Pagan in the month of July 2010 and gives details on significant
findings resulting from material collected on that survey. In addition, a full checklist of
terrestrial arthropods known from Pagan Island is given based on the current survey
material as well as previous published records. An appendix gives a full bibliography of
articles dealing with Pagan arthropods.
Staff from the U.S. Fish and Wildlife Service, Pacific Islands Office, and the U.S. Navy
conducted collections of terrestrial arthropods from 9-21 July 2010 using a variety of
collecting methods including Malaise flight intercept traps, yellow water pan traps, pitfall
traps, peanut butter traps for ants, aerial sweep netting, aquatic dip netting, aspirating, and
hand collections.
Thousands of terrestrial arthropod specimens resulted from these collections, which were
sorted by the collectors to order (to class for non-insects) and delivered to the Bishop
Museum for identification. A team of entomologists at the Bishop Museum identified a
total of 288 different taxa of terrestrial arthropods based on the survey, which included
228 new island records for Pagan (doubling the number of arthropods previously
recorded from Pagan) bringing the total number of terrestrial arthropod species known
from Pagan to 416. The full list of identified arthropods is given in Appendix II and
includes all previously published terrestrial arthropod records for Pagan as well as the
new records identified during this study.
To put the arthropod fauna into a proper historical context in order to better understand
their possible biological status on the island (e.g., endemic, native, nonindigenous), a
history of human habitation as well as previous collecting expeditions is given. The vast
majority of identified terrestrial arthropods are most likely nonindigenous, having arrived
on Pagan via a variety of mechanisms including transport by humans, supply shipments,
and commerce. Verification of true status requires study of the known distributions and
potential vagility of each species, which was outside the scope of this report.
Although the island has undergone numerous geophysical and human land use changes
resulting in what we are calling a synanthropic arthropod fauna, there are still pockets of
native arthropods that survive. Eight endemic species are recorded, three of them are new
to science. In addition, one new genus was found.
Evenhuis et al. — Pagan Island Arthropod Survey
The littoral zone has been a neglected area for previous collectors, and a number of new
marine and littoral faunal records have resulted from collecting during this survey.
None of the arthropods identified are any threat to the megapode or fruit bat populations
on the island. The crazy ant, Anoplolepis gracilipes, may pose a potential threat to food
resources of the bat and megapodes if the populations on the island ever form what are
called supercolonies. Their population levels now are large but not dangerous.
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BACKGROUND
Pagan Island (Map 1) is the largest and most complex biologically and geographically
among the northern islands of the Commonwealth of the Northern Mariana Islands
(CNMI). The island was named “Pagan” by Jesuits who traveled through the Marianas in
the 1600s. The name is one of the Christian names they gave to many of the islands in the
Northern Marianas. Since its first visitation by western Europeans in 1695, Pagan has
gone by many names including the following: Agan, Ile de Saint Ignace, Pagaon, Pagara,
Pagon, Paygan, Pragan, Prajan, Pemplie de Volcans, Remplie de Volcaus, San Ignace,
and Saint Ignace (Bryan, 1971).
It is an active volcanic island (18.10°N 145.76°E) approximately 320 kilometers
north of Saipan. It contains three volcanic cones, the highest of which and most active,
Mount Pagan in the northern part of the island, is 569.9 m [1870 ft] and whose last major
eruption was in 1981, which had ash deposits and lava flows that covered or partially
covered the village buildings and airfield and affected the shorelines of the two lakes in
the northwestern portion of the island. A smaller eruption in 2006 deposited ash in the
surrounding areas. The other two volcanoes are located in the southern portion of the
island and last erupted approximately 150 years ago.
Fig. 1. Pagan viewing south, showing Lake Sanhiyon in foreground,
separated from ocean by thick sand. Photo: Dan Polhemus.
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There are two natural lakes on the island, both in the north part and both containing
brackish water. They were formed approximately 200 years ago (Asakura et al., 1994).
The outer lake (Laguna Sanhiyon) [Fig. 1] is separated from the ocean by an
approximately 50-meter wide berm of sand. The inner lake (Laguna Sanhalom) [Fig. 2] at
one time had a hot mineral spring (see newspaper feature article by Ronck, 1975 for
photos and description), the surface portion of which disappeared after the eruption of
Mount Pagan in 1981 (warm water still percolates into the lake from below the surface).
Trusdell (2009) gives a concise summary of the geology of the island.
Fig. 2. Lake Sanhalom, Northern Pagan. Photo: Dan Polhemus.
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GENERAL HISTORY
Because changes in ecosystems due to anthropogenic factors play a large part in the
constituency of the vegetative and zoological composition of an island, we here present a
short history of habitation and human activities on Pagan.
Human habitation has been intermittent on the island, owing primarily to factors
such as volcanic eruptions and various commercial interests over time. Archeological
surveys and linguistic research suggest that the Marianas were first colonized about 3500
years ago. Little archeology has been done on Pagan owing to few good sites remaining
after volcanic eruptions and poor dating sources (e.g., there is no clay and pottery had to
be imported from the southern islands) (Russell, 1998). Despite this, research on a site on
the eastern coast of Pagan suggested it was occupied by 1300 C.E. (Egami & Saito,
1973).
After the Spanish explorer Magellan sailed through the area in 1521 on his
famous circumnavigation of the world, Spain declared the archipelago as a royal
possession. However, for the next 140 years the islands were virtually neglected as
Spanish activities there were few. In 1668, a Jesuit mission was established on Guam and
not long after its founding, priests began traveling northward from island to island in their
attempts to convert the native Chamorros to Christianity. Seeing that the remote ventures
to the islands to the north of their Guam base was becoming ineffectual in procuring
efficient conversions, the Spanish began a program called the reducción, which relocated
residents scattered on all the northern islands to one island: Saipan. The residents of
Pagan were relocated to Saipan in 1697 and, except for the rare exploratory expeditions
of the 19th century that sailed in to the area to make hydrographic studies, the island
slipped into obscurity once again.
In 1865 a serious attempt to re-introduce residents to Pagan was initiated by
American George Johnson, who brought 265 Carolinians to Pagan to produce copra. This
adventure was short-lived, though, as all the Pagan residents were sent to Saipan in 1869.
The reasons are unclear as to why the copra production was abandoned on Pagan but they
could well have been economic (Russell, 1998) since the northern islands were too far
removed from the economic center of Guam to sustain commercially viable operations.
In 1899 Germany purchased from Spain all the islands north of Guam, and their
administration was headquartered on Saipan. The 1899 census by the Germans gave the
population of Pagan then as 75 (Spennemann, 1999). In May 1901, Governor G. Fritz
visited the northern islands and made observations on the general geology and volcanic
activity (Fritz, 1902). Further observations, including those on the fauna and flora, were
made by the Czech zoologist and parasitologist Stanislaus Prowazek (1913).
World War I saw a changeover of administration of the islands from Germany to
the Japanese. In 1914, the islands of the northern Marianas became part of the Japanese
Mandated South Seas Islands through action of the League of Nations. A number of
Japanese scientists made expeditions during the 1930s and published accounts of the flora
(e.g., Hosokawa, 1934), submarine topography (Tayama, 1936), volcanic geology
(Tanakadate, 1940) as well as insects (see below for more details). During World War II,
the Northern Marianas were of little strategic or tactical value except Pagan. The
Japanese constructed an airfield on Pagan as well as associated troop barracks, storage
bunkers for bombs and fuel, and pillboxes and air-raid shelters. Manning this base were
Evenhuis et al. — Pagan Island Arthropod Survey
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over 2000 Japanese troops (Richard, 1957). After the surrender of the Japanese garrison
in September 1945, the Japanese were returned home and the surviving Chamorros were
all taken to Saipan due to extreme food shortages.
After World War II, the islands of the Marianas were administrated by the United
States Trust Territory. Several field projects were carried out on the fauna and flora; and
the military funded research on the geology. A small military contingent remained on the
island to maintain the airfield (Corwin et al., 1957).
In 1948, Chamorros and Carolinians decided to start up commercial operations in
the Northern Marianas again; and in 1951, 57 Chamorros were brought to Pagan to
produce copra. Copra production waxed and waned and eventually was abandoned
altogether by the 1970s; the remaining residents were maintained with supplies brought
in from government-run “field trip” ships a few times a year. The volcanic eruption of
Mount Pagan in 1981 forced the 53 residents of the island to flee to Saipan. The resultant
lava flows and ash destroyed much of the vegetation of the northern part of the island and
covered the airfield. The recent 2006 eruption again caused a significant amount of ash
and lava to cover the northern portion of the island so that there is a conspicuous lack of
vegetation and associated life forms in the immediate vicinity of the volcano.
Evenhuis et al. — Pagan Island Arthropod Survey
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PREVIOUS EXPEDITIONS TO PAGAN SURVEYING
TERRESTRIAL ARTHROPODS
Little attention has been paid to Pagan with respect to the island’s terrestrial invertebrate
fauna. Most explorers who had visited Pagan had either looked upon it from the ship and
made notes to charts, or upon landing, made only ethnological and geological
observations or commented on the vertebrate fauna [large mammals such as pigs having
been introduced to the island not long after Magellan’s sail through the island chain in the
1500s (Rodda, 2009)].
Fig. 3. The first insect collector on Pagan. “Mariano” who accompanied
Alfred Marche on his trip there in December 1887 (from Marche, 1982).
Evenhuis et al. — Pagan Island Arthropod Survey
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The first collection of insects was apparently made in 1887 by Antoine-Alfred
Marche (1844–1898), a naturalist and ethnologist at the Musée de l’Homme in Paris. He
had previously made trips to Africa, the Philippines, and Guam to do ethnology research
and had published accounts of his travels there. His trip to the Marianas in 1887–1888
was to be his last. In Manila shortly after arriving, he hired a collector, Mariano (Fig. 3)
and a cook and headed north to stop at various of the Mariana Islands. His account of his
December 1887 stay on Pagan during his trip through the Marianas [translated into
English in 1982 from the original (Marche, 1891) in a little-known French journal] gives
a detailed description of the island and is accompanied by the first known photograph of
Mount Pagan (Fig. 4) however, he says little of the biology of Pagan (Marche, 1982: 20):
“On Pagan, there are only a few Carolinians settled there to harvest coconuts, which trade is
carried on by Captain William. Hunting gave me meager results; birds are scarce; before the great
typhoon of 1884, they were in far greater number. No mammal, except for some pigs and wild
goats. Fresh water appears to be completely lacking on this island.”
He concluded his report of Pagan by giving an account of the material he collected
(Marche, 1982: 22):
“Upon my return to Guam, I prepared and sent my second shipment which contained eleven
objects for the Ethnographic Museum, about 200 skinned birds, 450 mollusks in alcohol, 300
skins, plus 500 insects in paper and in alcohol and about 100 species of plants with flowers and
fruits.”
Fig. 4. First known photograph of Mount Pagan, taken by Alfred Marche in 1887 (from Marche, 1982).
Evenhuis et al. — Pagan Island Arthropod Survey
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Although some of the birds he collected were studied by Oustalet (1895–1896), no one
has apparently bothered to trace the current whereabouts or existence of his insect
collections, which may still be in the Musée de l’Homme in Paris.
After Germany purchased the Marianas from Spain and began their governance,
two surveys were conducted that made natural history observations: Fritz (1902) and
Prowazek (1913). The latter gave a detailed report on the history, culture, and fauna and
flora of the Marianas but gave sketchy remarks concerning arthropods. Prowazek (1913)
discussed a number of species as occurring in the Marianas but does not specifically list
species occurring on Pagan.
When the Japanese administered the Pacific islands acquired through the League
of Nations, scientific expeditions were made in the 1930s to survey the natural history of
their newly acquired possessions in the Pacific. One of these was the Esaki Micronesian
Expeditions from 1936–1940, which resulted in a number of publications by specialists
including descriptions of many new species and the first published records of arthropods
from Pagan. Professor Teiso Esaki, entomologist at Fukuoka University, was in charge of
entomological investigations for the Japanese administration of Micronesia before World
War II. Almost 80 publications were generated from these surveys, although most dealt
with islands other than the northern Marianas.
