Annual Report - Panay Eco-Social Conservation Project

Transcription

Panay Eco-Social Conservation Project
(PanayCon)
Eighteenth Annual Report
January 2015
E. Curio (ed.)
PanayCon, Pandan Public Library, Pandan, Antique, Philippines
P.B. Box 42, Kalibo, Aklan 5600, Philippines
[email protected]
Under the umbrella of the NGO PhilinCon
In close cooperation with
Department of Environment and Natural Resources (Philippines)
University of the Philippines, Diliman, Quezon City (Philippines)
Frankfurt Zoological Society (Germany)
Erwin-Warth-Stiftung (Germany)
Ruhr-University Bochum (Germany)
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Frontispiece (overleaf): Front of our new T-shirt printed in 2015
Texts in English, Tagalog (Filipino) and Kinaray-a (local language spoken in Antique
Province, Panay)
From top to bottom:
From left to right: Philippine Eagle (Pithecophaga jefferyi). - Dulangan [Writhed-billed
Hornbill] (Rhabdotorrhinus waldeni, syn. Aceros waldeni) male. – The Philippine
Archipelago.
Boy with Salakot.
Spotted Deer (Rusa alfredi) male. – Banaue rice terraces. – Bayanihan spirit.
Rafflesia lobata, one of nine Philippine endemics. – Green Sea Turtle (Chelonia mydas)
Opposite: Back of T-shirt
From the living to the dead - extinction is for ever
Artwork by Helga S c h u l z e (Bochum); production of the t-shirt as a kind donation by
Claus S u d h o f f (Manila).
Impressum:
The eighteenth Report of PanayCon builds on contributions from
Curio, Eberhard
Dioso, Leocadio F.
Ebon Jr., Armelito
Faustino, Guillermo
Kühn-van Geldern, Rabea
Sanchez Jr., Enrique
Santillan, Rhea
Schwarz, Christian J.
and was edited by E. Curio
© PanayCon: no part of this report must be used without the written permission of the
PanayCon Mangement or the BOD of PhilinCon.
Pandan and Bochum, January 2016
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Acknowledgments
The PanayCon (earlier PESCP) received funding in 2015
Thanks
to the sponsors under the umbrella of the NGO PhilinCon
-
CAPE Foundation, President Macrina P. Lovina, Makati, Manila, Philippines
Prof. Dr. E. Curio, Ruhr-Universität Bochum, Bochum, Germany
Dr. Claudia Distler, Associate Professor, Owl Association Society, Ruhr-University Bochum,
Bochum, Germany
-
Mr. Antonio de Dios, President and General Manager of Transport Equipment
Corporation, Quezon City, Philippines
-
Mr. Jean-Michel Dujuypoo, Jardine Zoologique Tropical, La Londe-les-Maures, France
Erwin-Warth-Stiftung, President Hilde Stühlinger, Stuttgart, Germany
Frankfurt Zoological Society, Frankfurt, Germany
Mr. Georg Gewers, Architect, GPAI, GmbH Berlin, Berlin, Germany
Mohamed bin Zayed Species Conservation Fund, Abu Dhabi, United Arabian Emirates
Mrs. Helga Schulze, Ruhr-Universität Bochum, Bochum, Germany
Vogelschutz-Komitee (Bird Protection Comitee), President Prof. Dr. E. Schneider, Göttingen,
Niedersachsen, and Linum, Mark Brandenburg, Germany
In kind:
- Prof. Dr. Ursula Henke, Evangelische Fachhochschule, Bochum, Germany
- IDEXX Laboratories, Branch Germany, Ludwigsburg
- Leocadio F. Dioso, Director of the Leocadio Alonsagay Dioso Memorial Public
Library, Pandan, Antique, Philippines
- Richard Perron, Manager of Quantum Conversation, Oldenburg, Germany
- Dr. Jochen Reiter, Zoo Duisburg, Duisburg, Germany
- Helga Schulze, M.Sc., who is with Ruhr-University Bochum and Quantum
Conservation, gave freely of her priceless skills and time in designing and/or
perfecting the graphic presentation of PanayCon publications and reports
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Eighteenth Report 2015
An Update and Thorough Revision of the ‘Seventeenth Report 2014’
Title of Project and Time Period:
Panay Eco-Social Conservation Project (PanayCon). The time period covers the year 2015.
The project’s work is formalised under the aegis of a Memorandum of Agreement between
the DENR and Ruhr-University Bochum, renewed in 2012. A collecting permit, covering
collecting (plants, ectoparasites), and potential prey of the Marine Toad or Cane Toad
(Rhinella [Bufo] marinus), locally known also as ‘Hawaiian Frog’, is in the making. An a
dement with opportunistically obtained specimens (e.g. road kills) that represent new species
or a new distributional record in the pipeline. - Links with many environmentally concerned
agencies/ institutions are continuing to thrive and many others are developing: Erwin-WarthStiftung, President Hilde Stühlinger, and the CAPE Foundation, President Macrina P. Lovina,
absolutely vital for the project; the CAPE Foundation is aiming at costal development
including the instruction of fisher men and the conservation of marine wildlife. Furthermore,
the Foundation is no perspicacious of sponsoring six Forest Rangers that had educated as
WEOs (Wildlife Environmental Officers). Their delicate Jobs recall for arming them and
necogenations are underway mitigate their oftentimes risky situations. - And further
befriended supporter of the project is Leocadio F. Dioso. He hosts us by providing office
space in the Leocadio Alonsagay Dioso Memorial Public Library, Pandan, Antique,
Philippines. Furthermore, talks has been finalized with the University of the Philippines,
Diliman, Quezon City, alongside Los Baños by hiring Prof. Dr. E. Curio as Visiting
Professor to lecture, give seminars and supervising of Philippine Bachelor and Masters
Students.
In the 15th report the plans of the filinalisation of the project were detailed, i. e. the stepping
down on foreign personal to give way to key personal being Filipinos. The plans receive a
heavy blow when the management, including a German, found to for responsible for
embezzlement of funds. The ensuing chaos made three quarters of the staff jobless while the
segment of the reha and research facilities (8 people) by funds regenerated by Prof. Curio
could be remained until now. The case against is worn-doers to come to ends preliminarily.
The FZS recover almost completely its budget, had fired to the management staff under the
suspicion of embezzlement of their fund. – The project was victimized by two members of the
staff, incidence occurring at the station and the other serviced receiving during ascent to the
station. The theft concerned money. The victims were Ms. Santillan, our bookkeeper, and the
other incidence was theft sustained by Prof. Curio. In the latter case, the suspect admitted in
the theft during in a meeting in the brgy. hall of Bulanao, but failed to return the money and
disappeared in the day which he was supposed to bring back the stolen money. The
management is determined to follow up and retrieve the money by all legal means.
In the wake of this upheaval consultations led to an organization of staff as a result of which
emerged PhilinCon with the project PanayCon. In 2014, a new manager was hired leading a
new organizational structure (App. 1).
The Station ‘Sibaliw’ saw many visitors as usual (App. 2). For the first time members of the
University of the Philippines, led by Prof. Dr. M. Amante, were welcome by station staff. A venue
hammered out details for Prof. Curio joining the staff at Campus Diliman as Visiting Professor.
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Sad to say, BioCon, an NGO and an earlier outgrowth of PESCP (now PanayCon), failed all
along with its mandate of strengthening PESCP financially since its establishment 14 years
ago. Accordingly the founding of a new NGO PhilConserve by concerned citizens in 2005
laid the ground for fostering the hope for effective biodiversity conservation in Panay;
preparations toward fund-raising have been promoted by members of the BOD. To advance
on within-country support by non-governmental bodies born out through an on-going
correspondence with the BOD of BioCon, now in always entering in 14th year, so a far
without success. In the course of the ensuing correspondence latter of Prof. Curio never got an
answer the president of BioCon.
PanayCon gratefully acknowledges again the factual and moral support received from the
LGU of Pandan. I take this opportunity to extend my deep-felt gratitude to Hon. Jonathan D.
Tan, Municipal Mayor of Pandan, and the Head of the Pandan Department of Agriculture, Mr.
Ronald S. Sanchez, for their great understanding and perspicacity of giving leeway to their
staff in assisting PanayCon tremendously in its zeal of pushing its and the municipality’s
environment agenda. Accordingly I am pleased to mention the assistance of Mr. Arnold
Demegillo, Pandan’s MENRO and Agricultural Technologist, who took pains in advising
PanayCon in community liaison matters facilitating various technical problems. Mr.
Demegillo was promoted Head of the Municipal Disaster Risk Reduction Office.
As before, Prof. Dr. E. Schneider, President of the German ‘Bird Protection Commitee‘
(Göttingen, Linum), was circumspectly funding our ex situ work focused on the rehabilitation
and release of wildlife, specially endangered birds. Likewise, Mr. Antonio de Dios, President
and General Manager of Birds International, Inc., and of Transport Equipment Corporation,
and Mr. Georg Gewers, a Berlin architect, long term supporters of the project, donated funds
usual. However, among the European supporters diversion of funds going to the vast numbers
of refugees come from Near East and the Balkans were noticeable in the past year. This
competition, based one-sided in charity thinking, give the project a hard time.
To all these people and institutions we are deeply grateful and hope that they will support the
cause of both PanayCon and its umbrella NGO PhilinCon also in the future.
Editorial
The Philippines is hit hard by typhoons that increase in frequency and strength. There are on
average 20 typhoons every year, of which almost half make landfalls. These typhoons reach
wind speeds of 315 kph, spawning mega-storms like the 2013 typhoon ‘Yolanda’
(international name: Haiyan).
The damages are piling up, and yet, no governmental, comprehensive force has been put in
place to stem these natural calamities. Did I say ‘natural’? The incidence and strength are
natural, leaving aside climate change as a crucial determinant of those properties; however,
the severity of each single typhoon is man-made and is the primary cause of typhoons making
landfall. The severity can be measured as well: typhoon ‘Lando’ (international name: Koppu),
the latest, and according to a 68-year-old witness, the strongest one, is telling. It ravaged
Central and Northern Luzon, causing, according to governmental estimates, damages
amounting to PHP 9 billion (Euros 176 million), mostly affecting agricultural lands and
products. Also affected were 1.2 million persons, and the widespread flooding destroyed
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homes and roads. Even as far away as Manila, classes had to be suspended and flights
cancelled.
Among the most affected were the rice farms of Nueva Ecija, a situation which, according to
forecasts by the NEDA (National Economic and Development Authority), will triple the usual
amount of rice that the country needs to import to cover nationwide needs.
The rural population has been hit hardest. The extent of the damage to the vegetable and rice
fields, among other effects, have made the poorest segment of the population even poorer.
Already unable in the past to build decent homes, this group is now faced with a worse
situation: The demand for construction timber will increase and so will illegal logging. This
then is a two-pronged vicious circle: a need to generate timber for scale and construction
reduces the remaining forest cover need to stem flash-floods, causing the strength of the
typhoons to be man-made. This is one of the factors driving the calamity. Illegal logging is
the other driver. It plays in the hands of corruption in the lower ranks of bureaucracy and it
causes the loggers to become criminals, since the nature of logging is illegal.
The ‘natural’ disasters are man-made, due to the increasing denudation of the forests. If the
government wishes to address the country’s poverty problem, disasters will continue to cause
casualties and damages to property and preparing for typhoons will shrink to a minimum.
With an increase in the severity of typhoons, prospects are adverse for a program preparing
the country to more ‘natural’ disasters to come. The severity of typhoons is certainly a
consequence of climate change, but, also certainly, part of the problem is man-made and –
almost – avoidable through proper law-enforcement. (Source, in part, ‘Philippine Daily
Inquirer’.)
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Eighteenth Annual Report 2015. An Update and thorough Revision
of the ‘Seventeenth Report 2014’……………………………………….. 5
Editorial .................................................................................................................... 6
Executive Summary .................................................................................................9
1.
1.1
1.2
1.3
Conservation, Education, Rehabilitation ................................................. 11
Manager’s Report ......................................................................................... 11
Law enforcement report ................................................................................ 13
Animals under the care of PhilinCon in January 2014…………………… 26
2.
2.1
Conservation Research: Taxonomy and Biodiversity………………… 32
Gambaquezonia curioi – a new species of Gambaquezonia for the
Phlippine island Panay (Araneae: Salticidae) .............................................. 32
The praying mantids of Panay – an annotated checklist………………….. 32
2.2
3.