As part of these expeditions, the first collecting trip specifically to Pagan was
made in April 1940 by Fukuoka University entomologist Keizo Yasumatsu (Fig. 5) and
scientific records of arthropods from that trip were published in Yasumatsu (1940). It
included few records as the publication was only meant to be a short narrative of the
collecting trip. However, many other publications by Japanese colleagues on Pagan
arthropods soon followed, mostly basing their results on material collected by Yasumatsu
and his student Seiichiro Yoshimura during this trip in April 1940.
Fig. 5. Esaki Micronesian Expedition to Pagan, April 1940. Keizo Yasumatsu, center
(from Yasumatsu, 1940).
Evenhuis et al. — Pagan Island Arthropod Survey
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In 1946, shortly after the end of World War II, the Pacific Science Board of the
U.S. National Research Council was established. It was set up to “aid the scientists of
America who wish to engage in scientific investigations for which there is a need in the
Pacific area, to advise governmental and other agencies on scientific matters pertaining to
the Pacific, and to further international cooperation in the field of Pacific science.” The
Pacific Science Board cooperated with the U.S. Navy in investigating certain problems
that both felt were a priority for the Pacific area. One of these was finding ways to control
insect and related pests in Micronesia. The Insect Control Committee for Micronesia
(ISSM) was thus established and its priority research areas included research on the
following pests: (1) the giant African snail; (2) the rhinoceros beetle; (3) the banana root
borer; and (4) the Saipan coconut beetle (Bryan, 1949). The ICCM employed a staff
entomologist, Daniel B. Langford, who made trips to Micronesian islands from October
1947 to October 1949. He only visited Pagan on a trip from 5–18 May 1948 that also
included the islands of Anatahan, Alamagan, and Agrihan. It is not known how long he
spent on Pagan and what if any arthropods he may have collected.
In 1953, the entomological work of the Pacific Science Board and the office of
Naval Research were taken over by the Bishop Museum in Honolulu and with the
funding assistance of two grants from the National Science Foundation, the series Insects
of Micronesia was begun (Gressitt, 1954). The hundreds of resulting articles in 19
volumes offer to date the most detailed taxonomic accounts of insects occurring in
Micronesia, and many of these include records and descriptions of arthropods from
Pagan. The series was discontinued as a separate series by Bishop Museum in the 1990s
but was turned over to the University of Guam, which continues to publish articles in the
series within their journal Micronesica; the last of which was on lauxaniids (Sasakawa,
2009).
As part of the U.S. Army Corps of Engineers’ “Post Hostilities Mapping
Program” a geological survey was conducted of Pagan in the 1950s, the results of which
were published in Corwin et al. (1957). Along with the detailed geological report and
mapping, plants were listed and animals observed and discussed in the narrative.
However, remarks on arthropods were brief:
“More than a hundred species of insects, several spiders, two scorpions, two isopods, and a few
worms, centipedes, and millipedes have been collected. Of these only a few may be classed as
pests. Two species of flies are common and very annoying. Cockroaches and large beetles may do
some damage to supplies. Mosquitoes are chiefly nocturnal varieties and are harmless. Stinging
wasps are numerous in many groves of Casuarina trees. Although rarely fatal, the bites of the
scorpions and one large variety of centipede may cause considerable discomfort” (Corwin et al.,
1957: 109).
The Natural History Museum and Institute in Chiba, Japan conducted a biological
expedition to the northern Marianas in 1992 and results were published in a single
volume (Asakura & Furuki, 1994). This was the last formal expedition to Pagan to survey
arthropods prior to the current survey. The volume contains dozens of articles on the
results of this expedition and contains a checklist (Miyano, 1994) that forms the basis for
the current checklist in this report.
Since then, Pagan has only had sporadic visits primarily for floristic or vertebrate
surveys, or for geological concerns; with little work done on arthropods. One report
Evenhuis et al. — Pagan Island Arthropod Survey
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(Cruz et al., 2000) mentions insects being collected opportunistically on Pagan but no
specific records were listed in the report.
Lake
Sanhalom
Lake
Sanhiyon
Mt Pagan
Northern Pagan
Bandeera
Peninsula
Main
Camp
Tr a
n
e
Trans
Miari Cliffs
s ec
t1
ct 3
Apansanmeena
0
Collection Areas
and Trapping Locations
for Northern Pagan,
Pagan, CNMI
9-21 July 2010
0.5
1 Kilometers
collection_areas
malaise, pitfall, pan, UV light
malaise, pitfall, pan
GCS_WGS_1984
Datum: D_WGS_1984
Pagan
CNMI
1:40,000
UV light only
Map 2. Collection sites on northern Pagan, 9-21 July 2010.
Colored dots are sites where traps were set.
Yellow lines indicate collecting while hiking.
Evenhuis et al. — Pagan Island Arthropod Survey
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Middle Crater
Boat
Landing
Main
Camp
Southern Pagan
Lower
Shelf
Tr an
s
ect
Upper
Plateau
Tr an
s
ect 1
1
10
0
Collection Areas
and Trapping Locations
for Southern Pagan,
Pagan, CNMI
13-14 and
16-17 July 2010
collection_areas
malaise, pitfall, pan, UV light
malaise only
0.5
GCS_WGS_1984
Datum: D_WGS_1984
1 Kilometers
Pagan
CNMI
1:25,000
UV light only
Map 3. Collection sites on southern Pagan, 13–17 July 2010.
Dots are sites where traps were set.
Yellow lines indicate collecting while hiking.
Evenhuis et al. — Pagan Island Arthropod Survey
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CURRENT SURVEY AND LIST OF COLLECTING SITES
The current survey was conducted by three NAVPAC personnel (Cory Campora, Stephan
Lee, Justin Fujimoto) and USFWS staff members (Christa Russell, Mike Richardson)
from 6 to 21 July. A few specimens made prior to the main arthropod survey (collected
by E. Wosh on 25 June [during a herpetological survey] on south Pagan) were also
included in the identifications. Additionally, Dan Polhemus of USWFS (not a part of this
survey contract) made a separate survey to assess the aquatic insect fauna on Pagan and
the results of his survey are incorporated herein. Specimens were collected by hand and
sweep net from a variety of localities, usually while hiking from one locality to another.
However, the majority of specimens were collected through various trapping methods,
which were set up at specific localities and these are indicated below. A total of 10 sites
employed various trapping methods and are detailed below.
LIST OF COLLECTING SITES
Site 1 [= Campora field notes site 4] [Fig. 6]
Northern Pagan, Main Camp (old Shomushon village), near runway, 9 July 2010 and 16
July. Collectors: Mike Richardson, Christa Russell, Stephan Lee, Justin Fujimoto, Cory
Campora. [GPS: 18.12346°N, 145.760640°E]
Traps:
Malaise: 1 trap (9–10; 16–17 July collecting period)
Pitfall traps: 5 traps (9–10; 16–17 July collecting period)
Blacklight: 1 trap (night of 9, 16 July collecting period)
Additional collecting methods employed: general collecting.
Site 2 [= Campora field notes site 6] [Fig. 7]
Northern Pagan, Lake Sanhalom (west side), 10 July 2010. Collectors: Mike Richardson,
Christa Russell, Stephan Lee, Justin Fujimoto, Cory Campora. [GPS: 18.14814°N,
145.771950°E]
Traps:
Malaise: 1 trap (10–11 July collecting period)
Pitfall: 5 traps (10–11 July collecting period)
Pan traps: 5 traps (10–11 collecting period)
Additional collecting methods employed: aerial sweep netting, general collecting, aquatic
dip netting.
Evenhuis et al. — Pagan Island Arthropod Survey
Fig. 6. View of Collection Site 1: Main camp near Shomushon village, looking north.
Fig. 7. Collection Site 2: Lake Sanhalom, southwest side, showing Malaise trap and yellow pan traps
19
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Site 3 [= Campora site 10] [Fig. 8]
Isthmus on Pagan, Trail along coast from main camp (megapode transect #3). 12 July.
Collectors: Mike Richardson, Christa Russell, Stephan Lee, Justin Fujimoto, Cory
Campora. [GPS: 18.11891°N, 145.759970°E]
Traps:
Malaise: 1 trap (12–13 July collecting period)
Pitfall: 5 traps (12–13 July collecting period)
Pan traps: 5 traps (12–13 collecting period)
Additional collecting methods employed: general collecting
Fig. 8. Collection site 3. Megapode transect #3 along west coast south from Main Camp,
showing Malaise trap set up.
Site 4 [= Campora site 12] [Fig. 9]
Southern Pagan, South Camp area, southern end of island (megapode transect #10). 13
July 2010. Collectors: Mike Richardson, Christa Russell, Stephan Lee, Justin Fujimoto,
Cory Campora. [GPS: 18.064313°N, 145.715226°E]
Traps:
Malaise: 1 trap (13–14 July collecting period)
Pitfall traps: 5 traps (13–14 July collecting period)
Blacklight: 1 trap (night of 13 July collecting period)
Additional collecting methods employed: general collecting, aerial sweep netting.
Evenhuis et al. — Pagan Island Arthropod Survey
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Fig. 9. Collection Site 4. Southern Pagan camp site. Traps were set nearby.
Site 5 [= Campora site 16] [Fig. 10]
Northern Pagan, Bandeera Peninsula. 15 July 2010. Collectors: Mike Richardson, Christa
Russell, Stephan Lee, Justin Fujimoto, Cory Campora. [GPS: 18.126670°N,
145.756860°E]
Traps:
Malaise: 1 trap (15–16 July collecting period)
Pitfall: 5 traps (15–16 July collecting period)
Pan traps: 5 traps (15–16 collecting period)
Blacklight: 1 trap (night of 15 July collecting period)
Additional collecting methods employed: none.
Evenhuis et al. — Pagan Island Arthropod Survey
Fig. 10. Collection Site 5. Top of Bandeera Peninsula, looking east toward an active Mount Pagan,
showing Malaise trap in place.
Site 6
Southern Pagan, Main camp. 15–16 July 2010. Collectors: Mike Richardson, Christa
Russell, Stephan Lee, Justin Fujimoto, Cory Campora. [GPS: 18.065892°N,
145.714867°E]
Traps:
Blacklight: 1 trap (night of 15 July collecting period)
Additional collecting methods employed: none.
Site 7 [Fig. 11]
Southern Pagan, Middle crater. 15–16 July 2010. Collectors: Mike Richardson, Christa
Russell, Stephan Lee, Justin Fujimoto, Cory Campora. [GPS: 18.073481°N,
145.725095°E]
Traps:
Malaise: 1 trap (15–16 July collecting period)
Pitfall: 5 traps (15–16 July collecting period)
Additional collecting methods employed: aerial netting off of vegetation
22
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Fig. 11. Collection Site 7. Southern Pagan, middle crater, showing Malaise trap in place.
Site 8
Northern Pagan, south of Lake Sanhiyon, Ironwood forest. 17–19 July 2010. Collectors:
Mike Richardson, Stephan Lee, Justin Fujimoto. [GPS: 18.137694°N, 145.76903°E]
Traps:
Malaise: 1 trap (17–19 July collecting period)
Pan traps: 5 traps (17–19 collecting period)
Pitfall traps: 5 traps (17–19 July collecting period)
Additional collecting methods employed: aerial netting off of vegetation.
Site 9
Northern Pagan, Somushon village, north of motor pool. 17–18 July 2010. Collectors:
Mike Richardson, Stephan Lee, Justin Fujimoto. [GPS: 18.127526°N, 145.762736°E]
Traps:
Malaise: 1 trap (17–18 July collecting period)
Pan traps: 5 traps (17–18 collecting period)
Additional collecting methods employed: aerial netting off of vegetation.
Evenhuis et al. — Pagan Island Arthropod Survey
Site 10 [Fig. 12]
Eastern central Pagan, coconut grove. 20–21 July 2010. Collectors: Mike Richardson,
Stephan Lee, Justin Fujimoto. [GPS: 18.112823°N, 145.785874°E]
Traps:
Malaise trap: 1 trap (20-21 July collecting period)
Pan traps: 5 traps (20–21 collecting period)
Additional collecting methods employed: aerial netting off of vegetation.
Fig. 12. Collection Site 10. Coconut grove on east side of island, showing Malaise trap in place.
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SAMPLING METHODS
Collecting on Pagan was done with Malaise trapping, yellow water pan trapping, pitfall
traps, ultra-violet light traps, and aerial sweep nets. Malaise traps, pan traps and pitfall
traps were set up in various locations (see Maps 2, 3) to allow sampling in various
vegetative habitats. The ultra-violet light trap was set up near the main camp to allow
easy access during evening collecting. Aerial sweep netting and hand collecting were
conducted along transects and trails as indicated in yellow on the maps.
Descriptions of some of the collecting methods employed during this survey
include the following:
1. Malaise traps—Malaise traps are among the most productive samplers in terms
of species richness and number of specimens captured. Flying insects approach the
central panel of the trap and attempt to avoid it either by dropping to the ground or flying
upward. Those flying upward (the vast majority) are funneled into the collecting head of
the trap where they are killed. The collecting head is attached to the higher end of the
trap. Insects caught at the top, are killed by drowning in a collecting canister of 95%
ethanol. Those dropping to the ground can be collected in water pan traps.
2. Water pan traps—Shallow plastic pans (often yellow or white) are placed on
the ground or in trees to sample arthropods. The pans are filled with water and a
surfactant (soap solution) to allow trapped specimens to sink to the bottom of the pan and
drown. They are attractive to flying insects in both open areas and forest canopy.
3. Pitfall traps—These consist of small plastic cups placed into the ground with
the rim level with the surface. Pitfall traps collect most ground dwelling arthropods either
through adventive encounters to the trap or by baits. Arthropods fall into the trap and a
propylene glycol solution in the bottom of the cup kills the trapped arthropods.
4. Aerial nets—Aerial sweep nets with a fine mesh to collect smaller Diptera and
Hymenoptera are used to sample arthropods on vegetation, leaf litter, littoral habits,
beaches, and rocky intertidal reefs.
5. Hand collecting—Hand collecting, with or without the use of an aspirator, are
used for collecting arthropods on specific species of plants and to collect arthropods that
are difficult to collect using other techniques—such as those on craggy rock faces, in
small holes and crevices or on muddy substrates.
6. UV light traps—Ultra-violet (UV) light traps are used to collect flying insects
at night. Insects are attracted to the ultra-violet light reflecting off of a white sheet and are
collected off the sheet by hand or aspirator. These are especially good for collecting night
flying Lepidoptera (moths) as well as certain beetles and some aquatic Diptera.
7. Peanut butter bait—Spreading peanut butter on sticks or cards as a bait attracts
both sugar-loving and oil-loving ants, which can easily be transferred to collecting
containers. Peanut butter baiting was used sporadically during the survey.
Evenhuis et al. — Pagan Island Arthropod Survey
26
SURVEY RESULTS
Out of thousands of specimens collected during this survey, some 288 species of
terrestrial arthropods were identified [see quick summary in Table 1]. Previous to this
survey, 188 species of arthropods were published as occurring on Pagan Island. The
current survey brings the total known species to 416 and includes 228 new records to the
island. A full list of arthropods previously known from the literature from Pagan and
those identified during this survey is given in Appendix II.
Table 1. Quick Summary of Results of Identifications by Order
(orders in red are new order or class records to Pagan)
order or class
Acarina (mites, ticks)
Araneae (spiders)
Blattaria (roaches)
Chilopoda (centipedes)
Coleoptera (beetles)
Collembola (springtails)
Dermaptera (earwigs)
Diplopoda (millipedes)
Diptera (flies, gnats, midges)
Embiida (web spinners)
Hemiptera (true bugs)
Homoptera (aphids, scale insects)
Hymenoptera (wasps, bees, ants)
Isopoda (sow bugs, pill bugs)
Isoptera (termites)
Lepidoptera (moths, butterflies)
Mantodea (mantids)
Neuroptera (lacewings)
Odonata (dragonflies, damselflies)
Orthoptera (crickets, grasshoppers)
Phasmatodea (walking sticks)
Pseudoscorpionida (pseudoscorpions)
Psocoptera (book lice, bark lice)
Scorpionida (scorpions)
Siphonaptera (fleas)
Thysanoptera (thrips)
Thysanura (bristletails, silverfish)
totals
prev
12
11
5
1
23
4
0
0
21
0
20
16
42
2
0
10
1
3
4
3
1
3
1
2
1
2
0
this
3
36
4
2
41
6
2
2
69
1
12
7
47
1
1
17
1
8
3
14
0
2
8
1
0
2
1
total
15
43
7
2
57
9
2
2
82
1
31
20
69
2
1
22
1
9
4
15
1
5
9
2
1
4
1
new
records
3
33
2
1
34
5
2
2
61
1
10
5
25
0
1
12
0
6
0
12
0
2
8
0
0
2
1
188
288
416
228
The current survey lists 5 new order records (Dermaptera, Diplopoda, Embiida, Isoptera,
and Thysanura), 99 new family records, and 228 new species records for Pagan. We were
able to identify most all of the material collected except for groups that entailed special
preparation and taxonomic expertise (e.g., Acarina; Collembola; Homoptera [Coccoidea];
Evenhuis et al. — Pagan Island Arthropod Survey
27
Thysanoptera). However, even in those groups, we were able to identify at least to family
the larger and better-known taxa. Brief overviews of the orders of arthropods encountered
are given below with reference made to the more common or interesting taxa. Significant
finds are described in more detail in the next section.
Pagan shows a fairly normally composed arthropod community with most
elements represented that would be expected given the amount of human activity that has
taken place over its history; thus the island’s fauna can be said to be primarily
synanthropic with pockets of native populations still surviving. A fairly high proportion
of the arthropod fauna are introduced or are native elements of widespread species.
Despite the large number of introduced taxa, it is significant that there are 8 endemic
arthropods that are known only from Pagan (5 previously described; 3 new undescribed
endemic taxa found during this survey). Further study is needed to determine the relative
health of these populations. A few of these previously recorded endemic species were recollected during this survey, indicating that at least some still survive despite the humaninduced perturbations and volcanic activity.
Some species previously recorded were not found in this survey and, in some
cases, it could be a result of land use changes that currently do not support resources that
would have otherwise been there to allow a organism to survive; or, more probable, that
sampling was not done in enough areas to be able to re-collect these taxa. In addition, soil
sampling and use of Berlese or Tullgren funnels were not employed on this project,
which can help with obtaining microscopic soil and ground dwelling organisms such as
pseudoscorpions, mites, springtails, and small beetles and immature stages of many
holometabolous orders of insects.
A note on place-names
Many of the names on the labels of material collected by Yasumatsu in 1940 and
published in subsequent papers by Japanese colleagues as well as other scientists are no
longer used and their current equivalents are difficult to decipher since no one has tried to
trace them previously and some localities do not exist any longer. Based on a manuscript
list of names provided to one of us (LGE) by a Pagan resident in the 1970s and the report
by Pangelinan & Kapileo (1970) comparing with the Japanese transliterations of these
names by Yasumatsu ands others, we here give a list of a few of the old names used by
Yasumatsu and their current equivalents in the hopes it will help pinpoint collecting
localities for future workers.
Old Yasumatsu Name
Song-Song
Laguna
Current Equivalent
Shomushon
Laguna Sanhiyon
Regusa
Tarague, Darage
Regusa, Rugusa
Degusa
Taragie, Tarague
Males, Malasu
Maras, Marasu
Abansantate
Apasanmeena
General location (all North Pagan)
Main village
outer lake; also name of the
village there
location on eastern coast; site
of old Japanese hospital
old village on N coast;
archaeological site
location on NW coast between
Laguna Sanhiyon and Tarague
location near isthmus
Evenhuis et al. — Pagan Island Arthropod Survey
28
TAXON ACCOUNTS
ACARINA (mites, ticks) [Fig. 13]
Few mites were collected in this survey, with those
recovered being primarily in the pitfall traps. Only the
ixodid (unverified identification but probably represents
the cattle tick Boophilus microplus), the bee mite
(Pyemotes sp.; cf. Fig. 13) and an oribatid soil mite were
recorded in this survey. Previous surveys (which
involved mite specialists) recorded 12 species from
Pagan. Further specialized collecting including soil
samples and Berlese funnels and identification of
properly prepared specimens should identify many more
acari from the island. The Pyemotes mite was collected
from a tenebrionid beetle, which had 30 or so specimens
attached to its body. The mites were attached to the body
under the wings and covered the thoracic and abdominal
region.
Fig. 13. Pyemotes bee mite. New
Island Record. Photo: Wikipedia.
ARANEAE (spiders) [Fig. 14]
Few spiders had been previously recorded from Pagan. The only publication on spiders
from Pagan (Yoshida et al., 1996) listed 11 species in 8 families. Collections from this
survey significantly increased the known spider fauna with 33 new species records of
taxa from the island bringing the number of species known from Pagan to 43 in 22
families. Most are common widespread species in the Pacific. One conspicuous spider,
the common cane spider, Heteropoda sp. (Fig. 14) was found in numbers under fallen
logs in a mixed forest in north Pagan. These spiders are free-living predators that do not
spin webs. They naturally occur on the forest floor but sometimes enter buildings. They
usually prey on ground dwelling insects such as cockroaches, crickets, and silverfish.
BLATTARIA (roaches)
Previous to this survey, five roaches had been recorded from Pagan. All are synanthropic
species that are widespread with humans and commerce. This survey only found three
species with one new record, the widespread Pacific beetle roach, Diploptera dysticoides.
Unlike other roaches, beetle roaches are not normally found in buildings but instead
prefer leaf litter and soil.
Evenhuis et al. — Pagan Island Arthropod Survey
29
Fig. 14. Heteropoda sp. (Sparassidae) flushed out from under a log on
Megapode Transect #3. New Island Record. Photo: Cory Campora.
CHILOPODA (centipedes)
Centipedes are fairly ubiquitous in faunal surveys and are another component of a
synanthropic fauna. Previous narratives of Pagan mention “centipedes” (e.g., Corwin,
1957); and two taxa were recovered in this survey: an undetermined member of the
Geophilomorpha, and a Scolopendra sp., a genus that contains some of the larger taxa of
this group of arthropods. Scolopendra are commonly transported in shipping from island
to island throughout the Pacific.
COLEOPTERA (beetles) [Fig. 15]
Beetles are among the most diverse groups of insects in the world and as such also are
among the most diverse in faunal surveys of specific areas such as islands. Previous to
this survey, 23 species were recorded from Pagan, all fairly widespread species. We
identified 41 taxa collected on this survey, with 34 new records for the island, bringing
the total number of beetles known from Pagan to 57. Of these, the following are new
family records to the island: Aderidae, Anthribidae, Bostrichidae, Cleridae,
Corylophidae, Laemophloeidae, Melyridae, Mordellidae, Nitidulidae, Platypodidae,
Salpingidae, Silvanidae, and Staphylinidae. The vast majority of species identified during
this survey are nonidigenous taxa. Some of the coccinellid species listed were probably
Evenhuis et al. — Pagan Island Arthropod Survey
30
introduced at one time to control aphids on crops (the beetle immatures are predaceous on
aphids) when commercial agriculture was prevalent on the island.
Fig. 15. The long-horned beetle, Cersium unicolor unicolor (Cerambycidae). Photo: Darcy Oishi.
COLLEMBOLA (springtails) [Fig. 16]
Springtails are often neglected in faunal surveys due to their small size but they are a
major component of any ecosystem and are a potentially significant and biodiverse group
of arthropods. Previous to this survey, only 4 species were recorded from Pagan
including one endemic species (Sira fuscana Uchida, 1944) [Fig. 15]. Pan and pitfall
traps recovered species of Collembola during this survey, and we were able to identify 6
species (including the endemic Sira fuscana) and confirm 5 new island records. Time
needed to properly slide mount and key species was not enough to allow us to identify
material to any finer resolution than what is represented in Appendix II, but we expect
with future research and collecting, especially soil sampling utilizing Berlese funnels to
extract arthropods, there will be additional new records for the island.
Evenhuis et al. — Pagan Island Arthropod Survey
31
Fig. 16. Sira fuscana Uchida, an endemic springtail from Pagan (original illustration from Uchida, 1944).