3.1
3.2
3.3
Behavioural Ecology…………………………..........................................40
Ecology of a Bark Mantis (Haania sp., Mantodea)……………………….40
Eucharitid ant-parasitiod affects facultative ant-plant Leea manillensis:
top-down effects through three trophic levels……………………………. 45
Flight styles in some passerines…………………………………………...45
Appendices 1- 5: Overview………………………………………………………46
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Executive Summary
Conservation and Rehabilitation
1.1 Manager’s Report – Law Enforcement
The law enforcement portion of PhilinCon’s activities mainly involves the work and
accomplishments of our team of Forest Rangers (FRs). In recognition of their effectiveness,
they have been officially deputized as WEOs (Wildlife Enforcement Officers) by the
Department of Environment and Natural Resources (DENR).
Among their standard functions, our FRs carry out regular patrols of the forest areas of
Libertad, Pandan and Sebaste in Antique and the municipality of Buruanga in Aklan province.
During 2014(?), their work resulted, in 13 separate occasions, in the confiscation of more than
135 pieces of logs that had been cut illegally. These were subsequently turned over to the
PNP/ DENR.
The FRs’ monitoring activities also resulted in the discovery of a poachers’ camp in the
Sibaliw area (near PhilinCon’s field station), from which CAMP our FRs were able to
confiscate various paraphernalia used by the poachers.. These people belonged to a wellorganized and heavily armed group from Nabas and Cubai. They were known to hunt for
Warty Pigs, monkeys, hornbills and pigeons for food and for sale to the souvenir tradesmen in
Boracay.
Sadly, the arrests and material confiscations that have been effected by PhilinCon and its FRs
have not led to the illegal loggers, hunters and poachers being brought to justice in a court of
law. Despite this fact, however, PhilinCon will continue its activities for the essential
protection and conservation of our forests and its wildlife inhabitants
1.2 Animals under the Care of PhilinCon in 2014
Both species of hornbills (Aceros [further on called this way instead of the modern name
Rhabdotorrhinus], Penolopides), various raptors, a Grass Owl and a Hawksbill Turtle
(Dermochelys imbricata) were rehabilitated and released upon proper health checks by our
veterinarian Dr. Enrique Sanchez. The rescue facilities from the animals came from in those
in Mag-aba, Bulanao and Station ‘Sibalew’. The Table 1 gives the details of the species, sex,
ring no. of applicable and the circumstances of the recovery and the release. Animals
classified as ‘unsuitable for release’ are sent to the PAWD DENR Region 6 in Iloilo City.
2. Conservation Research: Taxonomy and Biodiversity
2.1 Gambaquezonia curioi – a new Species of Gambaquezonia from the Philippine Island
Panay (Araneae: Salticidae)
A jumping spider of the genus Gambaquezonia has been considered monotypic and endemic
to the Philippine Island of Luzon. A new species has been described from Panay, based on
genital characters of a male and female specimen. The species, Gambaquezonia curioi, is
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distinguished by the male copulatory organ (embolus) and the female sperm receiving
opening (epigynal sperm duct). The find highlights the need to further study of the Philippine
spider fauna.
2.2 The praying mantids of Panay – an annotated checklist
Praying mantids (Mantodea) are an insect order which is largely understudied in the
Philippines. The ~2,500 described species are thermophilous and strictly solitary predators
distributed mainly in the tropics. They exhibit a variety of lifestyles and morphological
ecotypes. No records from Panay are known so far. Therefore, from 2010 to 2015 the
mantodean fauna of Panay has been repeatedly sampled via manual search, light and pitfall
trapping. Seventeen species representing seven phylogenetic units (families) have been found,
at least five of which are new to science. The majority (11 species) is confined to forests, and
contains mostly endemic and subendemic taxa, while six species widespread in the
Philippines and SE Asia prefer open habitats. The impoverished mantodean fauna of Panay is
discussed with reference to the paleogeography of the island.
3. Behavioural Ecology
3.1 Ecology of the Bark Mantis (Haania sp., Mantodea)
The SE Asian genus Haania contains morphologically highly specialized praying mantids
living on mossy tree bark. Two undescribed species occur on Panay, the more abundant of
which was studied from February to March at Sibaliw station. Habitat requirements comprise
trees of at least 32 cm circumference, hit by the sun and with a moss cover of ~81%. Adults
preferred lower parts of a tree than did nymphs. 91% of nymphs and 84% of adults assumed
the geotropic position. Specimens usually rested on the lee side of the tree and outside of
direct sunlight, and at minimum distances of 14 cm (adults) and 5 cm (nymphs) of keeping
inside an inhabited patch of moss. Haania is a typical ambush species, spending 67% (adults)
vs. 96.7% (nymphs) of the time motionless. Territoriality is weakly expressed in nymphs, but
becomes more pronounced when specimens approach maturity. Ten translocation trials have
been performed, all of which revealed the resident mantid to be dominant over the intruder,
even involving cases of cannibalism. The obtained results are discussed with reference to
other mantodean taxa.
3.2 Eucharitid Ant-parasitoid Effects facultative Ant-plant Leea manilllensis: top-down
Effects through three trophic Levels
Facultative ant-plant mutualisms could often hard to detect, especially in tropical ecosystems.
Leea manillensis in Panay is indirectly protected against damage by the production of
extrafloral nectaries that attract ants. Unexpectedly an ant-parasitoid wasp (Chalcura sp.,
Eucharitidae) exerts a strong effect on the system, both on the plants and on the ants well. The
parasitoid altered the behaviour of the interacting ant-species, but also directly and indirectly
affected the plants’ fitness. The study gives an example of how top-down effects alter species
interactions and can have a massive effect on mutualisms and their beneficial outcome.
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3.3 Flight Styles in some Passerines
Semi-quantitative observations on bounding flight and steady (flapping) flight are performed
in two forest-dwelling passerines (Philippine Bulbul Hypsipetes philippinus, Great Tit Parus
major) in free flight. A long-standing hypothesis predicts bounding flight to preserve energy.
A crucial role in the respect is played for the bounding flight – speed relationship. In is
controversial whether a U-shaped bounding flight – energy expenditure curve best describes
the relationship. The monotony of a new relationship reflecting bounding flight to speed
argues for a flattened speed to J-shape energy expenditure as compared to a U-shape curve.
Whether bounding flight is saving energy as proposed would depend of the kinematics of
flight in free-flying such species such as the Bulbul and the Tit. - Further observations
tentatively assign a training role for ‘mock’ prey capture and ‘group acrobatics’ of the Bulbul
serving prey capture and predator evasion, respectively.
1.
Conservation, Education, Rehabilitation
1.1 Manager’s Report
by Christian J. Schwarz, MSc., Project Manager
The most pressing problem I had to deal with when I took over this position in July 2014 was
the lack of proper funding. When compared to our activities in the financially stable years
2000 to 2009, this shortage of funds had a negative impact on conservation efforts on Panay
in particular, and the Philippines in general.
In fact, our activities are now restricted to wildlife rehabilitation, forest monitoring, and
scientific research, all of them basically restricted to the Northwest Panay Peninsula and some
portions of southern Pandan. PanayCon had to pull out completely from the Central Panay
Mountain Range, including the Dulungan or Writhed-Billed Hornbill (Aceros waldeni,
recently reassigned to the genus Rhabdotorrhinus) nest-guarding scheme. Since 2009,
poaching of nestlings of this species, as well as of other birds, snaring of Warty Pigs, and
illegal logging have increased substantially in both areas, as discovered by our forest rangers,
and experienced by myself during three years of research on the NWPP.
Conservation in the NWPP Natural Park is now basically a matter of the Protected Area
Management Board (PAMB), headed by the Regional Executive Director of the DENR, but
mainly under the supervision of Protected Area Superintendent Rhodel Lababit. The PAMB
decides over long-term activities in the Protected Area; this concerns PanayCon’s scientific
and conservation research as well as large-scale, more or less sustainable “development”
projects, like mining activities or wind turbines. The declaration of the NWPP as a PA under
the NIPAS act in 2005 has still not passed the Congress. However, this delay allowed for
adjustment of the initial, very coarse boundary of the PA to include additional areas of good
forest. The area of the PA now encompasses 13.000 ha. The true extent of forest is unknown,
but probably exceeds 6.000 ha. Despite this success, the PAMB is mainly a decision-making
organ. It is not properly equipped or funded to actually implement conservation activities.
Therefore, forest monitoring still largely relies on a core team of six Philincon FRs. These
FRs are funded by our partner, CAPE Foundation, headed by Macrina P. Lovina (Makati,
Manila). Without CAPE’s support, no forest monitoring activities whatsoever would be
conducted on the NWPP.
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Illegal logging is well-organized now: the operators are usually armed, use silencers on their
chainsaws, and informants at strategic points warn them when FR or PNP activities are
detected. This makes it extremely difficult (and dangerous) for our small team to effectively
intercept logging or hauling activities. Nevertheless, we can account for several successful
apprehensions of illegally cut lumber (see below). Animal poaching is much more difficult to
stop. Finding unreported snares is basically a matter of chance, and poachers using guns are
mobile and difficult to corner. Besides, local people are much more willing to report timber
than animal poaching. Poachers also roam in the environs of Sibaliw station, as evidenced by
discoveries of snares, animal remains, sheds, and other indications of the poachers being
active in the area. We have always duly reported such discoveries to the DENR. Towards the
end of 2014, our rangers have also found evidence of an organized team of around 6-8
poachers, mainly from Cubai and Nabas, carrying firearms and snare equipment, and hunting
for Tarictic Hornbills, monkeys, and Warty Pigs. While warty pigs are usually hunted for
meat, their tusks as well as the monkey skulls and the Tarictic beaks and legs are sold to
unknown buyers in Boracay. There, they are converted into souvenirs and sold to unknowing
tourists.
On the positive side, we are receiving an
increasing amount of information on
stranded sea turtles and their nests, and in
2014 were able to release two adults in
the process. Several nests, some of them
already successfully hatched, have also
been found. This shows that past
awareness campaigns conducted by the
DENR, the CAPE foundation, and us,
have proved useful. There is no reason to
tap our shoulders though, since we still
get to know of turtles which have been
illegally slaughtered by Pandan and
Libertad fishermen.The number of
donated or confiscated Dulungans rose
from zero between 2004 and 2011 (during
the nest-guarding scheme) to four since
2012, indicating increased poaching
Confiscated male Dulungan in the Mag-aba facility
pressure on this critically endangered
hornbill species. Our nest-guarding
scheme proved very successful, both in terms of providing data that allowed an estimate of
the population size on Panay, as well as protecting this population. However, it was also
criticized as “unsustainable” by a development program evaluator. Such comments bypass
biological reality: development programs have to be sustainable; conservation programs do
not! Their success is only measured in terms of survival of the targeted species. This example
shows how much damage to conservation can be done by recommendations of biologically
untrained professionals in highly influential positions.
When I returned to the Philippines in July 2014, most damages caused by supertyphoon
Yolanda in November 2013 had already been fixed. However, Sibaliw station, including its
solar power system, still required some attention. The humid climate takes its toll on technical
equipment. Major repairs have been scheduled to take place in 2015, and the logistics and
paper work imposed by this task have been discussed and planned towards the end of 2014.
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It also became evident, even during my previous stays, that the food catering arrangements for
the station and the organization of porterages had suffered from increasing inefficiency over
the years, both in terms of money wasted and of quality of the food supplies purchased. My
first duty over the course of the next weeks was to increase efficiency and lessen unnecessary
wastage by re-organizing the Sibaliw food supply system. Weekly supplies are now
controlled by PhilinCon office staff, basically consisting of Ms. Rhea Santillan and myself.
The new measures have already proved successful, but leaves still room for further
improvements.
Before closing, I would like to acknowledge the continued generosity of our supporters. Mr.
Antonio de Dios, Manila, donated a significant amount to Philincon, as in the years before.
This is also true for Macrina P. Lovina, CAPE foundation, who sponsored our FR team.
David Wynne and family, Alexandria VA, USA, visited our project and repeatedly donated
valuable materials. A major donation by Hilde Stühlinger, Erwin Warth-Stiftung, Stuttgart,
financed the project manager position. Without such support, our work would not be possible.
The following months have been devoted to project proposals and further fund-raising.
Several ideas were discussed in the Philincon Board Meeting on October 15. Obtaining
release permits for three of our Dulungans and some raptors, and the renovation of the station
are also on the agenda. There is no rest in conservation.