DERMAPTERA (earwigs) [Fig. 17]
No earwigs were found previous to this study. Two species of earwigs were collected in
this study and mark the first records of the order from Pagan. One species, Euborellia
stali [Anisolabididae], is cosmopolitan and known from many localities in the western
Pacific, Japan, and SE Asia as well as also being found in Africa and India. It was
probably introduced to Pagan through the shipping of supplies. The other (Fig. 17) is as
an undetermined labidiid, but is conspicuous with its huge caudal forceps. Earwigs are
omnivores and will eat many types of organic matter.
Fig. 17. An undetermined labidiid earwig from Pagan. Photo: Darcy Oishi.
Evenhuis et al. — Pagan Island Arthropod Survey
32
DIPLOPODA (millipedes) [Fig. 18]
No millipedes had previously been recorded from
Pagan. Two species are recorded for Pagan for the
first time based on collection made during this
survey. One species, the rusty millipede,
Trigoniulus corallinus (Fig. 18), was collected and
identified. The species is native to Thailand and
Burma but has been introduced to many areas of
the Pacific and Indian Ocean, and also the west
Indies and North America (Shelley et al., 2006).
The other species, Harpaphes haydeniana is the
common yellow spotted garden millipede found
Fig. 18. Trigoniulus corallinus, the
throughout North America but not known
rusty millipede. New Island Record.
previously from the west Pacific. Millipedes live in
the soil and are detritovores and scavengers. Since
this survey did not do soil sampling, we expect more species to be found through further
more focused collecting for this group.
DIPTERA (flies, gnats, midges) [Figs. 19, 20]
In previous surveys, flies were not well
recorded with only 21 species in 10 families
being published. We identified 69 different
species of Diptera in 29 families that were
collected during this survey, with 61 of them
new records to Pagan, bringing the total
number of Diptera species known from
Pagan to 82. Of these, the following 19
families are new records to the island:
Anthomyzidae, Canacidae (beach flies),
Cecidomyiidae (gall midges),
Ceratopogonidae (biting midges),
Chamaemyiidae, Chironomidae (midges),
Chloropidae (frit flies, eye gnats),
Drosophilidae (pomace flies), Ephydridae
(shore flies), Keroplatidae (predaceous
gnats), Limoniidae (crane flies), Milichiidae,
Nannodastidae, Phoridae (coffin flies),
Platystomatidae, Psychodidae (moth flies),
Sciaridae (fungus gnats), Sphaeroceridae
(wrack flies), and Syrphidae (flower flies).
The probable reason for the significantly
large amount of new faunistic information on
Diptera is that few workers bother to identify
Fig. 19. New genus and new species
of hydrophorine dolichopodid fly from
rocky shores on Pagan. New Island
Record. Photo: Neal Evenhuis.
Evenhuis et al. — Pagan Island Arthropod Survey
33
Diptera outside of the nuisance, agricultural pest, or disease-causing species and leave the
others undetermined. In this report, we have made a special effort to identify as many
taxa as possible of this order to better show its true representation in the fauna.
Many of the species listed are synanthropic or associated with agricultural crops
as pests or biological control agents. However, the discovery here of a few new taxa and
the recollection of a previously recorded endemic taxon indicate that, despite the
tremendous amount of human and volcanic impact on the island, there are still pockets of
native species that are surviving. See below under SIGNIFICANT FINDINGS for more details
on the new marine taxa found (e.g., Fig. 19).
Fig. 20. The ubiquitous Musca sorbens, also known as the dog dung fly.
One of the most common and most annoying insects on Pagan.
EMBIIDA (web spinners)
No web spinners had been previously recorded from Pagan. The species collected and
identified in this survey marks the first record of this species from Pagan. Oligotoma
humbertiana is a common widespread species in southern and eastern Asia and the
western Pacific that is commonly introduced in islands through transport of soils.
HEMIPTERA (true bugs)
(sometimes referred to as “Heteroptera” and encompassing the Homoptera)
The true bugs include many plant and agricultural pests, therefore attention has been paid
to them in previous studies. Twenty taxa in 6 families had been identified from Pagan
Evenhuis et al. — Pagan Island Arthropod Survey
34
previous to this survey. Collections made here have resulted in identifications of 12
species, 10 of which are new records. Of these, one new undescribed endemic species
was discovered by Dan Polhemus and is tentatively placed in the dumping-ground genus
Lygus until further research can pinpoint its true generic status. Polhemus also identified
marine Hemiptera, all new records to the island [see his report: Polhemus (2010) for
details]. It is probable that collections made at the shore and in the littoral zone had not
been conducted previously so all marine-associated arthropods were not recorded
previous to this study. Many of the aquatic forms are widespread taxa with the ability to
colonize tropical island habitats. With the material identified during this project, the total
number of Hemiptera known from Pagan now stands at 31 species in 13 families.
HOMOPTERA (sucking bugs, aphids, scale insects)
(some have split this order into Auchenorrhyncha and Sternorhyncha)
Homopterans are primarily sap sucking bugs that include the aphids, mealybugs, and
scale insects. Previous to this survey, 16 homopterans had been recorded in the literature.
We identified 7 species, of which 5 are new records for the island. The total known
homopteran fauna on Pagan is now 20 species. The new record of the leafhopper
Acertagallia (Cicadellidae) may represent a recent introduction, as it was found in a
number of collections so is well established on the island. As such, one would expect that
it would have been collected previously. Leafhoppers are known to transmit plant
diseases but nothing is known of the hosts or biology of the specimens collected during
this study.
The collecting techniques utilized during this survey did not collect the scale
insects and mealybugs that are normally easily identifiable. With regard to those smaller
specimens that were found in the material collected, given the time available between
delivery of material and the due date for the report, the resolution for some of the
identifications of aphids and scale insects which normally require slide preparation and
mounting for identification was not necessarily to species level.
HYMENOPTERA (wasps, bees, ants, parasitica) [Figs. 21, 22, 23, 32]
Hymenoptera included some of the most commonly encountered insects on the island.
Previous to this survey, 42 species in 12 families had been identified and published. We
have identified 47 species based on the current collections, of which 25 are new island
records bringing the total hymenopteran fauna known on Pagan to 69 species in 20
families.
The social wasps recorded were no doubt introduced to the islands long ago as
they are found in some of the earliest surveys. A few of these wasps will enter buildings
and form nests [see Fig. 21], but others prefer to build nests in the forests which are
closer to potential prey which they use to nourish their young.
Evenhuis et al. — Pagan Island Arthropod Survey
35
Fig. 21. Social and solitary wasp nests (undetermined spp.) in abandoned building in Shomushon village.
Photo: Jennifer Stauffler.
One of the most commonly collected groups during this survey were ants. They
not only were tops in numbers of specimens but also had the highest species-diversity per
family for any arthropod known from Pagan. This is no doubt due to the extreme success
ants have at invading and colonizing new habitats such as islands.
Fig. 22. Anoplolepis gracilipes, the crazy ant, worker caste. Photo: Antweb.
Evenhuis et al. — Pagan Island Arthropod Survey
36
Fig. 23. Tetramorium smithi, worker caste. New Island Record. Photo: Will Haines.
Previous to this study, 18 species of ants were known from Pagan, mainly all
widespread species. During this survey, we identified 25 species including 10 new island
records, bringing the total known number of ants from Pagan to 28. None of these species
are endemic to the island and only a few may be native; the remainder are alien
introductions. Their presence poses the greatest threat to the native ecosystem as many
species are either predaceous on other invertebrates while others are seedeaters. Any
unchecked predator can cause deleterious effects on plant and animals though depletion
of resources and destruction of the flora. Although the most pervasive species found on
Pagan and collected in huge numbers (hundreds) in some of the traps (see Fig. 32), the
crazy ant, Anoplolepis gracilipes (Fig. 22), is not a major threat now but given its
capacity to form what are called “supercolonies” in island ecosystem situations, it may
become a serious pest [see SIGNIFICANT FINDINGS below for more information.
Five species of bees were previously recorded from Pagan (including one
endemic megachilid bee) but were not as commonly collected during this survey (only 1
was identified, the common honeybee Apis mellifera). This may be due to competition
with other more aggressive insects such as the social wasps and ants, both of which can
either attack and overcome bees or outcompete for food resources or even deplete them.
ISOPODA (sow bugs, pill bugs, roly-polys, wood lice) [Fig. 24]
Two unnamed taxa of terrestrial isopods have previously been recorded from Pagan.
During this survey, we collected one unnamed species in the family Armadillidae (pill
bugs). These crustaceans are debris feeders and are harmless (they are not pestiferous or
Evenhuis et al. — Pagan Island Arthropod Survey
37
of any medical importance). They require moisture to survive so will be found in moist
soil or under rocks or debris. They are nocturnal in feeding habits and not usually seen
during the day unless an overabundance in populations causes them to expand into
human habitations in search of moisture and shelter. Pill bugs are easily separated from
their close cousins, the cosmopolitan wood lice (Porcellionidae) by the fact that pill bugs
can roll up into a ball, wood lice cannot.
Fig. 24. Species of Armadillidae, a pill bug. Photo: Bugguide.net.
ISOPTERA (termites)
Termites are cosmopolitan but none had previously been recorded from Pagan. The
discovery of the subterranean termite Rhinotermes inopinatus marks the first record of
termites from Pagan. The species is a common widespread species found throughout the
Pacific.
LEPIDOPTERA (moths, butterflies) [Figs. 25, 26]
Butterflies and moths are conspicuous insects in any arthropod survey. Although many
lepidopterans are large and showy, there are not very many species known from Pagan.
Previous to this study, only 10 species in 4 families had been published. During this
study, using a variety of trapping methods as well as hand collecting, we have identified
17 species, of which 12 are new island records. The lepidopteran fauna on Pagan is now
22 species in 11 families.
Evenhuis et al. — Pagan Island Arthropod Survey
Fig. 25. The widespread Blue-branded king crow butterfly, Euploea eunice, female.
The caterpillars feed on fig trees. Photo: Darcy Oishi.
Fig. 26. The widespread (throughout the Pacific) Common eggfly butterfly, Hypolimnas bolina,
female. Caterpillars feed on a variety of plants. New Island Record. Photo: Darcy Oishi.
38
Evenhuis et al. — Pagan Island Arthropod Survey
39
One new record, the butterfly Hypolimnas bolina (Fig. 26), is a very widespread
Pacific species, and it is unusual that is had not been recorded previously from Pagan. It
was commonly collected during this survey, so has most likely been established on the
island for some time.
Light trapping was conducted at two sites during this survey, which is the best
method for collecting night flying insects such as moths. Despite the trapping, only a
relatively few specimens of moths were identified during this survey, which most likely
was as direct result of trapping during bad weather and/or bright, moonlit nights.
Some of the species identified based on collections made during this study
(Endotricha sp., Asymphrodes sp., and Anocharis sp.) belong to large genera that are very
diverse in the Pacific, so there is a high probability that eventual species-level
identification may show that some are native or even endemic species on Pagan.
MANTODEA (mantids)
Only one species of praying mantis, Orthodera burmeisteri, a widespread species
throughout the Pacific was previously recorded from Pagan and was also collected on this
survey. Mantids are general predators and a natural component of most ecosystems.
However, the species here was undoubtedly introduced long ago to the island. Mantids
produce egg sacs (oothecae), a foam-like substance is produced by the mantid
surrounding the eggs that hardens and protects the eggs while they are developing. These
oothecae are usually attached to twigs or leaves.
Fig. 27. Ant lion larva from Pagan attacking crazy ant. Photo: Darcy Oishi.
Evenhuis et al. — Pagan Island Arthropod Survey
40
NEUROPTERA (lacewings, ant lions) [Fig. 27]
Previous to this study, three species of neuropterans, two lacewings and an ant lion, were
known from Pagan. Collections made during this study have identified 6 new species
records including the first record of a brown lacewing (Hemerobiidae). In addition, larvae
and adults of ant lions were collected, of which the adult was identified as a species of
Myrmeleon, which is a new record to the island.
ODONATA (dragonflies, damselflies) [Fig. 28]
Four species of odonates were previously recorded from Pagan: one damselfly (Ischnura
aurora) and three dragonflies. Three of the four species were recovered during this study.
Dan Polhemus (pers. comm.) indicated that two of the three dragonflies may include a
misidentification, so that there really may only be 3 species that occur from the island. A
conspicuous absence on Pagan is the widespread and long-distance flier, Pantala
flavescens. It occurs on most Pacific islands and is known to fly more than 1500 km over
oceans to get from one freshwater source to another. The species is known from other
islands in the Marianas but has not been seen on Pagan.
Fig. 28. The dragonfly Diplacodes bipunctata, male, from American Samoa. Photo: Dan Polhemus.
ORTHOPTERA (crickets, grasshoppers) [Figs. 29, 30]
Only three orthopterans were previously recorded from Pagan prior to this survey. Based
on collections made during this project, 15 taxa have been identified including 14 new
records. These include six acridids (grasshoppers), two katydids, three gryllids (crickets),
Evenhuis et al. — Pagan Island Arthropod Survey
41
two tetrigid grasshoppers, and the rare ant inquiline, Myrmecophilus leei. See below
under SIGNIFICANT FINDINGS for more details on this last species.
Fig. 29. Unidentified tetrigid grasshopper. New Island Record. Photo: Darcy Oishi
Fig. 30. The ant inquiline, Myrmecophila leei. New Island Record. Photo: Neal Evenhuis
PHASMATODEA (walking sticks, stick insects)
One species of stick insect, the widespread Acanthograeffea denticulata, was previously
recorded from Pagan. No collections of this or any other stick insects were made during
this survey, but we expect the species still exists. It is known as a pest of coconut and
coconut groves are abundant on Pagan, so it should still be there. This is an introduced
Evenhuis et al. — Pagan Island Arthropod Survey
42
insect that has been on the island for quite a long time. It was first recorded by
Yasumatsu (1940) from collections made on Pagan in April 1940.
PSEUDOSCORPIONIDA (pseudoscorpions)
Pseudoscorpions are tiny (usually no more than 2 mm in length) arthropods that live in
leaf litter and under tree bark where they prey on microscopic invertebrates. Three
species had previously been recorded from Pagan (all taken from soil samples). Two
species were identified during this survey, both new records for the island.
Pseudoscorpions are not commonly found on tropical islands so the finding of all these
different species on Pagan is notable. Further research should be carried out to determine
why.
PSOCOPTERA (book lice, bark lice)
Bark lice are common forest dwellers where they are found in leaf litter or under
decaying bark of trunks, logs, or twigs. One species, Caecilius analis was previously
recorded (Thornton, 1981) but not collected during this survey. However, eight other
species were identified from collections made during this survey, which are all new
records for the island. As these species are a common and diverse group on other Pacific
islands, we believe further more rigorous collecting focusing on this group should
produce more species records from Pagan.
SCORPIONIDA (scorpions) [Fig. 31]
Previous records report two species of scorpions (one unnamed) from Pagan. One of
these two species, Liocheles australasiae (Fig. 31) was found in one of the craters on
south Pagan and identified during this survey. It is a small (ca. 2 cm long), common and
widespread species throughout the Pacific islands. The sting of scorpions can be painful
in some species but is rarely fatal. The sting of Liocheles australasiae is no worse than a
bee sting. The paucity of material of collected during this survey despite the numerous
pitfall traps throughout the island lends support to the presumption that their presence is
very rare and as such they pose no threat or danger to humans or other animals on Pagan.
Evenhuis et al. — Pagan Island Arthropod Survey
43
Fig. 31. The scorpion, Liocheles australasiae, commonly found throughout the Pacific.
Approximately 2 cm long. Photo: Ryo Kenzaki.
SIPHONAPTERA (fleas)
The only known flea recorded from Pagan is the cat flea. It was not collected on this
survey, but feral cats still abound on the island and it is presumed that the associated fleas
still occur there as well.
THYSANOPTERA (thrips)
Two species of thrips were previously recorded from Pagan. Two thrips were found in
the material collected but only one could be identified to species level due to time
constraints. More specimens should be expected, as thrips are ubiquitous arthropods in all
island faunas. Specialized collecting techniques focusing directly on host plants may be
necessary to secure specimens for study. Thrips are known to be pests on crops. It is not
known whether or not the two species previously recorded or the one collected in this
study are pestiferous.
THYSANURA (bristletails, silverfish)
Silverfish are ubiquitous creatures usually associated with human habitation. They are
detritus feeders and are pests in offices and libraries where they feed on paper.
Evenhuis et al. — Pagan Island Arthropod Survey
44
SIGNIFICANT FINDINGS
Invasive species
HYMENOPTERA: FORMICIDAE
ANOPLOLEPIS GRACILIPES (SMITH, 1857) [Figs. 22, 32]
Probably the most prevalent arthropod in the survey is the crazy ant, Anoplolepis
gracilipes. This highly invasive species is found worldwide and in some places where it
has become introduced (e.g., Christmas Island in the Indian Ocean), it forms
supercolonies (Abbott, 2006). These are multiple colonies with queens but no
competition or aggravated behavior takes place between colonies allowing high densities
of the ants in small areas where they can devastate plant and invertebrate biota to use as
food resources for the burgeoning nests. No supercolonies were observed on Pagan;
however, the high levels of abundance of the ants in pitfall and pan traps (e.g., Fig. 32)
may be an indicator that this species may eventually form a supercolony on Pagan.
Fig. 32. Single sample of the crazy ant, Anoplolepis gracilipes,
collected from a pitfall trap on Pagan. Photo: Darcy Oishi.
Evenhuis et al. — Pagan Island Arthropod Survey
45
Selected Rare, New, or Unusual Species
DIPTERA: DOLICHOPODIDAE
HYDROPHORINAE - NEW GENUS, NEW SPECIES [Fig. 19]
A new genus and species of hydrophorine marine dolichopodid fly was collected off
rocks on the west coast of Pagan. This is the first record for Pagan of this new taxon,
which is otherwise known only from Guam. It should also occur on the islands between
Guam and Pagan but have yet to be collected. Recent collections on Saipan by Dan
Polhemus in September 2010 did not find any.
This taxon is found along rocky shores in the splash zone (Fig. 33). It is
predaceous on other small invertebrates both as immatures and adults. Immatures were
not found but should be found in small rock holes and crevices in association with algae
and other littoral invertebrate organisms.
Fig. 33. Rocky shoreline on south side of Bandeera peninsula where a new genus of marine
dolichopodid flies was collected. Photo: Dan Polhemus.
ORTHOPTERA: MYRMECOPHILIDAE
MYRMECOPHILA LEEI KISTNER & CHONG 2007 [FIG. 30]
This rare and unusual creature is an inquiline in ant nests. It was found associated with
Anoplolepis on the south portion of Pagan and is the first record of this family from the
island. It appearance here on Pagan is unusual in that, previous to this survey, it was
originally described and known only from mainland Malaysia. It is most likely that this is
Evenhuis et al. — Pagan Island Arthropod Survey
46
much more widespread than previously known but has been neglected in previous
collections because of its superficial resemblance to immature cockroaches. Samples
from collecting on other areas between SE Asia and Pagan should be checked for this
interesting ant inquiline.
THREATS TO FRUIT BATS OR THE MEGAPODE FAUNA
No direct threats to either megapodes or the fruit bat were found in this survey.
Theoretically, predator and parasitic arthropods that deplete food resources may have an
indirect affect on populations, but there is no evidence to suggest this has or will happen
in the near future by any of the arthropods identified in this survey.
ACKNOWLEDGMENTS
The staff of the library and archives of the Bishop Museum and Hamilton Library at the
University of Hawaii at Manoa assisted with obtaining some of the literature. Al
Samuelson (beetles) and Kelly Miller (embiids) kindly assisted with identifications. Dan
Polhemus allowed use of information from his aquatic insect survey report in this overall
arthropod report. Maps 2 and 3 were prepared by Cory Campora. Many of the photos
used in this report were kindly provided by Christa Russell, Cory Campora, and Dan
Polhemus. Eiko Lynch assisted with transliterating the 1940 Yasumatsu localities. We
especially acknowledge the efforts of the collectors of this survey, Christa Russell, Mike
Richardson, Cory Campora, Stephan Lee, and Justin Fujimoto. Neither erupting
volcanoes, sweltering heat during “death marches” across the island, charging bulls,
constant companionship of “overly attentive” flies, thousands of ants billowing out of
nests, nor spam, spam, and even more spam did not deter them from their goal of making
an extensive and successful survey of the island of Pagan in only a few days.
LITERATURE CITED
Abbott, K.L. (2006) Spatial dynamics of supercolonies of the invasive yellow crazy ant,
Anoplolepis gracilipes, on Christmas Island, Indian Ocean. Diversity and Distributions
12(1): 101–110.
Akasura, A. & Furuki, T. (editors) (1994) Biological expedition to the northern Mariana
Islands, Micronesia. Natural History Research, Special Issue 1: 1–344.
Akasura, A., Ohba, T., Miyano, S., Furuki, T., Kurozumi, T. & Harada , H. (1994)
Outline of the biological expedition to the Northern Mariana Islands, Micronesia. Natural
History Research, Special Issue 1: 1–11.
Evenhuis et al. — Pagan Island Arthropod Survey
47
Bryan, E.H. Jr. (1949) Economic insects of Micronesia. Report of the Insect Control
Committee for Micronesia 1947–1948. National Research Council, Washington, D.C. 29
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the Marshall, Caroline and Mariana Islands. Pacific Science Information Center,
Honolulu.
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Pagan, Maria Islands. Intelligence Division, Office of the Engineer, U.S. Geological
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Schutzgebieten 15: 96–118.
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Hosokawa, T. (1934) Preliminary account of the vegetation of the Marianne Islands
group. Bulletin of the Biogeographical Society of Japan 5: 124–172. [In Japanese.]
Marche, A. (1891) Rapport général sur une mission aux îles Mariannes. Nouvelles
Archives des Missions Scientifiques et Litteraires 1: 241–280.
Marche, A. (1982) The Mariana Islands. Translated from the French by Sylvia E. Cheng.
Micronesian Area Research Center, University of Guam. 52 pp.
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Marianas. Report for US Fish & Wildlife Service, Honolulu. 20 pp.
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48
Prowazek, S. von (1913) Die Deutschen Marianen. Ihre Natur und Geschichte. A. Barth,
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Entomological News 117: 239–240.
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Islands 1899–1914. Commonwealth of the Northern Mariana Islands, Division of
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D.A. (editors), Encyclopedia of islands. University of California Press, Berkeley. 1074
pp.
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Evenhuis et al. — Pagan Island Arthropod Survey
APPENDICES
49
Evenhuis et al. — Pagan Island Arthropod Survey
50
APPENDIX I.
BIBLIOGRAPHY OF PAGAN TERRESTRIAL ARTHROPODS
The following is a complete list of all known publications specifically listing Pagan
terrestrial arthropods including articles that mention arthropods as occurring on the island
but without a scientific name. Many more articles list terrestrial arthropods with a low
geographical resolution of only “Marianas” or “Northern Marianas”. If those articles do
not specifically mention Pagan, they are not included below.
Adams, P.A. (1959) Myrmeleontidae and Chrysopidae. Insects of Micronesia 8(2): 13–
33.
Akasura, A. & Furuki, T. (editors) (1994) Biological expedition to the northern Mariana
Islands, Micronesia. Natural History Research, Special Issue 1: 1–344.
Akasura, A., Ohba, T., Miyano, S., Furuki, T., Kurozumi, T. & Harada , H. (1994)
Outline of the biological expedition to the Northern Mariana Islands, Micronesia. Natural
History Research, Special Issue 1: 1–11.
Aoki, J. (1994) Oribatid mites of the Northern Mariana Islands, Micronesia. 1. Uracas
and Maug islands. Natural History Research, Special Issue 1: 181–194.
Barber, H.G. (1958) Heteroptera. Lygaeidae. Insects of Micronesia 7(4): 173–218.
Beal, R.S. (1961) Coleoptera: Dermestidae. Insects of Micronesia 16(3): 109–131.
Beardsley, J.W. (1963) The Coccidae of Micronesia (Homoptera). PhD dissertation,
University of Hawaii at Manoa. iv + 324 pp.