1.2 Law enforcement Report
by E. Sanchez Jr., C. J. Schwarz, R. Santillan, A. Ebon, and F. Guillermo
Introduction
The implementation of “law enforcement activity” is one of the main sub-projects of
PhilinCon, administered by its operational arm PanayCon. The sub-project workers are
otherwise known as “Bantay Gubat” or Forest Rangers/Guards. To legalize the operations
undertaken by our forest rangers against the illegal activities destroying our forests, the
rangers are duly deputized as WEO (Wildlife Enforcement Officer) by the office of the
DENR Region 6. The WEO is renewed every year, and valid in the municipalities of Libertad,
Pandan, and Sebaste. In contrast to adequately funded WEOs in previous years, our FR team
currently comprises only six full-time rangers. Additional rangers may join the team on a
daily basis, if necessary. Despite being a small team and allotted with limited allowances, our
FRs account for several successful apprehensions outlined below.
Background
In the year 2001 the first and at the time only forest guard was hired to act as a so-called
“forest monitor”. When his efforts were recognized as a success, forest monitoring was added
as one of the top priorities among the project’s objectives, in order to help our conservation
partner, the DENR, to protect the last lowland forest of the North West Panay Peninsula. The
number of FRs was later increased to 18. Due to their profound knowledge, we hired former
hunters, timber poachers and other people previously engaged in other illegal activities
harming our precious forests in the NWPP and the CPMR. By acting as Forest Rangers, they
could receive an income and bring in their experience without negatively affecting the forest.
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Unfortunately, in the last quarter of 2009, almost all rangers lost their jobs due to
mismanagement by previous project members, eventually leading to the pull-out of a major
sponsor. Despite receiving no payment, our FRs continued to undertake operations on a
voluntary basis.
Achievements 2014
Last year, our rangers had 48 operations, undertaken in the municipalities of Pandan, Sebaste
and Libertad (Antique), and Buruanga (Aklan), respectively. Illegal logging and poaching
were documented 13 times. On nine instances, a total of over 135 (!) pieces or logs of
illegally cut trees could be apprehended and turned over to the PNP/DENR. Notable is the
canter stopped with the help of the PNP on June 8 at Centro Norte, Pandan, loaded with 66
pieces of illegally cut Laua-an and Libtog timber. A matter of great concern are also the
paraphernalia confiscated at a poacher camp close to Sibaliw station. They belonged to a well
organized and heavily armed group of six to eight poachers from Nabas, Aklan and Cubai,
Libertad, who are regularly hunting warty pigs, monkeys and hornbills for supplying the
souvenir trade in Boracay.
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Table 1. Forest and wildlife monitoring successes: Upper row: turtle nest monitoring at San
Andres in December 2014. Middle left: illegally cut timber apprehended in February 2013.
Middle right to lower right: Poacher camp in the NWPPNP. Middle right: cooking place.
Lower left: snaring equipment. Lower right: shotgun ammo and feathers of poached tarictic
hornbill.
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Forest ranger activities in the year 2014.
DATE AND
TIME
LOCATION
ACTIVITY
FOREST
RANGERS
/WEOs
INVOLVED
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Ibanez, Cerwin
Matinong, Jose
Dujali, Joven
January 04,
2014
7:00 P.M.
Brgy. San Andres, Pandan,
Antique
Foot patrolling
/monitoring
January 06,
2014
1:00 P.M.
Brgy Talisay, Pandan,
Antique
Monitoring
Confiscation/
Apprehension
January 08,
2014
8:00 A.M.
Antique
Monitoring
Confiscation
Apprehension
Matinong, Jose
Guillermo, Faustino
10:00 A.M.
PhilinCon Office
L.A. Dioso Public Library
Bldg.
Brgy. Baybay, Pandan,
Antique
Orientation on
Wildlife
Enforcement
Officers
January 16,
2014
9:00 A.M.
Brgy. San Andres, Pandan
Antique,
Follow-up
monitoring
January 21-22,
2014
7:00 A.M.
Sibaliw Research Station
NWPP
Foot patrol
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Fernandez, Freddie
Dujali, Joven
Santillan, Rhea
Domingo, Rudy
Manga, Joeman
Ibanez, Cerwin
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
Dujali, Joven
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
REMARKS
The team did not find
any signs of illegal
activities during an
overnight operation
After a close
monitoring, 5 pieces of
Dao timber with the
dimensions of 2x4x16
were successfully
confiscated by the PNP
Pandan, headed by
Chief PI Jose Partisala.
After a follow-up
monitoring together
with PNP Pandan,
headed by SPO1
Ranchez Barsubia and
company, the team
found Narra trees that
had been logged
illegally:
1 pc. timber 16x20x7
1 pc. log 26x30x7
1 pc. log 24x28x7
WEOs IDs were given
to PhilinCon WEO staff
by DENR personnel in
the persons of For.
Amadona Rana and
Ms. Selma Joy Barcival.
The team confirmed that
the logged Narra were
still there.
The team did not find
activities, despite
previously received
17
Domingo, Rudy
February 13-14,
2014
6:00 P.M.
Brgy. Duyong, Pandan,
Antique
Monitoring
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Dujali,Joven
February 26,
2014
6:00 P.M.
Brgy. Nauring, Pandan,
Antique
Monitoring
Guillermo, Faustino
Ebon, Armelito Jr,
Matinong, Jose
Dujali, Joven,
Domingo, Rudy
March 7-9, 2014
8:00 A.M.
Brgy. Maramig to Sibaliw
Research Statiion
(NWPP-PA)
Conduct survey
on the uprooted/
damaged trees
caused by
typhoon Yolanda
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
Domino, Rudy
Mangga, Joeman
March 11, 2014
7:00 P.M.
Antique
Foot patrolling/
monitoring
Follow-up
Monitoring
March 14, 2014
6:30 P.M.
Mt. Muntili, Brgy. Duyong,
Pandan,
Antique
Patrolling,
Monitoring
Confiscation
Ebon, Armelito Jr.
Faustino, Guillermo
Matinong, Jose
Dujali, Joven
Ebon, Armelito Jr.
Guillermo, Faustino
Domingo, Rudy
Fernandez, Freddie
reports about illegal
hunters.
During an overnight
monitoring, the team
found an Ughayan tree
that had been cut down
and sliced into timber.
The timber had already
been delivered.
Despite received reports
on an illegal boat hull
made of a Mugni tree,
during an overnight
monitoring the team
could not confirm the
delivery of the boat hull.
The team found and
listed different species
of uprooted trees caused
by Yolanda: 3 Red
Laua-an, 2 Malaboyo, 1
Baid, 1 Natu, and 2
Malakbakan
After a whole night of
monitoring, the team did
not find any Mugni boat
hull, despite a
previously received
report from a concerned
citizen.
Instead, together with
PNP Pandan, the team
saw a tricycle with a
temporary plate No.
064409, owned by Mr.
Roque Carino of Brgy.
Fragante, loaded with
timber to be delivered to
a certain Mr. Hernani
Baraca of Brgy.
Nauring.
2 pieces of Narra
timbers with the
dimensions of 2x8x7
were found by the team
together with the PNP
Pandan along the road
18
March 15, 2014
7:00 P.M.
Antique
Patrolling
Monitoring
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
April 5-6, 2014
7:00 P.M.
Sitio Tabay, Brgy. Patria,
Pandan, Antique
Foot patrolling
monitoring
Ebon, Armelito Jr.
Matinong, Jose
April 7-8, 2014
7:00 P.M.
Antique
Foot Patrolling
Monitoring
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
Dujali, Joven
Domingo, Rudy
Fernandez, Freddie
April 14-15,
2014
8:00 P.M.
Northwest Panay Peninsula
(Sibaliw Research Station)
Foot patrolling
Monitoring
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Fernandez, Freddie
Dujali, Joven
to Mt. Muntili. The
retrieved timbers were
brought to the PNP
station for proper
custody.
from an informant about
a boat hull made of
Mugni being about to be
delivered from Brgy.
Fragante, passing
Pukatod River, down to
Nauring River, the team
did not find this
reported hull during a
whole night of
monitoring. So, a
follow-up monitoring
will be conducted.
Despite previously
received reports about
Laua-an timber to be
delivered from Mt.
Bantulinao, Sitio San
Juan, Brgy. San Roque,
after two nights of
operation the team did
not find any traces of
illegally cut timber.
Because of a report
from a concerned citizen
about Narra timber to be
delivered from Mt.
Montili (between the
Boundary of Brgy.
Duyong and Brgy.
Tingib), the team
conducted an overnight
monitoring but did not
find any signs of illegal
activities.
On the first day of
monitoring the team
camped overnight at Mt.
Jubo and Mt. Tabyaka.
During an overnight
foot patrolling no signs
of illegal activities could
19
April 30, 2014
10:00 A.M.
Antique
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
June 08-09,
2014
6:00 P.M.
Sitio Calabanog, Brgy.
Idiacacan to Brgy. Nauhon,
Sebaste, Antique
Monitoring
Apprehension
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
Domingo, Rudy
Frenandez, Freddie
June 12-14,
2014
8:00 P.M.
Protected Area
Monitoring
Foot patrol
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
Dujali, Joven
Fernandez, Freddie
Domingo, Rudy
be detected.
On the second day the
team passed by the
Station and went down
to Bulanao together with
Prof. Eberhard Curio.
Monitoring was
conducted after
information was
received from an
informant through PNP
Chief of Police, Jose
Partisala. However, the
team found out that
mahogany had been cut
instead of the reported
Narra tree.
A white elf canter with
plate No. POD 845
loaded with 66 fletches
of both Laua-an and
Libtog timber was
apprehended at Centro
Norte, Pandan, Antique,
through the help of the
PNP Pandan, headed by
their Officer-in-Charge,
PI Bryan Alamo.
It turned out that the
drivers had been Mr.
Bernie Elegino y
Agravante and Mr. Bien
Berte y Lomugdang,
both residents of Idio,
Sebaste. The owner of
the timber was Mr.
Oliver Berte y
Lomugdang.
The team heard a gun
fired at Mt. Liktinon,
and heard it again on the
following day. While
searching the area where
the gun had been fired,
the team saw 2 hunters
with a home-made
shotgun and a 22 caliber
handgun. These two
20
June 15, 2014
7:30 P.M.
Brgy. Tingib to Brgy. Patria,
Pandan, Antique
Foot Patrolling/
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Fernandez, Freddie
July 12-13, 2014
8:00 A.M.
Brgy. Barusbus to Brgy.
Inyawan, Libertad, Pandan,
Atique
Foot Patrol
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
Domingo, Rudy
Fernandez, Freddie
July 19, 2014
7:00 A.M.
Antique
Monitoring
Guillermo, Faustino
Ebon, Armelito Jr.
Fernandez, Freddie
Matinong, Jose
Domingo, Rudy
July 19, 2014
7:30 P.M.
Antique
Foot patrolling
Follow-up
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Fernandez, Freddie
Dujali, Joven
hunters were seen going
back to their area at
Sitio San Juan, Brgy.
San Roque, Libertad,
Antique.
Because of previously
received reports from a
concerned citizen that
Narra timber was to be
delivered in Brgy.
Patria, the team
coordinated with Brgy.
Capt. Hon. Carlito
Garzon of Brgy.Tingib,
in order to stay at his
area, since it was a
possible pathway of the
Narra delivery.
However, the team
found nothing after
follow-up monitoring.
Despite of received
reports about
timber to be delivered,
during an overnight
monitoring the team did
not find any signs of
illegal activities.
through a phone call
about ongoing cutting of
Gmelina trees within a
timberland area (DUPA
Site), the following
monitoring did not
produce any Gmelina
timber, nor had a
chainsaw been heard
operating.
Follow-up
monitoring/Foot
patrolling was
conducted. The team
found only 2 pieces of
2x8x8 Gmelina timber
left of the 500 board feet
of Gmelina timber that
had already been
21
July 19, 2014
7:00 P.M.
Brgy. Dumrog, Pandan,
Antique
Foot patrol
monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Fernandez, Freddie
Matinong, Jose
August 02-03,
2014
6:00 A.M.
Brgy. Fragante to Brgy.
San Andres, Pandan, Atique
Foot Patrol
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
Domingo, Rudy
Fernandez, Freddie
August 06, 2014
7:00 P.M.
Brgy. Sto. Rosario, Pandan,
Antique
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
Domingo, Rudy
Fernandez, Freddie
August 6, 2014
9:00 A.M.
Brgy. Zaldivar, Pandan,
Antique
Rescue and
release of a
hawksbill turtle
Dr. Sanchez,
Enrique D. Jr.
delivered. It was owned
by DUPA Chairman Mr.