Beardsley, J.W. (1966) Homoptera. Coccoidea. Insects of Micronesia 6(7): 377–562.
Bickel, D.J. (1994) Diptera: Dolichopodidae Part I. Sciapodinae, Medeterinae and
Sympycninae (part). Insects of Micronesia 13(8): 361–406.
Bohart, R.M. (1957) Diptera: Culicidae. Insects of Micronesia 12(1): 1–85.
Cartwright, O.L. & Gordon, R.D. (1971) Coleoptera: Scarabaeidae. Insects of Micronesia
17(4): 257–296.
Carvalho, J. (1956) Heteroptera. Miridae. Insects of Micronesia 7(1): 1–100.
Chapin, E.A. (1957) Scorpionida. Insects of Micronesia 3(2): 65–70.
Chapin, E.A. (1965) Coleoptera: Coccinellidae. Insects of Micronesia 16(5): 189–254.
Evenhuis et al. — Pagan Island Arthropod Survey
51
Clouse, R.M. (2007) The ants of Micronesia (Hymenoptera: Formicidae). Micronesica
39(2): 171–295.
Corwin, G., Bonham, L.D., Terman, M.J. & Viele, G.W. (1957) Military geology of
Pagan, Maria Islands. Intelligence Division, Office of the Engineer, U.S. Geological
Survey, Washington, D.C. 259 pp.
Cruz, J., Arriola, L., Johnson, N. & Beauprez, G. (2000) Wildlife and vegetation surveys
Pagan 2000. Technical Report #7 CNMI-DFW, unpublished. 68 pp.
Darlington, P.J. Jr. (1970) Coleoptera: Carabidae including Cicindelinae. Insects of
Micronesia 15(1): 1–49.
Esaki, T. (1940) [South Seas Islands’ injurious insect fauna and their control.]
Shokobutsu oyobi Dobutsu 8: 274–280. [In Japanese.]
Esaki, T. (1943) [Scorpions of Micronesia]. Acta Arachnologica 8(1-2): 1–5. [In
Japanese.]
Esaki, T. (1943) [Fauna of injurious insects of Micronesia.] Shokobutsu oyobi Dobutsu
11: 839–843. [In Japanese.]
Esaki, T. & Ishihara, T. (1943) Hemiptera Micronesica, I. Nabidae. Mushi 15: 69–75.
Fukushima, I. (1947) Noctuidae of Micronesia (Lepidoptera). Mushi 18(1): 1–22.
Gressitt, J.L. (1954) Introduction. Insects of Micronesia 1: 1–257.
Gressitt, J.L. (1956) Coleoptera: Cerambycidae. Insects of Micronesia 17(2): 61–183.
Gross, G. (1963) Coreidae, Neididae, and Nabidae. Insects of Micronesia 7(7): 357–390.
Hardy, D.E. & Adachi, M.S. (1956) Diptera: Tephritidae. Insects of Micronesia 14(1): 1–
28.
Hiramatsu, N. 1994. Preliminary list of Uropodidae (Acari: Gamasida) from the northern
Mariana Islands, Micronesia. Natural History Research, Special Issue 1: 179–180.
Hodkinson, I.D. (1983) The psyllids (Homoptera: Psylloidea) of the Austro-Oriental
Pacific and Hawaiian zoogeographical realms. Journal of Natural History 17: 341–377.
Hopkins, G.H.E. (1961) Siphonaptera. Insects of Micronesia 14(4): 91–107.
Itoh, R. (1994) Symphypleona (Insecta: Collembola) from the Northern Marian Islands,
Micronesia. Natural History Research, Special Issue 1: 217–218.
Evenhuis et al. — Pagan Island Arthropod Survey
52
Kasahara, S. (1994) Carabidae (Insecta: Coleoptera) of the Mariana Islands, Micronesia.
Natural History Research, Special Issue 1: 221–222.
Kevan, D.K.McE. & Vickery, V.R. (1997) An annotated provisional list of nonsaltatorial orthopteroid insects of Micronesia, compiled mainly from the literature.
Micronesica 30(2): 269–353.
Kono, H. (1942) Die Rüsselkäfer auf den Micronesien. Insecta Matsumurana 16(4): 13–
21.
Krombein, K.V. (1949) The aculeate Hymenoptera of Micronesia I. Scoliidae, Mutillidae,
Pompilidae, and Sphecidae. Proceedings of the Hawaiian Entomological Society 13:
367–410.
Krombein, K.V. (1950) The aculeate Hymenoptera of Micronesia II. Colletidae,
Halictidae, Megachilidae and Apidae. Proceedings of the Hawaiian Entomological
Society 14: 101–142.
Kulzer, H. (1957) Coleoptera: Tenebrionidae. Insects of Micronesia 17(3): 185–256.
Marche, A. (1891) Rapport général sur une mission aux îles Mariannes. Nouvelles
Archives des Missions Scientifiques et Litteraires 1[1889]: 241–280.
Marche, A. (1982) The Mariana Islands. Translated from the French by Sylvia E. Cheng.
Micronesian Area Research Center. 52 pp.
Mathis, W.N. (1993) Studies of Gymnomyzinae (Diptera: Ephydridae), IV: a revision of
the shore-fly genus Hecamede Haliday. Smithsonian Contributions to Zoology 541: 1–46.
Miyano, S. (1994) Insects of the Northern Mariana Islands, Micronesia, collected during
the expedition. Natural History Research, Special Issue 1: 199–215.
Miyano, S. (1994) Some ecological observations of social wasps (Insecta: Hymenoptera:
Vespidae) in the Northern Mariana Islands, Micronesia. Natural History Research,
Special Issue 1: 237–245.
Miyano, S. (1994) Results of fruit fly (Insecta: Diptera: Tephritidae) trapping in the
Northern Mariana Islands, Micronesia. Natural History Research, Special Issue 1: 247–
248.
Nakata, S. (1961) Some notes on the occurrence of Phasmotodea in Oceania. Pacific
Insects Monograph 2: 107–121.
Nomura, S. (1943) Zur Kenntnis der Aphodiiden aus Mikronesien (Coleoptera:
Scarabeidae). Mushi 15: 77–82.
Evenhuis et al. — Pagan Island Arthropod Survey
53
Saito, A. (1994) Records of some Cerambycidae (Insecta: Coleoptera) from the Northern
Mariana Islands, Micronesia. Natural History Research, Special Issue 1: 227–228.
Sasakawa, M. (2009) Diptera: Lauxaniidae. Insects of Micronesia 14(9): 303–327.
Snyder, F.M. (1965) Diptera: Muscidae. Insects of Micronesia 13(6): 191–327.
Spencer, K.A. (1961) A synopsis of the Oriental Agromyzidae (Diptera). Transactions of
the Royal Entomological Society of London 113(4): 55–100.
Spencer, K.A. (1963). Diptera: Agromyzidae. Insects of Micronesia 14(5): 135–162.
Tadauchi, O. (1994) Bees of the Mariana Islands, Micronesia, collected by the Expedition
of the Natural History Museum & Institute, Chiba (Hymenoptera, Apoidea). Esakia 34:
215–225.
Takahashi, R. (1942) Some species of Aleyrodidae, Aphididae and Coccidae in
Micronesia (Homoptera). Tenthredo 3(4): 349–358.
Terayama, M., Miyano, S. & Kurozumi, T. (1994) Ant fauna (Insecta: Hymenoptera:
Formicidae) of the Northern Mariana Islands, Micronesia. Natural History Research,
Special Issue 1: 231–236.
Thornton, I.W.B. (1981) The Psocoptera of the Hawaiian Islands. Parts I and II.
Introduction and the nonendemic fauna. Pacific Insects 23: 1–49.
Townes, H. (1958) Hymenoptera: Ichneumonidae, Stephanidae, and Evaniidae. Insects of
Micronesia 19(2): 37–87.
Tuthill, L.D. (1951) Records and descriptions of some Micronesian Psyllidae
(Homoptera). Proceedings of the Hawaiian Entomological Society 5: 273–278.
Tuthill, L.D. (1964) Homoptera: Psyllidae. Insects of Micronesia 6(5): 353–376.
Uchida, H. (1944) Collembola von Mikronesien. Bulletin of the Tokyo Science Museum
17: 1–23. [In Japanese with German summary.]
Van Zwaluwenberg, R.H. (1957) Coleoptera: Elateridae. Insects of Micronesia 16(1): 1–
66.
Vickery, V.R., Kevan, D.K.McE. & English, M.-L. (1999) Gryllacridoidea,
Rhaphidophoroidea and Tettigonoidea. Insects of Micronesia 5(4): 11–83.
Watanabe, C. (1945) Note on some Micronesian Braconidae (Hymenoptera). Mushi
16(9): 47–58.
Evenhuis et al. — Pagan Island Arthropod Survey
54
Watanabe, C. (1958) Hymenoptera: Eucharidae. Insects of Micronesia 19(2): 19–34.
Wheeler, M.R. & Takada, H. (1964) Diptera: Drosophilidae. Insects of Micronesia 14(6):
163–242.
Williams, D.J. (1994) Distribution of the Pacific coconut mealybug, Dysmicoccus cocotis
(Maskell) and of a new related species on coconut in southern Asia (Hemiptera:
Coccoidea: Pseudoccidae). Journal of Natural History 28: 365–371.
Wood, S.L. (1960) Coleoptera: Platypodidae and Scolytidae. Insects of Micronesia 18(1):
1–73.
Yasumatsu, K. (1940). [An excursion to the island of Pagan, Marianas]. Akitu 2(4): 180–
188. [In Japanese.]
Yasumatsu, K. (1940) Evaniidae of Micronesia (Hymenoptera). Kontyu Kenkyu 4(1-2):
19–21.
Yasumatsu, K. (1941) Beitrag zur Kenntnis der Echthromorpha-Fauna Mikronesiens
(Hym., Ichneumonidae). Insecta Matsumura 15: 141–145.
Yasumatsu, K. (1941) Sphecoidea of Micronesia. III. Family Larridae. Mushi 14: 44–47.
Yasumatsu, K. (1942) Apoidea of Micronesia. III. Records of the genera Megachile,
Heriades, Ceratina and Prosopis. Tenthredo 3(4): 355–348.
Yasumatsu, K. (1942) Notes on some Micronesian Elasmidae and Eucharidae
(Hymenoptera, Chalcidoidea). Insecta Matsumurana 16: 151–158.
Yasumatsu, K. (1942) [Stick insects’ eggs.] Takarazuka Kontyû 18: 1–20. [In Japanese.]
Yasumatsu, K. (1945) Vespoidea of Micronesia. 2 (Hymenoptera). Mushi 16(8): 35–45.
Yasumatsu, K. & Yoshimura, S. (1945) Some chalcidoid parasites of Saissetia nigra and
S. hemisphaerica in Micronesia (Hymenoptera). Mushi 16(7): 29–34.
Yoshida, M., Tanikawa, A., Ikeda, H., Tanaka, H., Nishikawa, Y. & Kamura, T. (1996)
The spiders of the Northern Mariana Islands, Micronesia. Journal of the Natural History
Museum and Institute, Chiba 4(1): 59–64.
Evenhuis et al. — Pagan Island Arthropod Survey
55
APPENDIX II. CHECKLIST OF PAGAN ARTHROPODS
(taxa in italics are synonyms or misidentifications [that appeared in the literature] of current valid names)
Order/Family
ACARINA (mites,
Ameroseiidae
Eviphididae
Ixodidae
Laelapidae
Ologamasidae
Oribatidae
Podocinidae
Pyemotidae
Rhodacaridae
Uropodidae
Taxon*
ticks)
Ameroseius sp. 2
Eximiris sp.
prob. Boophilus microplus
Hypoaspis sp. 1
Hypoaspis sp. 2
Gamasiphis sp. 1
Gamasiphis sp. 2
Gen. sp.
Podocinum jamaicense Evans & Hyatt, 1957
Pyemotes sp.
Rhodacarus sp.
Metagynella sp. A
Trigonuropoda sp. B
Trigonuropoda sp. D
Uroobovella sp. C
ARANEAE (spiders)
Anyphaenidae
Gen. sp.