Rhodel Lamigas. Based
on our investigation,
Mr. Demetrio Francisco
of Brgy. Tingib had
been the chainsaw
operator. It could be
confirmed that it had not
been the first time of
illegal cutting of trees at
the DUPA site.
on Laua-an timber, the
team did not find any
signs of illegal activities
during an overnight
patrol.
Despite information
received from a
concerned citizen
regarding some illegally
sliced Narra timber to
be delivered, the team
did not find any
sightings of illegal
activities after two
nights of monitoring/
patrolling.
Despite of reports from
a concerned citizen
about sliced Mugis
timber in various
dimensions about to be
delivered, the team did
not find any sightings of
illegal activities after a
whole night of
monitoring/patrolling.
On August 6, 2014, a
hawksbill turtle was
captured by fishermen
from Brgy. Zaldivar, but
soon after turned over to
PhilinCon in the person
of Dr. Sanchez Jr.. After
2 injections of treatment
22
August 13-15,
2014
7:00 P.M.
Brgy. Barusbus, Libertad,
Pandan, Antique
and w/in the NWPP-PA
Foot patrol
monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Tenorio, Joeserey
Domingo, Rudy
August 17, 2014
8:30 A.M.
Brgy. Dumrog, Pandan,
Antique
Foot patrol
monitoring that
led to a
successful
apprehension.
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Dujali, Joven
Domingo, Rudy
Fernandez, Freddie
August 18, 2014
6:00 P.M.
Brgy. Centro Norte, Pandan,
Antique
Monitoring
Apprehension
Santillan, Rhea
Guillermo, Faustino
August 19, 2014
5:00 A.M.
Mt. Tawidwid, Brgy. Luhod
Bayang, Pandan, Antique
Monitoring
Matinong, Jose
Ebon, Armelito Jr.
August 25, 2014
3:20 A.M.
Sitio San Juan, Brgy. San
Roque, Libertad to Brgy.
Patria, Pandan, Antique
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
with antibacterial and
fat soluble vitamins, the
said turtle was released
on August 8.
After a whole night of
monitoring operations
from Mt. Bulalis via Mt.
Palhe to Mt. Tabyaka,
the team did not find
activities in the area.
After valuable
information from a
concerned citizen, the
following operation led
to a very successful
apprehension of 17
pieces of Natu timber
with a total of 196 bd.
ft.. Full police assistance
was provided by the
Office of PNP Pandan,
headed by PNP Insp.
Bryan Alamo Chief of
Police/Station
Commander.
33 pcs. = 342.6 bd. ft. of
illegally cut and
transported Mugis tree
lumber were
apprehended at the
police checkpoint
conducted by the PNP
Pandan, headed by PNP
Insp. Bryan Alamo. The
information was
gathered by the two
WEOs listed in this
report.
No sign of illegal
activities at the area
could be detected during
a whole night of
Despite information
received from a
concerned citizen
regarding illegal
23
September 5-6,
2014
8:00 P.M.
Mt. Igpangi, Brgy. Magaba, Pandan, Antique
Monitoring
September 10,
2014
7:00 P.M.
NWPP-PA
Monitoring
9:00 P.M.
Roque, Libertad to Brgy.
Brgy. Sto. Rosario to Brgy.
Mag-aba, Pandan, Antique
Monitoring
September 1516, 2014
8:00 P.M.
Sitio Kaligdon, Brgy. Magaba, Pandan, Antique
Monitoring
5:00 P.M.
Mt. Talangban, Brgy. Magaba to Brgy. Tingib, Pandan,
Antique
Foot Patrol
Monitoring
September 232014
10:00 A.M.
Brgy. Sto. Rosario,
Pandan, Antique
Monitoring
Ebon, Armelito Jr.
Matinong, Jose
7:00 P.M.
Brgy. Tingib, Pandan,
Antique
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
September 12,
2014
7:00 P.M.
Monitoring
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Fernandez, Freddie
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
Ebon, Armelito Jr.
Matinong, Jose
Guillermo, Faustino
Domingo, Rudy
timber to be delivered
from Sitio San Juan to
Sebaste, the team did
not find any traces of
illegal activities.
No sign of illegal
activities in the area
could be detected after a
whole night of
about illegal cutting of
timber at night-time, the
team did not find any
sighting of illegal
activities during an
overnight monitoring.
Despite information
received from a
concerned citizen
regarding the illegal
slicing of a Mugni tree,
the team found out
during monitoring that
only coconut trees were
at the reported area.
Despite information
received from a
concerned citizen
regarding illegal
timber to be delivered
from Brgy. Tingib to
Pandan, the team did not
detect any illegal
24
October 1, 2014
8:00 P.M.
Brgy. San Roque, Libertad,
Antique
Monitoring
Ebon, Armelito Jr.
Matinong, Jose
Guillermo, Faustino
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Guillermo, Faustino
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
October 15,
2014
3:00 P.M.
Roque, Libertad, Antique
Monitoring
October 20-21,
2014
9:00 A.M.
Sitio Atabay, Brgy. Patria,
Pandan to Sitio San Juan,
Libertad, Antique
Foot patrol
Monitoring
October 23,
2014
10:00 P.M.
Sitio San Juan, Libertad,
Antique
Monitoring
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Santillan, Rhea
October 25-26,
2014
4:00 P.M.
Idiacacan, Pandan, Ant.
Foot patrol
Monitoring
Ebon, Armelito Jr.
Matinong, Jose
Domingo, Rudy
Guillermo, Faustino
Fernandez, Freddie
November 0305, 2014
1:00 P.M.
Brgy. Nauhon, Sebaste,
Antique
Monitoring
Foot patrol
Ebon, Armelito Jr.
Matinong, Jose
Guillermo, Faustino
Domingo, Rudy
November 1213, 2014
9:00 A.M.
Brgy. Candari, Brgy. Sto.
Rosario to Brgy. Guia
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
Fernandez, Freddie
November 21,
2014
Brgy. Mag-aba, Pandan,
Antique
General Clean up
Ebon, Armelito Jr.
Guillermo, Faustino
activities during their
monitoring.
Monitoring conducted
after request by PASU
Rhodel Lababit
During an overnight
monitoring the team did
not find a reported
illegally cut Kamagong
tree.
Despite received
information regarding a
white elf delivering
illegal timber from Sitio
San Juan to North
Sebaste, no vehicle
could be spotted during
overnight monitoring.
The team conducted a
foot patrol to Mt.
Kabuluan, where a boat
hull was reported to be
located, but found
nothing after an
overnight operation.
Despite information
received from a
concerned citizen
regarding the illegal
poaching of timber,
intended to be used as
boat hull, the team
found nothing after 2
days of monitoring/foot
patrol.
During an overnight
monitoring conducted
by the team, there were
no sightings of
previously reported
electro-fishing in the
area.
General cleaning of
Mag-aba Rescue and
25
9:00 A.M.
Matinong, Jose
Domingo, Rudy
Fernandez, Freddie
November 2527, 2014
8:00 A.M.
Protected Area
Foot patrol
Monitoring
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Domingo, Rudy
Fernandez, Freddie
Mangga, Joeman
December 0910, 2014
7:00 P.M
Idiacacan to Brgy. Patria,
Pandan, Antique
Monitoring
Foot patrol
Ebon, Armelito Jr.
Matinong, Jose
Guillermo, Faustino
Domingo, Rudy
Fernandez, Freddie
Santillan, Rhea
Rehabilitation Facility
in preparation of an
upcoming visit of
tourists.
When the team reached
Palhe Cave, they found
21 pieces of ropes, 1
cooking pot and 4
plates. These
paraphernalia were
presumably owned by
illegal hunters active in
the area. Targeted
species had obviously
been wild pigs and
endangered birds, esp.
Tarictics.
The team started to
conduct a turtle nest
monitoring from Sitio
Calabanog down to
Brgy. Patria. A
concerned citizen
provided information on
a potential nest at
around 6:30 in the
morning. At around 10
P.M. the team arrived at
said nesting site at Brgy.
San Andres, but the
turtles had already
hatched successfully a
month ago. In order to
confirm the information,
the team started to dig
out the nest and found
shells of newly hatched
turtles. Mr. Ricky
Tamboong showed us 2
rescued newly hatched
turtles which were about
2 weeks old. We asked
him why he did not
inform us in time, and
he explained that he had
informed their fish
warden, Mr. Bobby
Tindog, who informed
26
December 25,
2014
9:30 A.M
Brgy. Cabugao, Pandan,
Antique
Monitoring
Apprehension
Ebon, Armelito Jr.
Guillermo, Faustino
Matinong, Jose
Santillan, Rhea
December 2627, 2014
9:00 A.M
Antique
Monitoring
Ebon, Armelito Jr.
Matinong, Jose
Guillermo, Faustino
Domingo, Rudy
Fernandez, Freddie
neither the project nor
the office of the
MENRO.
In response to
information received
about illegal cutting of a
Narra tree, the team
apprehended, together
with the PNP Pandan
headed by Chief PI
Brian Alamo, some
Narra timber owned by
Mr. Aokly Alvaro. The
timber is now under
custody of Brgy. Capt.
Benito Rubite.
The team found 2
Tipolo trees that had
been already cut down,
and 1 Mugni tree which
had been already sliced
into timber and hauled
away. The 2 Tipolo
have not yet been
finished because the
chainsaw operator had
an accident while
cutting it.
1.3 Animals under the care of PhilinCon Year 2014
by E. Sanchez Jr., DVM
Sibaliw
Admission
Date/Origin
Brgy. Alegre, Sebaste ,
Antique
2004
Animal
Species
Dulungan
(Aceros
waldeni)
Sex
Ring#
Remarks
male
0031
Slated for release target date January
of 2015
June 2000
Brgy. Idiacacan, Pandan,
Antique
May 2002
sent up to Sibaliw 2008
2 Dulungans
(Aceros waldeni)
Female
Female
0035
0036
Slated for release target date is January
of 2015.
Tarictic
(Penelopides
Female
0033
27
June 6, 2007
Brgy. Bagumbayan,
Pandan , Antique
May 24, 2007
Sebaste, Antique
sent up to Sibaliw
September 21, 2010
panini)
Tarictic
(Penelopides
panini )
Taricitc
(Penelopides
panini)
Tarictic
(Penelopides
panini)
September 18, 2012
Sitio Bulabog, Brgy.
Balabag, Boracay Island,
Malay, Aklan
Dulungan
(Aceros waldeni)
(immature)
June 20, 2014
2 Dulungans
(Aceros waldeni)
(immature)
Female
0151
Male
104
Female
0164
Mortality due to
super-typhoon
Yolanda in November
2013
Surrendered to
DENR, CENRO
PAWCZMS Boracay
through PhilinCon
by Mr. Toby Mockel,
and facilitated by Mr.
Jason Probyn and Del
Nano, owners of
Nigui-Nigui Resort,
Boracay Island , and
Mr. Jun Aguirre from
the Philippine News
Agency (PNA). The
hornbill has been sent
to Sibaliw Station
after 60 days of
quarantine in Bulanao
Reha Facility.
Surrendered by Mr. J.
Amar of Brgy.
Importante, Mun. of
Tibiao, Antique ,
through the efforts of
Tibiao Fish Spa
owner Nick Calawag
of Brgy. Malabor.
Mortality of 1 bird
due to stress caused
by environmental
factors. Turn-over to
PhilinCon in a DENR
ceremony (DENR
Caravan) in Libertad,
Antique.
Ring #
Remarks
Sent to Bulanao in
August 2011.
Mortality of 1 female
during super-typhoon
Yolanda on
Male
Female
Sent up to Sibaliw
February 11, 2009
Bulanao Rehabilitation Facility
Admission Date/Origin
July 2011
Brgy. Calabanog, Pandan,
Antique
Animal Species
3 Tarictics
(Penelopides
panini)
(nestlings)
Sex
Male
Female
Female
28
November 30, 2011
San Andres, Pandan,
Antique
Crested Serpent
Eagle
(Spilornis cheela
holospilus)
(adult)
December 20, 2011
Religious Group Rally
against mining in Antique
Province , San José Public
Plaza
2 Brahminy
Kites
(Haliastur indus)
June 20, 2014
2 Tarictics
(Penelopides
panini)
(immature)
November 8, 2013.
Admitted to Mag-aba
Reha. Information on
the whereabouts of
this bird was provided
to PhilinCon office
staff by our partner
NGO, CAPE
Foundation. The
turnover to our FR’s
has been witnessed by
the President of
CAPE, Macky
Lovina.
Mortality due to
super-typhoon
Yolanda in November
2013.