Araneidae
Argiope appensa (Walckenaer, 1841)
Argiope sp. 2
Neoscona sp.
Neoscona theisi (Walckenaer, 1842)
Clubionidae
Gen. sp. 1
Gen. sp. 2
Gen. sp. 3
Corinnidae
Gen. sp.
Gnaphosidae
Gen. sp. 1
Gen. sp. 2
Linyphiidae
Gen. sp.
Liocranidae
Apostenus sp.
Lycosidae
Gen. sp.
Pardosa sp.
Schizocosa sp. 1
Schizocosa sp. 2
Schizocosa sp. 3
Miturgidae
Cheiracanthium sp.
Nesticidae
Gen. sp. 1
Gen. sp. 2
Ooponidae
Gamasomorpha sp.
Pholcidae
Gen. sp.
Pisauridae
Dolomedes sp.
Salticidae
Athamas whitmeei Pickard-Cambridge, 1877
Hasarius adansoni Savigny & Audouin, 1825
Plexippus sp. 1
Plexippus sp. 2
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Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
ARANEAE (spiders) [continued]
Salticidae
Plexippus paykulli (Audouin, 1826)
Sassacus sp.
Thorelliola ensifera (Thorell, 1877)
Scytodidae
Scytodes sp.
Sparassidae
Heteropoda sp.
Olios sp.
Tengellidae
Gen. sp.
Tetragnathidae
Gen. sp.
Theridiidae
Argyrodes sp.
Coleosoma floridanum Banks, 1900
Thomisidae
Thomisius sp.
Gen. sp.
Titanoecidae
Gen. sp.
Titanoeca? sp.
Zodariidae
Zodarion? sp.
BLATTARIA (roaches)
Blaberidae
Diploptera dytiscoides (Serville, 1838)
Blatellidae
Blatella lituricollis (Walker, 1868)
Onychostylus notulatus (Stål, 1861)
Blattidae
Periplaneta americana (Linnaeus, 1758)
Periplaneta australasiae (Fabricius, 1775)
Pycnoscelidae
Pycnoscelus indicus (Fabricius, 1775)
Pycnoscelus surinamensis (Linnaeus, 1758)
CHILOPODA (centipedes)
Scolopendridae
Scolopendra? sp.
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x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
“centipede”
Geophilomorpha gen. sp.
COLEOPTERA (beetles)
Aderidae
Aderus sp.
Anthribidae
Araecerus fasciculatus (De Geer, 1775)
Araecerus vieillardi (Montrouzier, 1860)
Bostrichidae
Xylopsocus capucinus (Fabricius, 1787)
Xylothrips flavipes (Illiger, 1801)
Buprestidae
Agrilus auriventris Saunders, 1873
Carabidae
Agadroma sp.
Gnathaphanus licinoides Hope, 1842
Stenolophus smaragdulus (Fabricius, 1798)
Cerambycidae
Ceresium unicolor unicolor (Fabricius)
Micronesiella mariana (Gressitt, 1956)
x
x
x
x
x
x
x
x
x
x
x
x
x
Sciadella mariana
Phloeopsis meridiana (Ohbayashi, 1941)
x
Sciadella meridiana
Chrysomelidae
Prosoplus marianarum Aurivillus, 1908
Brontispa mariana Spaeth, 1937
x
x
Planispa castaneipennis Chujo, 1937
Gen. sp.
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
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COLEOPTERA (beetles) [continued]
Cleridae
Necrobia rufipes (De Geer, 1775)
Coccinellidae
Coleophora inaequalis (Fabricius, 1775)
Cryptolaemus montrouzieri Mulsant, 1853
Menochilus sexmaculatus (Fabricius, 1781)
Nephus roepkei (Fluiter, 1938)
Telsimia nitida Chapin, 1926
Corylophidae
Sericoderus sp.
Curculionidae
Camptorrhinus dorsalis (Boisduval, 1835)
Curculio sp.
Deretiosus ficae Zimmerman, 1942
Memectetorus cf. setulosus (Boheman, 1859)
Microcryptorhynchus “sp. 2”
Myllocerus sp.
Rhabdocnemis obscura (Boisduval, 1835)
Dermestidae
Dermestes ater De Geer, 1774
x
x
Elateridae
x
Endomychidae
Laemophloeidae
Melyridae
Mordellidae
Nitidulidae
Platypodidae
Salpingidae
Scarabaeidae
Scolytidae
Silvanidae
Staphylinidae
Tenebrionidae
Conoderus pallipes (Eschscholtz, 1829)
Lacon modestus (Boisduval, 1835)
Prodorasterius sp.
Simodactylus cinnamomeus (Boisduval, 1835)
Trochoideus desjardinsi Guérin-Méneville, 1857
Laemophloeus sp. nr. minutus
Gen. sp.
Dellamora castanea (Boheman, 1858)
Carpophilus davidsoni Dobson, 1952
Carpophilus sp. nr. davidsoni?
Carpophilus humeralis (Fabricius, 1798)
Carpophilus dimidiatus (Fabricius, 1792)
Carpophilus sp.
Phylloplatypus pandani Kato, 1998
Prostomiinae, gen. sp.
Adoretus sinicus Burmeister, 1854
Aphodius lividus (Olivier, 1789)
Ericryphalus sylvicola (Perkins, 1900)
Hypothenemus birmanus (Eichhoff, 1878)
Hypothenemus eruditus Westwood, 1836
Cryptamorpha desjardinsi (Guérin-Méneville, 1844)
Silvanus nr. unidentatus (Fabricius, 1792)
Egadroma sp.
Derosphaerus rotundicollis (Castlenau, 1840)
Gonocephalum adpressiforme Kaszab, 1951
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
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x
x
x
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x
x
x
x
x
x
x
x
x
x
x
x
x
x
Gonocephalum sp.
Gonocephalum incisum Blanchard, 1853
Palorus papuanus Kaszab, 1939
COLLEMBOLA (springtails)
Entomobryidae
Gen. sp.
Sira fuscana Uchida, 1944
Isotomidae
Gen. sp. 1
x
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x
x
x
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Order/Family
Taxon*
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COLLEMBOLA (springtails) [continued]
Isotomidae
Gen. sp. 2
Neelidae
Neelus minimus (Willem, 1900)
Poduridae
Gen. sp.
Sminthuridae
Cyphoderus albinus Nicolet, 1841
Sminthurides sp.
Sphaeridia pumilis (Krausbauer, 1891)
x
x
x
x
x
x
DERMAPTERA (earwigs)
Anisolabididae
Euborellia stali (Dohrn, 1864)
Labidiidae
Gen. sp.
x
x
DIPLOPODA (millipedes)
Trigoniulidae
Trigoniulus corallinus (Gervais, 1847)
Xystodesmatidae
Harpaphe haydeniana (Wood, 1864)
x
x
DIPTERA (flies, gnats, midges)
Agromyzidae
Japanagromyza eucalytpi paganensis
Spencer, 1963
Japanagromyza sp.
Liriomyza nr. brassicae (Riley, 1885)
Melanagromyza atomella Malloch, 1914
Pseudonapomyza spicata (Malloch, 1914)
Anthomyzidae
Amalygdops sp.
Calliphoridae
Chrysomya megacephala (Fabricius, 1794)
Chrysomya rufifacies (Macquart, 1843)
Hemipyrellia tagaliana (Bigot, 1877)
Canacidae
Nocticanace peculiaris Malloch, 1933
Cecidomyiidae
Cecidomyiinae Gen. sp.
Lestremia sp.
Ceratopogonidae
Dasyhelea sp. A
Dasyhelea sp. B
Dasyhelea sp. C
Forcipomyia sp.
Chamaemyiidae
Leucopis sp.
Chironomidae
Chironomus longilobus (Kieffer, 1916)
Thalassomyia maritima Wirth, 1947
Chloropidae
Cadrema pallida (Loew, 1865)
Gen. sp.
Siphunculina nitidissima Kanmiya, 1982
Culicidae
Aedes albopictus (Skuse, 1894)
Aedes nocturnus (Theobald, 1903)
Aedes saipanensis Stone, 1945
Anopheles subpictus indefinitus (Ludlow, 1904)
Culex annulirostris marianae Bohart & Ingram, 1946
Dolichopodidae
Chrysosoma mariana Bickel, 1994
Chrysotus sp.
Krakatauia micronesiana Bickel, 1994
Hydrophorinae - New gen. n. sp.
Tachytrechus sp.
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
DIPTERA (flies, gnats, midges) [continued]
Dolichopodidae
Thinophilus sp.
Plagiozopelma flavipodex (Becker, 1922)
Sympycninae - New gen. n. sp.
Drosophilidae
Drosophila ananassae Doleschall, 1858
Leucophenga nigriventris (Macquart, 1843)
Ephydridae
Allotrichoma sp.
Atissa antennalis Aldrich, 1931
Hecamedoides sp.
Paralimna fusca Bock, 1988
Paralimna lineata de Meijere, 1908
Scatella septempunctata Malloch, 1933
Zeros sp.
Keroplatidae
Neoplatyura n. sp.
Lauxaniidae
Homoneura acrostichalis (de Meijere, 1915)
Homononeura sp.
Limoniidae
Dicranomyia basifusca Alexander, 1919
Dicranomyia obesula Edwards, 1927
Dicranomyia pontophila (Tokunaga, 1940)
Lonchaeidae
Lamprolonchaea sp.
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x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Gen. sp.
Milichiidae
Muscidae
Lonchaea sp.
Desmometopa gressitti Sabrosky, 1983
Desmometopa tarsalis Loew, 1866
Milichiella lactiepennis (Loew, 1865)
Atherigona orientalis Schiner, 1858
Atherigona excisa (Thomson)
Nannodastidae
Phoridae
Platystomatidae
Psychodidae
Sarcophagidae
Sciaridae
Sphaeroceridae
Syrphidae
Tephritidae
Haematobia exigua (de Meijere, 1903)
Musca domestica Linnaeus, 1758
Musca sorbens Wiedemann, 1830
Orchisia costata (Meigen, 1826)
Pygophora respondens (Walker, 1859)
Stomoxys calcitrans (Linnaeus, 1758)
Nannodastia sp.
Dohriphora cornuta (Bigot, 1857)
Megaselia setaria (Malloch, 1912)
Puliciphora sp.
Scholastes carolinensis Enderlein, 1924
Psychoda mediocris Quate, 1959
Sarcophaga gressitti Hall & Bohart, 1951
Sarcophaga karnyi (Hardy, 1927)
Sarcophaga misera (Walker, 1849)
Sarcophaga peregrina (Robineau-Desvoidy, 1830)
Scythropochroa quadrispinosa Steffan, 1969
Coproica hirtula (Rondani, 1880)
Poecilosomella punctipennis (Wiedemann, 1824)
Gen. sp.
Lathyrophthalmus arvorum (Fabricius, 1787)
Bactrocera ochrosiae (Malloch, 1942)
Chaetodacus dorsalis, misid.
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
DIPTERA (flies, gnats, midges) [continued]
Tephritidae
Euphranta lemniscata (Enderlein, 1911)
Spathulina acroleuca Schiner, 1868
Ulidiidae
Notogramma cimiciforme Loew, 1868
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x
x
x
x
Gen. sp.
EMBIIDA (web spinners)
Oligotomidae
Oligotoma humbertiana (Saussure, 1896)
HEMIPTERA (true bugs)
Anthocoridae
Buchaniella sodalis (White, 1878)
Lasiochilus marianensis Usinger, 1946
Lasiochilus swezeyi Usinger, 1946
Coreidae
Leptocorixa acuta (Thunberg, 1783)
Leptoglossus australis (Fabricius, 1774)
x
x
x
x
x
x
Leptoglossus membranaceus (Fabricius)
Deltocephalidae
Gerridae
Hermatobatidae
Lygaeidae
Mesoveliidae
Miridae
Nabidae
Plataspidae
Reduviidae
Saldidae
Veliidae
Melanacauthus margineguttatus Distant, 1911
Balclutha rufofasciata (Merino, 1936)
Halobates flaviventris Eschscholtz, 1822
Hermatobates sp.