Birds were donated
for rehabilitation and
future release. 1 bird
is a mortality of
super-typhoon
Yolanda.
Turned over to
PhilinCon by the
DENR during a
ceremony in Libertad,
Antique. Surrendered
to the DENR Office
by Nick Calawag.
Mag-aba Rehabilitation Facility
Admission date/Origin
July 2007
Brgy. Pajo, Libertad,
Antique
Animal Species
Crested Serpent
Eagle
(Spilornis cheela
holospilus)
(adult)
May 2011,
Brgy. Fragante, Pandan,
Antique
Grass Owl
(Tyto capensis)
August 11, 2012
Sitio Burabod, Brgy.
3 Crows
(Corvus
Sex
Ring #
0799
Remarks
Slated for release,
targeted schedule is
January 2015.
Surrendered to DENR
through PhilinCOn by
Mr. Nonilon Dioso,
And facilitated by SB
Reynaldo Dioso and
PhilinCon WEOs’
together with
Christian Schwarz.
Slated for release,
targeted schedule is
January 2015.
Surrendered to DENR
CENRO PAWCZMS
29
Callan, Sebaste, Antique
brachyrhynchos)
September 24, 2012
Poblacion, Kalibo, Aklan
Dulungan
(Aceros waldeni)
(adult)
January 16, 2013
Honey Buzzard
(Pernis steerei)
(immature)
August 8, 2013
Visayan Spotted
Deer
(Rusa alfredi)
(juvenile)
December 7, 2013
Changeable
Hawk Eagle
(Spizaetus
chirratus)
(subadult)
Female
32442 w/
Wilhelmshaven
Vogelwarte
Helgoland
inscription
through PhilinCon by
Arnaldo Nepomuceno
& Hermie Yac-yac.
Released after a
month (30 days) of
quarantine.
Surrendered to
PhilinCon on behalf
of CENRO Kalibo by
Grace MapesoQuimpo. The bird has
probably been in
captivity for 14 years,
as estimated by the
owner. She had
inherited the bird
from her father.
Mortality due to
pectoral muscle
atrophy and bacterial
infection.
Turn-over was
facilitated by Brgy.
Captain of Baybay,
Pandan, Hon. Michael
O. Condez, Liga
President of the
Association of
Barangay Chairmans,
Pandan, Antique.
Turned over by
CENRO San José,
Antique, upon the
recommendation of
PAWCZMS Region 6
RTD Office (Carlo
Custodio). Initially
confiscated by the
members of the PNP
Mobile Group
Bugasong, Antique,
during their patrol
operations in the
hinterland barangays.
Rescued by a
fisherman from Brgy.
San Andres, Pandan,
Antique on December
2007. Said raptor
perched on his banca
and appeared weak
and tired. Its last
resort had been to
hunt in the fish caught
by the fisherman out
of his banca. The bird
30
was probably unable
to find prey, which
had probably become
very scarce due to
super-typhoon
Yolanda. Upon
capture, the fisherman
informed the MENRO
(Municipal
Environment and
Natural Resources
Officer) of Pandan,
who immediately
called the PhilinCon
Reha in Mag-aba,
Pandan.
July 2014
August 6-7, 2014
Crested Serpent
Eagle
(Spilornis cheela
holospilus)
(immature)
Hawksbill Turtle
(Eretmochelys
imbricata)
(immature)
not determined
Rehabilitation and
Release: The raptor
was brought up to
good health, and after
a necessary
quarantine period of
30-45 days it was
finally released on
January 30, through
an effort of the GIZ
ForClim Project 2014.
The release has been
documented by a TV
journalist from
Germany.
Turned over to
PhilinCon after being
received by our
FR/WEO Rudy
Domingo.
Turned over to
PhilinCon. The donor,
Mr. Jesus Patiño,
captured the said
animal because it had
been weak and floated
on the sea surface,
appearing emaciated.
Treatment regime
included antibacterial
medications and
vitamins A, D and E.
Both medications
have been
administered
intramuscularly. The
turtle was released on
August 8, 2014
31
Note: Admitted animals which are classified as “Unsuitable for release” are sent to the
PAWD DENR Region 6 in Iloilo City.
Release of Changeable Hawk-Eagle in Magaba on January 30, 2014. Left: Dr. Enrique
Sanchez, Jr. explaining Mag-aba reha facility to local and international witnesses, including a
German film team. Right: Eagle female being released.
Release of Hawksbill Turtle in Barangay Zaldivar, Pandan, on August 8, 2014.
32
2.
Conservation Research: Taxonomy and Biodiversity
2.1 Gambaquezonia curioi - a new species of Gambaquezonia from the Philippine island
Summary. - The jumping spider genus Gambaquezonia has been considered monotypic and
endemic to the Philippine Island Luzon. Here, we describe a new species from the Philippine
Island Panay, based on genital characters of a male and female specimen. The new species,
Gambaquezonia curioi, is distinguished by the shape and structure of the male embolus and
tibial apophysis and the epigynal sperm duct. Finally, we highlight the necessity of further
arachnological exporations on the Philippine Archipelago (App. 3).
2.2 The praying mantids of Panay – an annotated checklist
By Christian J. Schwarz, MSc., Project Manager
Introduction
The praying mantids (Mantodea) are a dictyopteran order of exclusively carnivorous insects,
with about 2,500 described species (SVENSON et al. 2015). All species are thermophilous and
strictly solitary predators distributed mainly in the tropics where they exhibit a variety of
lifestyles and morphological ecotypes (PRETE et al. 1999, EHRMANN 2002, SCHWARZ 2003).
Like most invertebrates, this group of insects is largely understudied in the Philippines, all
data available to date being in fact decades-old expedition collections from Luzon or
Mindanao, or without any specific locality. The most important contributions are those of
STÅL (1877), HEBARD (1920), WERNER (1922, 1926) and BEIER (1966). Preliminary
catalogues of the Philippine mantodean fauna were compiled by BRUNER (1915) and WERNER
(1926). The Western Visayas are notoriously undersampled, with only a few specimens from
Negros known in the literature (HEBARD 1920, ROY 2011). From 2010 to 2015 the author was
the first to study the Panay mantodean fauna on a regular basis. All habitat types have been
repeatedly sampled in the course of the cane toad study via manual search, light and pitfall
trapping. The mantodean results of these investigations are compiled in the following list. It is
expected to be a fairly complete account of the species occurring on the island given the fact
that no new species have been added to the list since 2012. All species are new records for the
island, several are new to science and will be described subsequently.
The systematic arrangement of the order is currently in a state of successive revision, since
recent phylogenetic studies (YAGER & SVENSON 2008, SVENSON & WHITING 2009, WIELAND
2013, LEGENDRE et al. 2015) are largely incongruent with the classic system established by
BEIER (1964) and widely used since (e. g. EHRMANN 2002, OTTE & SPEARMAN 2005).
Therefore, a revised system based on recent phylogenetic hypotheses is used here.
Information on distribution, life history, primary defense strategy, and habitat type is given
for each species. Life history strategies include arbusticolous and arboricolous ambush
hunters, hanging on the underside of leaves in rainforests or inhabiting weeds in open
habitats, and semi-cursorial bark and ground mantids (see EDMUNDS & BRUNNER 1999 and
SCHWARZ 2003 for a review). Bark mantids can be divided into several subtypes, depending
on the type of tree they inhabit and on the adaptations to this habitat. The ground mantid niche
33
is poorly represented in SE Asia (SCHWARZ & KONOPIK 2014), even more so on Panay, which
only harbors one species with mostly terricolous nymphs (see below).
With regards to the preferred habitat, the species on Panay can be roughly be grouped into
two categories. Forest-dependent species are confined to closed canopy conditions and today
restricted to primary and secondary dipterocarp forests, savaged plantations and neophyte
stands. The second group is made up of species preferring open habitats. Initially, these taxa
were confined to the upper canopy, tree fall gaps, landslide areas, and forest edges, but today
they inhabit degraded man-made habitats like agricultural field margins, scrubland, and
ruderal places. This group of mantids is naturally very rare in primary forests but was able to
profit from human activity at the expense of forest taxa.
Mantids are masters of camouflage and employ several different strategies to disguise
themselves from prey and predators. Generalist morphotypes rely on homochromy or crypsis
(background-matching color pattern). These species are often polyphonic and able to adopt
the colors and pattern of the background. Other taxa employ disruptive coloration and
morphology (e. g. lobes) to break up the body outline. However, when removed from their
habitat, members of these groups still look like edible insects. Morphologically more
specialized taxa evolved to closely resemble certain structures of the habitat, like twigs or
leaves, or mimic inedible or harmful arthropod models like ants. Their camouflage does not
rely on a matching background to be effective.
Species list
Fam. Leptomantellidae
01. Leptomantella lactea (Saussure, 1870)
- Widely distributed in SE Asia.
- Confined to forests.
- Arboricolous, generalist ambush hunter
inhabiting the canopy.
- Female exhibits a primitive form of brood care: it
stays close to its ootheca and lacks aggression
towards newly hatched nymphs.
02. Aetaella bakeri Hebard, 1920
- Endemic in the Philippines, with a close relative
in Borneo.
inhabiting the understorey and lower canopy.
- Female exhibits a primitive form of brood care: it
stays close to its ootheca and lacks aggression
towards newly hatched nymphs.
34
Fam. Nanomantidae
03. Pliacanthopus sp.
- New genus record for the Philippines, with close
relatives in Malaysia.
- Species description in progress
Fam. Gonypetidae
04. Amantis aeta Hebard, 1920
- Endemic in the Philippines, with close relatives
in SE Asia.
- Adults are generalist ambush hunters inhabiting
the understorey and lower canopy, while nymphs
are semi-cursorial and mainly terricolous (the only
Panay mantid species sampled by pitfall traps). A
terricolous or nearly terricolous lifestyle in both
nymphs and adults is ancestral for the genus, A.
aeta being an exception in this regard.
- Nymphs and adults engage in foreleg waving
(references in SCHWARZ & KONOPIK 2014). Nymphs resort to ant mimicry; later instars mimic
the aggressive trap-jaw ant Odontomachus philippinicus.
05. Compsomantis mindoroensis Beier, 1942
- Described from neighboring Mindoro, but also
common on Panay.
- Found in open habitats.
- Arbusticolous; this bark mantis subtype prefers
slender branches to sit on.
- Nymphs and adults engage in foreleg waving.
Upon approaching threats, the mantids switch
position to the opposite side of the twig.
06. gen. n. sp. n. aff. Compsomantis
- Genus and species new to science, description in
progress.
- Endemic on Panay.
- Confined to forests, but preferring sunny places
in secondary forests and at forest edges.
- Living on tree bark, prefers young trees.
- Nymphs and adults engage in foreleg waving.
Upon approaching threats, the mantids switch
35
position to the opposite side of the trunk and run up several centimeters before flattening
against the bark.
07. Theopompa sp.
- Possibly new to science; the status of this species
is currently being investigated.
- Living on tree bark, preferring the canopy of
larger trees.
- Upon an approaching threat, the mantids flatten
themselves; if threat persists, they quickly run up
the trunk.
Fam. Haaniidae
08. Haania sp. “short wing”
- New to science, description in progress.
- Endemic on Panay.
- Lives on mossy tree bark at low heights,
representing a morphologically and behaviorally
distinct subtype of bark mantids.
- The ecology of this species has been described
elsewhere (SCHWARZ 2014, KÜHN-VAN GELDERN
et al. this volume).
09. Haania sp. “long wing”
- New to science, description in progress.
- Endemic on Panay.
- Lives on mossy tree bark, representing a
morphologically and behaviorally distinct subtype
of bark mantids.
- In contrast to the preceding species, this congener
prefers dipterocarp bark heterogeneously covered
in mosses, lichens, and small vines, climbs higher
up the trees, and has a slightly divergent phenology (SCHWARZ 2014).
Fam. Hymenopodidae
10. Creobroter meleagris Stål, 1877
in SE Asia.
- Arbusticolous, generalist ambush hunter
inhabiting the inflorescences of weeds and scrubs.
36
- This species represents the “flower mantis” ecotype in the Philippines, characterized by
disruptive coloration and morphology (lobes). Most commonly found on Ageratum
conyzoides and Mikania cordata (both Asteraceae), sometimes also on Stachytarpheta
jamaicensis (Verbenaceae) and small individuals of Melastoma polyantha (Melastomataceae).
11. Odontomantis euphrosyne Stål, 1877
in SE Asia.
- Arbusticolous, generalist ambush hunter
inhabiting weeds and scrubs.
- Nymphs resort to ant mimicry, mimicking
Camponotus, Plagiolepis and similarly-looking
ants (MATHEW 1935).
12. Acromantis sp.
- The status of this species is currently being
investigated; the genus is widely distributed in SE
Asia.
- Arboricolous; on Panay this species is a generalist
ambush hunter inhabiting the canopy. Other
species may prefer weeds and scrubs in open
habitats.
- Nymphs resort to ant mimicry.
Fam. Deroplatyidae
13. Tagalomantis manillensis (Saussure, 1870)
- Endemic in the Philippines, known so far only
from Luzon and Panay; a close relative occurs on
Sulawesi.
- Redescription in progress.
- Arboricolous ambush hunter inhabiting the
understorey and lower canopy.
- This species represents the “stick mantis”
ecotype on Panay, characterized by an elongate
body and associated behavior enhancing stick
resemblance. Female exhibits brood care: it guards
its ootheca and lacks aggression towards newly
hatched nymphs.
Fam. Mantidae
14. Statilia pallida Werner, 1922
37
- Endemic in the Philippines, with close relatives in SE Asia.
- Graminicolous and arbusticolous, generalist ambush hunter inhabiting grasses and weeds.
- The species exhibits green/brown polyphenism.
15. Tenodera aridifolia (Stoll, 1813)
- Graminicolous and arbusticolous, generalist
ambush hunter inhabiting grasses and weeds.
- The species exhibits a green/brown polyphenism.
16. Hierodula patellifera (Audinet-Serville, 1839)
- Arbusticolous and arboricolous, generalist
ambush hunter inhabiting weeds and bushes.
Freshly hatched nymphs resort to ant mimicry,
mimicking the weaver ant Oecophylla
smaragdina.
17. Hierodula cf. vitreoides Giglio-Tos, 1912
- Possibly endemic to the Philippines, the status of
this species is currently being investigated.
Freshly hatched nymphs resort to ant mimicry,
mimicking the weaver ant Oecophylla
smaragdina.
Discussion
When compared with other island groups (PAICs) of the archipelago (HEBARD 1920,
WERNER 1922, 1926, BEIER 1966), the mantid fauna of Panay is rather impoverished, lacking
several SE Asian genera known to occur on Luzon or Mindanao, like Metallyticus,
Amorphoscelis, Tropidomantis, Phyllothelys, Deroplatys, Euchomenella and Mesopteryx.
This picture will probably turn out to be also roughly true for the Western Visayas as a whole,
even though Negros harbors at least two species which have not yet been recorded from
Panay (ROY 2011, pers. obs.).
Most genera are represented on Panay by only one, often endemic species, with two
exceptions. First, the speciose genus Hierodula, exemplifying the “typical” green generalist
mantid in Asia and Australasia, has two species on Panay which occupy two different niches.
38
H. patellifera occurs in disturbed habitats, while H. cf. vitreoides requires more shade and
replaces the former in primary forests. Heterogeneous secondary forests and savaged
plantations may harbor both species.
More interesting is the occurrence of two species of Haania on Panay. This mantodean group
is phylogenetically much older than the Philippine islands (SVENSON & WHITING 2009) and
turns out to have undergone a radiation event after colonizing the archipelago, comparable to
the frog genus Platymantis. Aside from the two species on Panay, at least five other
Philippine and several Sundaian species await description (Schwarz & Stiewe in prep.).
Endemism level is moderate: 5 out of the 17 species recorded from the island are endemic,
corresponding to a rate of 29%. The most unusual find is the single specimen of the new
Pliacanthopus species, the first member of this genus recorded for the Philippines. It is
possible that this or a related species occurs on other islands but has escaped detection so far.
The reasons for the low number of taxa contrasting with fair levels of endemism is found in
the relative isolation of the Greater Negros-Panay PAIC with respect to the remaining
archipelago in particular, and of the Philippines to SE Asia in general. While the Greater
Luzon and the Greater Mindanao PAIC are today separated from adjacent SE Asian mainland
(and from each other) by only one oceanic barrier, a second oceanic barrier separates the
Western Visayas, acting as an additional dispersal filter. This confirms the rather low
transoceanic dispersal abilities of mantodeans when compared to other groups (in fact it is
rather similar to that of mammals), underscored by a very low diversity on oceanic islands
(SCHWARZ & KONOPIK 2014 and references therein).
Another, maybe more important reason is the comparatively young age of the Greater NegrosPanay PAIC, which is estimated to be not older than 3.5 Ma (STEPPAN et al. 2003). Most
other Philippine islands have provided dry land for longer periods of time (HALL 2009,
STEPPAN et al. 2003, LOHMANN et al. 2011), providing colonization and persistence
opportunities. In fact, speciation in most investigated taxa seems to be older than the PAICs
created by glacial cycles and to reflect more the Miocene to Pliocene geologic history of the
archipelago than Pleistocene sea level changes (STEPPAN et al. 2003, SILER et al. 2010, 2011,
2012). For example, emergent parts of the terranes making up present-day Sulawesi were
connected with each other and with the northern Philippines via volcanic island chains until
the Middle Miocene (15 Ma). Genera like Tagalomantis must have colonized Sulawesi and
the Philippine archipelago from Sundaland by that time, since latest after 10 Ma, only the
southern Philippines (where this genus does not occur) retained an island stepping stone
connection to Sulawesi. In contrast, genera like Amorphoscelis and Deroplatys are missing
from the central and northern Philippines, including Panay. They must have colonized the
archipelago after 5 Ma, when stepping stones connected northern Borneo and Mindanao, and
Borneo and Palawan, respectively (HALL 2009, LOHMANN et al. 2011).
References
BEIER M. (1964) Blattopteroidea, Mantodea, p. 849-970 in: BRONN H. G. (ed.), Klassen und
Ordnungen des Tierreichs. Fünfter Band: Arthropoda. III Abteilung: Insecta. Geest & Portig,
Leipzig.
BEIER M. (1966) Noona Dan Papers No. 29. Die Mantiden der Noona Dan Expedition nach
den Philippinen und Bismarck Inseln. Entomologiske Meddelelser 34: 361-370, 3 pl.
39
BRUNER L. (1915) Preliminary catalogue of the orthopteroid insects of the Philippine Islands.
University Studies 15: 195-281.
EDMUNDS M. & BRUNNER D. (1999) Ethology of Defenses against Predators, p. 276-299 in:
PRETE F. R., WELLS H., WELLS P. H. & HURD L. E. (eds), The Praying Mantids. The
Johns Hopkins University Press, Baltimore & London.
EHRMANN R. (2002) Mantodea – Gottesanbeterinnen der Welt. Natur und Tier Verlag GmbH,
Münster.
HALL R. (2009) Southeast Asia’s changing palaeogeography. Blumea 54: 148-161.
HEBARD M. (1920) Studies in Malayan, Papuan, and Australian Mantidae. Proceedings of the
Academy of Natural Sciences of Philadelphia 71: 14-82, 2 pl.
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sp., Mantodea), p. 40, in: CURIO E. (ed.), Panay Eco-Social Conservation Project
(PanayCon)
Eighteenth
Annual
Report.
Available
as
download
at
www.panaycon.org/index.html.
LEGENDRE F., NEL A., SVENSON G. J., ROBILLARD T., PELLENS R. & GRANDCOLAS P. (2015)
Phylogeny of Dictyoptera: Dating the origin of cockroaches, praying mantises and
termites with molecular data and controlled fossil evidence. PLoS ONE 10 (7):
e0130127, doi:10.1371/journal.pone.0130127.
LOHMANN D. J., BRUYN M. DE, PAGE T., RINTELEN K. VON, HALL R., NG P. K. L., SHIH H.-T.,
CARVALHO G. R. & RINTELEN T. VON (2011) Biogeography of the Indo-Australian
Archipelago. Annual Review of Ecology, Evolution, and Systematics 42: 205-226.
MATHEW A. P. (1935) Transformational deceptive resemblance as seen in the life history of a
plant bug (Riptortus pedestris), and of a mantis (Euantissa pulchra). Journal of the
Bombay Natural History Society 37: 803-813, 1 pl.
OTTE D. & SPEARMAN L. (2005) Mantida species file – Catalog of the mantids of the world.
Insect Diversity Association, Publication Number 1, Philadelphia.
PRETE F. R., WELLS H., WELLS P. H. & HURD L. E. (1999) The Praying Mantids. The Johns
Hopkins University Press, Baltimore and London.
ROY R. (2011) Les Amorphoscelinae Indo-Malais (Mantodea, Amorphoscelidae). Revue
française d’Entomologie (N. S.) 32 (1-2): 65-92.
SCHWARZ C. J. (2003) Diets and habitat preferences of neotropical praying mantids
(Dictyoptera: Mantodea, Burm. 1838). Diploma thesis at the University of Würzburg.
SCHWARZ C. J. (2014) Anspruchsvolle Zwerge – Die Tarnkünstler der Gattung Haania. Bugs
6: 24-29.
SCHWARZ C. J. & KONOPIK O. (2014) An annotated checklist of the praying mantises
(Mantodea) of Borneo, including the results of the 2008 scientific
expedition to Lanjak Entimau Wildlife Sanctuary, Sarawak. Zootaxa 3797 (1):
130-168.
SILER C. D., DIESMOS A. C., ALCALA A. C. & BROWN R. M. (2010) Phylogeny and
biogeography of Philippine bent-toed geckos (Gekkonidae: Cyrtodactylus) contradict a
prevailing model of Pleistocene diversification. Molecular Phylogenetics and Evolution
55: 699-710.
SILER C. D., DIESMOS A. C., ALCALA A. C. & BROWN R. M. (2011) Phylogeny of Philippine
slender skinks (Scincidae: Brachymeles) reveals underestimated species diversity,
complex biogeographical relationships, and cryptic patterns of lineage diversification.
Molecular Phylogenetics and Evolution 59: 53-65.
SILER C. D., OAKS J. R., WELTON L. J., LINKEM C. W., SWAB J. C., DIESMOS A. C. & BROWN
R. M. (2012) Did geckos ride the Palawan raft to the Philippines? Journal of
Biogeography 39: 1217-1234.
STÅL C. (1877) Orthoptera nova ex insulis Philippinis descripsit. Öfversigt af Kongliga
Vetenskaps-Akademiens Förhandlingar 34 (10): 33-58.
40
STEPPAN S. J., ZAWADZKI C. & HEANEY L. R. (2003) Molecular phylogeny of the endemic
Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic
archipelago. Biological Journal of the Linnean Society 80: 699-715.
SVENSON G. J. & WHITING M. F. (2009) Reconstructing the origins of praying mantises
(Dictyoptera, Mantodea): the role of Gondwanan vicariance and morphological
convergence. Cladistics 25: 468-514.
SVENSON G. J., HARDY N. B., CAHILL WIGHTMAN H. M. & W IELAND F. (2015) Of flowers and
twigs: phylogenetic revision of the plant-mimicking praying mantises (Mantodea:
Empusidae and Hymenopodidae) with a new suprageneric classification. Systematic
Entomology doi: 10.1111/syen.12134.
WERNER F. (1922) Philippine mantids, or praying insects. Philippine Journal of Science 21:
147-157, 1 pl.
WERNER F. (1926) Zur Kenntnis der Mantodeen der Philippinen. Konowia 5: 227-232.
WIELAND F. (2013) The phylogenetic system of Mantodea (Insecta: Dictyoptera). Species,
Phylogeny, and Evolution 3: 1-306.
YAGER D. D. & SVENSON G. J. (2008) Patterns of praying mantis auditory system evolution
based on morphological, molecular, neurophysiological, and behavioural data.
Biological Journal of the Linnean Society 94: 541-568.
3. Behavioural Ecology
3.1 Ecology of a Bark Mantis (Haania sp., Mantodea)
By Rabea Kühn-van Geldern and Christian J. Schwarz
Introduction and Methods
Haania sp. male (left) and female (right).
1
The SE Asian genus Haania Saussure, 1871, contains morphologically highly specialized
praying mantids living on mossy tree bark. Ten species are currently known but the genus is
more diverse than previously known; several species await description. Panay harbors two
undescribed species, one of which is more abundant. Previous observations had revealed that
the behavior of Haania differs considerably from typical generalist mantids (SCHWARZ 2014),
so a more detailed study was conducted from February to March at Sibaliw station. Only one
location in the primary forest is known to sustain a population of this mantid. However, it also
occurs in the secondary forest, albeit at lower densities. The season was already advanced for
this species, nevertheless 19 specimens (5 females, 1-2 males, 2-3 subadults and ten younger
nymphs) could be observed. Observation periods lasted
↓
from 11 to 17 h. Bark mantids exhibit site fidelity
(SCHWARZ 2003), so usually a specimen could be
encountered on the same tree on consecutive days.