Bedunia pagana Barber, 1958
Cligenes marianensis Usinger, 1946
Nysius pulchellus Stål, 1859
Pachybrachius nigriceps (Dallas, 1852)
Mesovelia vittigera Horvath, 1895
Campylomma breviceps Usinger, 1946
Campylomma brunneicollis Usinger, 1946
Campylomma lividicornis Reuter, 1912
Camplyomma nr. wakeana Schuh, 1984
Creontiades pallidifer (Walker, 1873)
Eurystylus costalis unicolor Poppius, 1911
Lygocoris kusaiensis (Carvalho, 1956)
“Lygus” n. sp.
Taylorilygus pallidulus (Blanchard, 1852)
Trigonotylus dohertyi (Distant, 1904)
Reduviolus capsiformis (Germar, 1837)
Coptosoma xanthogramma (White, 1842)
Physoderes minor Usinger, 1946
Scadra rufidens Stål, 1859
Saldula palauana Drake, 1961
Halovelia bergrothi Esaki, 1926
Microvelia diluta Distant, 1909
HOMOPTERA (sucking bugs, aphids, scale insects)
Aleyrodidae
Gen. sp.
Neomaskellia bergii (Signoret, 1868)
Aphididae
Aphis gossypii Glover, 1877
Gen. sp.
Aradidae
Mezira sp.
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x
x
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x
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x
x
x
x
x
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x
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x
x
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x
x
x
x
x
x
x
x
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Cicadellidae
Acertagallia sp.
Order/Family
Taxon*
HOMOPTERA (sucking bugs, aphids, scale insects) [continued]
Coccidae
Saissetia coffeae (Walker, 1852)
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x
Saissetia hemisphaericum
Pseudococcidae
Saissetia nigra (Nietner, 1861)
Saissetia oleae (Bernard, 1782)
Hemiberlesia lataniae (Signoret, 1869)
Lepidosaphes esakii Takahashi, 1939
Icerya aegyptiaca (Douglas, 1890)
Bulbostethus transversalis Ruckes, 1963
Geotomus pygmaeus (Dallas, 1851)
Piezodorus hybneri (Gmelin)
Dysmicoccus cocotis (Maskell, 1890)
Psyllidae
Ferrisia virgata (Cockerell, 1893)
Trioza guama Caldwell, 1942
Diaspididae
Monophlebidae
Pentatomidae
x
x
x
x
x
x
x
x
x
x
Dysmicoccus saipanensis (Siraiwa, 1933)
x
x
x
x
x
Trioza propria Tuthill, 1951
Scutelleridae
Coleotichus breddini Shouteden, 1905
HYMENOPTERA (wasps, bees, ants)
Agaonidae
Ceratosolen sp.
Liporrhopalum? sp.
Sycoscapter sp.
Aphelinidae
Aneristus ceroplastae Howard, 1895
Apidae
Apis mellifera Linnaeus, 1758
Bethylidae
Scleroderma sp.
Braconidae
Apanteles sp.
Doryctes sp.
Phaneratoma sp.
Elasmidae
Elasmus sp.
Eucharidae
Chalcura upeensis Fullaway, 1913
Eulophidae
Hemiptarsenus varicornis (Girault, 1913)
Tetrastichinae Gen. sp.
Eumenidae
Delta esuriens (Fabricius) ssp.
Gen. sp.
Rhynchium brunneum (Fabricius, 1793)
Subancisatrocerus sp.
Evaniidae
Evania appendigaster (Linnaeus, 1758)
Formicidae
Anoplolepis gracilipes (Smith, 1857)
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Anoplolepis longipes Jerdon, 1851
Camponotus chloroticus Emery, 1897
Cardiocondyla kagutsuchi Terayama, 1999
Cardiocondyla obscurior Wheeler, 1929
Cardiocondyla tjibodana Karavaiev, 1935
Hypoponera punctatissima (Roger, 1859)
Iridomyrmex anceps (Roger, 1863)
Monomorium australicum Forel, 1907
Monomorium chinense Santschi, 1925
Monomorium destructor (Jerdon, 1851)
Monomorium floricola Jerdon, 1851
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
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HYMENOPTERA (wasps, bees, ants) [continued]
Formicidae
Monomorium sechellense Emery, 1894
x
Monomorium fossulatum Wilson & Taylor, 1967
Nylanderia bourbonica (Forel, 1886)
Odontomachus simillimus (Smith, 1858)
x
x
x
x
Odontomachus haemotodus (Linnaeus, 1758), misid.
Paratrechina longicornis (Latreille, 1802)
Pheidole sp. B
Pheidole nindi Mann, 1919
Pheidole umbonata Mayr, 1870
Platythyrea parallela (Smith, 1859)
Strumigenys emmae (Emery, 1890)
x
x
x
x
x
x
x
x
x
Quadristruma emmae (Emery, 1890)
Halictidae
Ichneumonidae
Ichneumonidae
Tapinoma melanocephalum (Fabricius, 1793)
Tapinoma sp.
Technomyrmex difficilis Forel, 1892
Tetramorium bicarinatum Nylander, 1846
Tetramorium lanuginosum Mayr, 1870
Tetramorium simillimum (Smith, 1851)
Tetramorium smithi Mayr, 1879
Tetramorium tonganum Mayr, 1870
Homalictus vexator (Krombein, 1950)
Echthromorpha agrestoria insidiator (Smith, 1863)
Echthromorpha agrestoria conopleura Krieger, 1909
Megachilidae
Trathala flavoorbitalis (Cameron, 1907)
Heriades paganensis Yasumatsu, 1942
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Echthromorpha conopleura Krieger, 1909
x
x
Eriades sp.
Mymaridae
Pompilidae
Scelionidae
Sphecidae
Megachile laticeps Smith, 1853
Megachile fullawayi Cockerell, 1914
Gen. sp.
Paracyphononyx pedestris (Smith, 1855)
Scelio sp.
Chalybion bengalense (Dahlbom, 1845)
Liris aurulenta (Fabricius, 1787)
x
x
x
x
x
x
x
x
Liris opulenta (Le Peletier, 1845)
Liris aurata (Fabricius, 1787)
Motes manilae (Ashmead, 1905)
Sceliphron nr. laetum Smith, 1856
Trypoxylon nr. thaianum Tsuneki, 1961
Trichogrammatidae Trichogramma sp.
Vespidae
Delta esuriens (Fabricius, 1787)
Odynerus paganensis Yasumatsu, 1945
x
x
x
x
x
x
x
x
x
Odynerus mariannensis Bequaert & Yasumatsu
Odynerus haemorrhoidalis quinquecinctus (Fabr.)
Pachyodynerus nasidens (Latreille, 1832)
Polistes olivaceus (de Geer, 1773)
Ropalidia marginata sundaica van der Vecht, 1941
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
ISOPODA (sow bugs, pill bugs, wood lice)
Armadillidae
Gen. sp. 1
Fam.
Gen. sp. 2
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x
x
ISOPTERA (termites)
Rhinotermitidae
Prorhinotermes inopinatus Silvestri
LEPIDOPTERA (moths, butterflies)
Cosmopterygidae
Asymphorodes sp.
Gen. sp.
Crambidae
Autocharis sp.
Geometridae
Chloroclystis scintillata Prout, 1932
Scopula homodoxa Meyrick, 1886
Hesperiidae
Badamia exclamationis (Fabricius, 1775)
Lycaenidae
Petrelaea dana (de Niceville, 1884)
Noctuidae
Achaea serva (Fabricius, 1775)
Callopistria maillardi (Guenée, 1862)
Earias ochrophylla Turner, 1902
Melanitis leda (Linnaeus, 1758) ssp.
Mocis fragalis (Fabricius, 1775)
Mocis undata (Fabricius, 1775)
Nymphalidae
Euploea eunice (Quoy & Gaimard, 1824)
Hypolimas bolina (Linnaeus, 1758)
Papilionidae
Papilio polytes Linnaeus, 1758 ssp.
Papilio xuthus Linnaeus, 1767
Pterophoridae
Gen. sp.
Pyralidae
Cryptolabes sp.
Endotricha sp.
Etiella nr. grisea Hampson, 1903
Tineidae
Erecthias sp.
MANTODEA (mantids)
Mantidae
Orthodera burmeisteri Wood-Mason, 1889
NEUROPTERA (lacewings, ant lions)
Chrysopidae
Brinckochrysa scelestes (Banks, 1911)
Chrysoperla externa (Haqen, 1861)
Chrysoperla krakatauensis Tsukaguchi, 1988
Mallada basalis Walker, 1852
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Chrysopa basalis Walker
Plesiochrysa oceanica (Walker, 1852)
x
Chrysopa oceanica Walker
Hemerobiidae
Myrmeleontidae
Gen. sp.
Distoleon bistrigatus (Rambur, 1842)
Myrmeleon sp.
ODONATA (dragonflies, damselflies)
Agrionidae
Ischnura aurora (Brauer, 1865)
Libellulidae
Diplacodes bipunctatus (Brauer, 1865)
x
x
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
ODONATA (dragonflies, damselflies) [continued]
Libellulidae
Macrodiplax cora (Brauer, 1867)
Rhyothemis regia chalcoptilon (Brauer, 1867)
ORTHOPTERA (crickets, grasshoppers)
Acrididae
Aiolopus thalassinus dubius Willemse, 1923
Aiolopus thalassinus tamulus (Fabricius, 1781)
Cyrtacanthridinae Gen. sp. 1
Cyrtacanthridinae Gen. sp. 2
Cyrtacanthridinae Gen. sp. 3
Locusta migratoria manilensis (Mayen, 1835)
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x
x
x
x
x
x
x
x
x
x
x
Locusta migratoria
Conocephalidae
Gryllidae
Myrmecophilidae
Tetrigidae
Tettigoniidae
Valanga excavata (Stål, 1861)
Valanga nigricornis nr. ssp. carolinensis
Euconocephalus nasutus (Thunberg, 1815)
Gryllodes sigillatus (Walker, 1859)
Gryllus sp.
Teleogryllus oceanicus (Le Guillou, 1841)
Myrmecophilus leei Kistner & Chong, 2007
Gen. sp. 1
Gen. sp. 2
Mecopoda elongata (Linnaeus, 1758)
PHASMATODEA (walking sticks)
Phasmatidae
Acanthograeffea denticulata (Redtenbacher, 1908)
PSEUDOSCORPIONIDA (pseudoscorpions)
Atemnidae
Oratemnus samoanus
Chernetidae
Smeringochernes guamensis Beier, 1957
Tyrannochthonius chamarro Chamberlin, 1947
Olpiidae
Beierolpium oceanicum (With, 1907)
Undetermined
Gen. sp.
PSOCOPTERA (book lice, bark lice)
Archipsocidae
Archipsocus sp.
Caeciliidae
Caecilius analis Banks, 1931
Caecilius sp. nr. marianus Thornton et al., 1972
Ectopsocidae
Ectopsocus sp.
Lepidopsocidae
Lepidopsocus sp.
Myopsocidae
Phlodotes sp.
Peripsocidae
Peripsocus sp.
Pseudocaciliidae
Pseudocaecilius sp.
Psocidae
Ptycta sp.
SCORPIONIDA (scorpions)
Ischnuridae
Liocheles australasiae (Fabricius, 1775)
Fam.
Gen. sp.
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Evenhuis et al. — Pagan Island Arthropod Survey
Order/Family
Taxon*
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SIPHONAPTERA (fleas)
Pulicidae
Ctenocephalides felis felis (Bouché, 1835)
x
THYSANOPTERA (thrips)
Phlaeothripidae
Allothrips megacephala Hood, 1908
Allothrips sp.
Phlaeothripinae Gen. sp.
Dexiothrips madrasensis (Ananthakrishnan, 1964)
x
x
x
x
THYSANURA (bristletails, silverfish)
Fam.
Gen. sp.
TOTALS
Grand total of species known from Pagan -- 416
x
188
*Taxa listed in boldface are endemic species known only from Pagan Island.
288
Evenhuis et al. — Pagan Island Arthropod Survey
APPENDIX III. CORY CAMPORA FIELD NOTES
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Evenhuis et al. — Pagan Island Arthropod Survey
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Evenhuis et al. — Pagan Island Arthropod Survey
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Evenhuis et al. — Pagan Island Arthropod Survey
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