However, several individuals vanished during the
course of the study. The following parameters were
recorded: circumference of the inhabited tree, relative
moss cover up to 1 m height, occurrence of vines,
distance of mantid to the ground, cardinal direction,
orientation of mantid, distance of mantid from margin
of moss cover, and the direction of sun and wind with
respect to the mantid. Intraspecific interactions and
territorial behavior was also assessed by displacing
selected individuals to trees occupied by a conspecific.
Numerical values are given as mean ± SD.
←
Results and Discussion
1. Morphology and primary defenses
These mantids exhibit crypsis, and disruptive
coloration and morphology as adaptations to their
habitat. Their green ground color is mottled with dark
and light. Disruptive effects breaking up the body
outline are achieved through the lobes on head,
pronotum, and abdomen, annulated walking legs, and
two pairs of bright lateral V-like patterns above the
walking legs coxae. The walking legs are long, slender
and stalk-like, holding the body of the mantid above
the moss cover. Young nymphs are pale green and
become successively brighter green in the course of the
post-embryonic developOothecae
Haania sp. female camouflaged on
mossy bark
ment.
Oothecae are laid on
structures protruding from
the moss and are on
average 5 mm long and
2-3 mm wide and high.
In contrast to other bark
mantids (HILL et al. 2004,
SCHWARZ, this volume)
Haania does not spiral up
or down the trunk upon
an approaching threat, but
bends the body and crouches against the moss. Only
Habitat
42
when poked repeatedly they start to run away using all six pairs of legs. Adult males are good
fliers and sometimes attracted to lights.
2. Habitat requirements
Haania sp. only inhabits dry trees covered by short mosses and hit by the sun. Therefore, it
avoids most parts of the primary forest, which are too shady and too humid, and prefers steep
slopes or adjacent secondary forests. Moss cover (at 100 cm height) of the trees inhabited by
the study specimens averaged 81 ± 9%. Average circumference of inhabited trees was 94 ± 62
cm. A possible explanation for the avoidance of thin trees is that those are shaken too much
by the wind. Horizontal branches are likewise avoided.
3. Behavior
Orientation
Geotropy is strongly supported in this species, as has been in other bark mantids (e. g.
LIEFTINCK 1953, LADAU 2003, SCHWARZ 2003): 91% of nymphs and 84% of adults assumed
the geotropic position. Of the remaining, 8% of nymphs and 13% of adults chose the upright
position, while just 1-3% of specimens rested
sideways.
The adaptive significance of geotropic behavior may
be related to successful prey capture. Hunting
against the direction of the light presumably
impedes proper object recognition. Most bark
mantids observed by the second author assume the
geotropic orientation by day. In contrast, the upright
position is often assumed by night (KROMBEIN
1963). These orientation switches between day and
night have not been observed in Haania and Astape
(LIEFTINC K 1953, Schwarz pers. obs.).
Perch height
Adults prefer the lower parts of the tree, resting at
an average height of 34 ± 10 cm. In contrast,
nymphs have been found to rest at heights of 83 ±
15 cm. The maximum height an individual has been
found at was 120 cm.
Wind and sun avoidance
Almost all specimens (90% of adults and 100% of
First instar nymph
nymphs) rested on the lee side of the tree. This was
also true with regard to the position of the mantid
with respect to the sun: 97% of adults and 100% of nymphs rested outside direct sunlight.
The low heights preferred by the mantids in combination with their avoidance of excessive
sun and wind may be a consequence of their habitat requirements, defined by low substrate
humidity but high atmospheric humidity. Moss cover is more extensive at low heights in
those parts of the forest inhabited by populations of this mantid. Also, wind stress is lower
near the ground than at several meters off the ground, causing less disturbance (which equals
to energetic loss) and desiccation stress to the individual. This may also be the reason for the
localized occurrence of these mantids: long-fronded mosses in other parts of the primary
43
forest are too moist (because they dry out too slowly after rains), and impede proper
locomotion.
Distance to moss cover margin
In most cases the mantids rested directly on the moss, keeping a distance of 24 ± 10 cm
(adults) and 10 ± 5 cm (nymphs) to the margin of the moss cover. This is in concordance with
their cryptic defense strategy. Mantids are able to perceive suitable backgrounds and prefer
those in favor of non-matching resting places (reviewed in EDMUNDS & BRUNNER 1999 and
BERG et al. 2011).
Cardinal direction
There was a difference in the preferred cardinal direction between adults and nymphs: while
56% of adults rested on the southern side of the tree, with North being the second-most
preferred side (26%), nymphs preferred the northern and western sides (44% and 26%,
respectively). This difference may be a consequence of the location, since the studied nymphs
inhabited a different part of the forest with a different wind direction. Thus, the chosen resting
side seems to be not influenced by cardinal direction per se, but by other abiotic factors like
position of the sun and prevailing direction.
Activity
Haania sp. is less cursorial than other bark mantids and can be regarded as a typical ambush
species which spends most of the day motionless. For adults, this amounts to 67% of the time.
This roughly equals the amount of time spent motionless by the arboricolous ambush species
Sphodromantis lineola (ZACK 1978), and is somewhat less than the 75.3% recorded for the
bark mantis genus Liturgusa (SCHWARZ 2003). Prey capture and ingestion is a rare event,
accounting for 1% of observation time. This is much less than the values recorded by
SCHWARZ (2003) for Liturgusa (11.6%). This may be explained by the fact that the latter
genus feeds mainly on ants, which are a common prey item, in contrast to the rather rare prey
encounters most mantids are faced with.
Other activities observed in Haania are grooming (11%) and locomotion (21%). The time
required for grooming roughly corresponds to the values for other species (16.7% in S.
lineola, 11% in Liturgusa), while locomotion (not related to prey capture) is more common in
Haania. It is mostly caused by wind driving the mantid to the lee side (15%), and as such may
be a characteristic of the peculiar habitat of this species. Nymphs are more sedentary than
adults, spending 96.7% of the time motionless.
4. Intraspecific interactions
Territorial behavior was tested by removing a subadult or adult specimen from its home tree
and placing near a conspecific at about 15-20 cm. This experiment was not performed with
young nymphs, since they show a much more greater tolerance to conspecifics than is the case
in subadults and adults. In the field, several young and medium-sized nymphs may share a
tree, while this is very rarely the case with adults. Only six specimens were available for this
experiment due to the advanced season. Some specimens were used for more than one trial,
allowing 10 trials to be performed.
In all cases (100%) the resident mantid showed dominant behavior, expressed either as active
(attack, 50%) or passive (standing its ground, 50%) territory defense. The intruder fled in all
cases. The attacks of the resident mantid resulted in one intruder being killed; four trials have
been prematurely ended by the experimenter to avoid the killing of the attacked specimen. A
44
high degree of territoriality in adults had already been noted by SCHWARZ (2014) and partly
coincides with observations on another Haania species from Palawan (Schwarz unpubl.) and
on the closely related genus Astape on Java (LIEFTINCK 1953). In the latter cases, the territory
of a female was also defended against intruders, but in contrast to the Panay species, more
than one specimen may be found on a single tree.
The mantids of the genus Haania differ in several respects from typical ambush mantids
which had served as models in scientific studies so far (see PRETE et al. 1999 and BERG et al.
2011 for an overview). This study was conducted towards the end of the adult’s phenological
timeframe, so obtaining a good sample size (particularly of adult males) proved
problematical. Additional translocation experiments involving adults of both sexes and
nymphs will help to deepen our understanding of the intraspecific behavioral repertoire of this
mantid species, including sexual interaction, prey capture, and (the lack of?) territorial
aggression towards nymphs. Observation time should be extended into the night to account
for circadian shifts in activity patterns.
References
BERG M., SCHWARZ C. J. & MEHL J. E. (2011) Die Gottesanbeterin, Mantis religiosa. Die
Neue Brehm-Bücherei Bd. 656, Westarp Wissenschaften, Hohenwardsleben.
EDMUNDS M. & BRUNNER D. (1999) Ethology of Defenses against Predators, p. 276-299 in:
PRETE F. R., WELLS H., WELLS P. H. & HURD L. E. (eds), The Praying Mantids. The
Johns Hopkins University Press, Baltimore & London.
HILL P., HOLWELL G., GÖTH A. & HERBERSTEIN M. (2004) Preference for habitats with low
structural complexity in the praying mantid Ciulfina sp. (Mantidae). Acta Oecologica
26: 1-7.
KROMBEIN K. V. (1963) Behavioral notes on a Floridian mantid, Gonatista grisea (F.)
(Orthoptera, Mantidae). Entomological News 74 (1): 1-2, 1 pl.
LADAU J. (2003) Prey capture in a mantid (Gonatista grisea): Does geotropy promote
success? Canadian Journal of Zoology 81: 354-356.
LIEFTINCK M. A. (1953) Biological and ecological of a bark haunting mantid in Java
(Orthopt., Mantoidea). Transactions of the Ninth International Congress of Entomology
2: 125-134.
PRETE F. R., WELLS H., WELLS P. H. & HURD L. E. (1999) The Praying Mantids. The Johns
Hopkins University Press, Baltimore and London.
SCHWARZ C. J. (2003) Diets and habitat preferences of neotropical praying mantids
(Dictyoptera: Mantodea, Burm. 1838). Diploma thesis at the University of Würzburg.
SCHWARZ C. J. (2014) Anspruchsvolle Zwerge – Die Tarnkünstler der Gattung Haania. Bugs
6: 24-29.
SCHWARZ C. J. (this volume) The Praying Mantids of Panay – an Annotated Checklist, p. …
in: CURIO E. (ed.), Panay Eco-Social Conservation Project (PanayCon) Eighteenth
Annual Report. Available as download at www.panaycon.org/index.html.
ZACK S. (1978) Description of the behavior of praying mantis with particular reference to
grooming. Behavioural Processes 3: 97-105.
45
3.2 Eucharitid ant-parasitoid affects facultative ant-plant Leea
manillensis: top-down effects through three trophic levels
By Christoph Schwitzke, Brigitte Fiala, K. Eduard Linsenmair, Eberhard Curio
Abstract. – Facultative ant–plant mutualisms are variable systems, shaped by a number of
biotic and abiotic factors. Especially in tropical ecosystems, the generally assumed
mutualistic benefits are often hard to prove. We studied the system Leea manillensis on the
Philippine island Panay and its indirect defence mechanism against herbivory by producing
extrafloral nectar therewith attracting ants. Unexpectedly, we found an ant-parasitoid wasp
from the genus Chalcura (Eucharitidae) to have a strong influence on the system, on ants as
well as on plants. The parasitoid not only altered the behaviour of interacting ant species, but
also directly and indirectly affected the plants’ fitness. This study demonstrates how top-down
effects may alter species interactions and have a massive effect on mutualisms and their
beneficial outcome (App. 4).
3.3 Flight Styles in some Passerines
By Eberhard Curio
Abstract. – Semi-quantitative observations of the circumstances account under which
bounding flight and steady (flapping) flight are performed by two forest-dwelling passerines
of two different lineages (Pycnonotidae, Philippine Bulbul Hypsipetes philippinus; Paridae,
Great Tit Parus major) in free flight. Bounding flight is one of two intermittent flight styles,
to the exclusion of undulating flight, in passerines weighing ≤100g. A new bounding flightspeed relationship is apt to shed light on the power output of bounding flight. In line the
function found in most flying animals for metabolic energy expenditure against speed the
relationship support a flattened J-curve as compared to U-shaped curve found in some birds;
this is based on the monotony of the parameters of bounding as a function of speed. Whether
bounding flight is saving energy as proposed by the work of Rayner et al., and of Tobalske in
tunnel wind experiments, would depend to the kinematics of flight in free-flying birds such as
the Bulbul and the Tit. – Further observations demonstrate that both steady flight and
bounding flight are used in foraging and in chases of two birds of the same species, whilst
‘mock’ prey capture and ‘group acrobatics’ in the Bulbul are tentatively assigned a training
role serving prey capture and predator evasion, respectively. – Observations of Tree Sparrows
(Passer montanus) support the idea of flapping flight changing abruptly into the still phase of
bounding flight and is therefore consistent with ’body morphing’ in the Bulbul (App. 5).
46
Appendices 1 – 5: Overview
App. 1
Staff of PhilinCon
Organizational Structure and Staff
App. 2
Rhea Matining
Visitors of Research Station ‘Sibaliw’
App. 3
M. Freudenschuss,
A. Grabolle &
H. Krehenwinkel
Gambaquezonia curioi – a new Species of Gambaquezonia
from the Philippine Island Panay (Araneae: Salticidae).
Arachnology in press
App. 4
C. Schwitzke, B.
Fiala, K. E.
Linsenmair
& E. Curio
App. 5
E. Curio
Eucharitid Ant-parasitroid effects facultative Ant-plant
Leea manillensis: Top-down Effects through three trophic
Levels.
Arthropod-Plant Interactions 9: 497-505.
DOI 10.1007/s11829-015-9391-y.
Flight Styles in some Passerines.
Ecol. Birds (Ökol. Vögel) XX. 201X: 00-00, in press
Appendix 1
Organizational Stucture of PhilinCon
Appendix 2
VISITORS OF SIBALIW RESEARCH STATION
Year 2014
Name
Affiliation
Kristin Hagel
Former student of RuhrUniversitat Bochum
Paul French
United Kingdom
Rhea Santillan
PhilinCon/PanayCon staff/
WEO
Faustino Guillermo
Forest Ranger/ Wildlife
Enforcement Officer
Joven Dujali
Forest Ranger/Wildlife
Enforcement Office
Jose Matinong
Enforcement Officer
Rudy Domingo
Enforcement Officer
Armelito Ebon Jr.
Enforcement Officer
Nestor Bagac
PhilinCon/PanayCon Staff
Brendan Sloan
Local Government Officer London
Dr. Maren Gaulke
Project Herpetologist
Dr. Hendrik Freitag
Ateneo De Manila
Marivine M. Santos
National Museum of the
Philippines
Clister V. Pangantihon
National Museum of the
Philippines
Gersom Operiano
Field Assistant
Dr. Friedhelm Goeltenboth
Visayas Statte University Baybay,
Leyte
Prof. Dr. Eberhard Curio
PhilnCon/PanayCon Founder
Date
Reason for Visit
January 10, 2014
visit
January 21-24,2014
birdwatching
January 21-22,2014
Project visit, conduct monitoring
January 21-22,2014
January 21-22,2014
January 21-22,2014
January 21-22,2014
January 21-22,2014
January 21-22,2014
Feb. 15-17, 2014
Birding
Feb. 24-26, 2014
Herpetological research
Feb.24-26, 2014
Field sampling w/ Phil.National
Museum
Museum
Feb.24-26, 2014
Feb.24-26, 2014
Museum
Site establishment
April 2-4,2014
Project visit
April 3-15, 2014
Project visit
Scientific Adviser
Prof. Maragtas Amante
University of the Philippines
Diliman
Director Rene N. Rollon
Diliman
Dr. Benjamin M. Vallejo
Diliman
Jonathan M. Madrid
Mardonie Cruz
Arnel Telesforo
Panay Bird Club
Ruperto Quitag
Panay Bird Club
Emmanuel Lerona
Panay Bird Club
Frederick Arceo
Panay Bird Club
Gersom Operiano
Field Assistant
Joeman Mangga
Field Assistant/FR/WEO
Luciano Palmieri
Philippine National Museum
Collaborator
Lilian Rodriguez
Philppine Natinal Museum
Collaborator
Prashant Sharma
Collaborator
Ronald Clouse
Collaborator
Perry Buenavente
Philppine National Museum
Collaborator
David Wynne
London
Marlon Legurpa
University of Visayas
Prof. Sasha Dioso
University of Visayas Miag-ao
Frances Anthea Redison
Educator
Arthur Hortillo
April 14-15,2014
Discuss field course and MOA
With Prof. Curio
April 1-15, 2014
with Prof. Curio
April 1-15, 2014
with Prof. Curio
April 1-15, 2014
April 21-23, 2014
with Prof. Curio
with Prof. Curio
Bird watching
April 21-23, 2014
Bird watching
April 21-23, 2014
Bird watching
April 21-23, 2014
Bird watching
May 13-20, 2014
Site establishment
May 13-20, 2014
Site establishment
May 13-14, 2014
Research Voulnteer
May 13-14, 2014
Research Volunteer
May 15-17, 2014
Research Volunteer
May 15-19, 2014
Research volunteer
May 15-19, 2014
Research volunteer
July 3-4, 2014
Bird watching/ site seeing
August 7-8, 2014
Site seeing/exploring
August 7-8, 2014
August 7-8, 2014
August 27-29, 2014
Install and repair solar power
April 1-15, 2014
Solar Technician
Ryan James Posadas
Solar Technician
Lena Scheimann
PhilinCon/Ruhr Universitat
Bochum
Prof. Dr. Eberhard Curio
PhilinCon/PanayCon Founder
Scientific Adviser
Christian Schwarz, MSc.,
Project Manager/ PhD student
Lena Scheimann
PhilinCon/Ruhr Universitat
Bochum
Delizia Polli
Switzerland
Luca Hauri
Switzerland
Fernando Sierra
Worldwide Cover-Switzerland
Prepared by: Rhea A. Santillan
Project Bookkeeper
August 27-19, 2014
Install and repair solar power
Sept.9-Oct.17,2014
Study of Biology/Research
volunteer
Sept. 10-22, 2014
Project visit/ Research
October 22-27, 2014
Project visit/ research volunteer
October 30-Nov.30,
2014
Study of Biology/ Research
volunteer
November 10-11,
2014
Nov. 10-11, 2014
Site seeing/ bird watching
Nov. 16-17, 2014
Site seeing/bird watching
Site seeing/bird watching
Appendix 3
Gambaquezonia curioi - a new species of Gambaquezonia from the Philippine island
Running title: A new Philippine jumping spider species
By Mario Freudenschuss, Arno Grabolle and Henrik Krehenwinkel
Key words: Panay, taxonomy, new species, Salticidae, jumping spiders, Philippines,
biodiversity
Summary. - The jumping spider genus Gambaquezonia has been considered monotypic and
endemic to the Philippine Island Luzon. Here, we describe a new species from the Philippine
Island Panay, based on genital characters of a male and female specimen. The new species,
Gambaquezonia curioi, is distinguished by the shape and structure of the male embolus and
tibial apophysis and the epigynal sperm duct. Finally, we highlight the necessity of further
arachnological exporations on the Philippine Archipelago.
Arachnology in press.
Appendix 4
Eucharitid ant-parasitoid affects facultative
ant-plant Leea manillensis: top-down
effects through three trophic levels
Christoph Schwitzke, Brigitte Fiala,
K. Eduard Linsenmair & Eberhard Curio
Arthropod-Plant Interactions
An international journal devoted to
studies on interactions of insects, mites,
and other arthropods with plants
ISSN 1872-8855
DOI 10.1007/s11829-015-9391-y
1 23
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Author's personal copy
DOI 10.1007/s11829-015-9391-y
ORIGINAL PAPER
Eucharitid ant-parasitoid affects facultative ant-plant Leea
manillensis: top-down effects through three trophic levels
Christoph Schwitzke1 • Brigitte Fiala1 • K. Eduard Linsenmair1 • Eberhard Curio2
Received: 9 March 2015 / Accepted: 25 July 2015
Springer Science+Business Media Dordrecht 2015
Abstract Facultative ant–plant mutualisms are variable
systems, shaped by a number of biotic and abiotic factors.
Especially in tropical ecosystems, the generally assumed
mutualistic benefits are often hard to prove. We studied the
system Leea manillensis on the Philippine island Panay and
its indirect defence mechanism against herbivory by producing extrafloral nectar therewith attracting ants. Unexpectedly, we found an ant-parasitoid wasp from the genus
Chalcura (Eucharitidae) to have a strong influence on the
system, on ants as well as on plants. The parasitoid not only
altered the behaviour of interacting ant species, but also
directly and indirectly affected the plants’ fitness. This
study demonstrates how top-down effects may alter species
interactions and have a massive effect on mutualisms and
their beneficial outcome.
Keywords Chalcura Extrafloral nectaries Ants Mutualistic interaction Philippines
Handling Editor: Heikki Hokkanen.
Electronic supplementary material The online version of this
article (doi:10.1007/s11829-015-9391-y) contains supplementary
material, which is available to authorized users.
& Christoph Schwitzke
[email protected]
1
Department of Animal Ecology and Tropical Biology,
Biozentrum, Am Hubland, 97074 Würzburg, Germany
2
Conservation Biology Unit, Ruhr-Universität Bochum, ND
1/31 Universitaetsstrasse 150, 44801 Bochum, Germany
Introduction
Interactions between organisms can be very complex and
are often yet unexplored (Dáttilo et al. 2014), especially in
tropical forests. Within ecosystems, members of different
trophic levels are interacting with each other by regulatory
forces. According to their main direction and frequency,
these are described as top-down and bottom-up regulations
(e.g. Power 1992; Oksanen and Oksanen 2000; Ernest and
Brown 2001; Báez et al. 2006). The trophic cascade model
combines both forces. These forces can flow up and down
the trophic structure, and organisms on any level can have
dramatic effects on the composition of the whole community (Leibold 1989; Schmitz 1993).
Compared with investigations of predation and competition as structuring forces, studies that address the roles of
mutualism and parasitism in shaping communities are
underrepresented. Mutualistic interactions between species
are widespread in tropical forests, playing an important role
in ecosystem functioning (Thompson 2006). For instance,
ant–plant protection mutualisms have served as model
systems to study mechanisms promoting species coexistence and trophic cascades, and they are known for many
plants from different taxonomic groups (Heil and McKey
2003; Bronstein et al. 2006). Protection mutualisms can
have far-reaching effects in food webs, because they
involve not only direct (and often indirect) trophic
exchanges between the two mutualists, but also interactions
with additional trophic levels (Dyer 2008). In these associations, ants are defending a valuable food resource such
as extrafloral nectar (EFN) and thus are indirectly protecting the entire plant or plant parts, respectively, against
herbivores. Many studies demonstrated a positive effect of
EFN on the fitness of the host plant, but others failed to
find a benefit (ÓDowd and Catchpole 1983; Mody and
123
Christoph Schwitzke et al.
Linsenmair 2004; Horvitz and Schemske 1984; Bronstein
1998, do Nascimento and Del-Claro 2010). The different
results of these studies might be caused by the fact that in
general these associations are characterized by a low
specificity and high variability, with plants attracting a
broad range of ant species (Schemske 1982; Diaz-Castelazo et al. 2010) as well as other arthropods (Koptur 1992).
While ant–plant interactions have been a research focus
in different tropical regions, from the Philippines even
basic knowledge about ant–plant interactions is lacking.
And still more basically, the Philippine ant fauna is very
poorly explored (General and Alpert 2012), and an
unknown number of species has probably already been lost,
due to the fact that only about 11.2 % of primary forests
were left in 2010 through large-scale anthropogenic
destruction (FAO UN 2010). We found that the shrub Leea
manillensis (Leeaceae) offering EFN and food bodies (FB)
on its shoots was regularly visited by ants. However, none
of the hitherto performed studies could prove any benefit
derived by the plant due to the ants’ presence and their
potential protection against herbivores, but the genus Leea
has been reported to be involved in facultative mutualisms
in other parts of Southeast and East Asia (Fiala and Linsenmair 1995; Meng et al. 2011). The original aim of our
study was to investigate this yet unknown ant–plant association and its mutualistic quality. Unexpectedly, this system became complicated by a third trophic level—an antparasitoid wasp of the genus Chalcura (Hymenoptera:
Eucharitidae: Eucharitinae). This wasp turned out to be
abundant and to have a massive impact on plants as well as
on ants.
Parasitoids are important organisms and abundant
members of nearly all terrestrial communities, essentially
contributing to the high biodiversity of tropical ecosystems
(Godfray 2007). Their species richness (and therefore
possible functional relevance) has been markedly underestimated in the tropics (see, e.g. Veijalainen et al. 2012;
Hrcek et al. 2013). The Eucharitidae is the only insect
family known to comprise only parasitoids specialized on
ant brood (Heraty 1985; Heraty and Darling 1984;
reviewed in Lachaud and Pèrez-Lachaud 2012). All
members of this family, where the life cycle is known,
develop as koinobiont, larval–pupal ectoparasitoids
(Lachaud and Pèrez-Lachaud 2012). Females place their
eggs away from the host on plant tissue. The active firstinstar larvae (planidia) are responsible for gaining access to
the larval ant host via various phoretic behaviours,
involving attachment to ant workers or intermediate hosts
carried into the colony by ant workers (Heraty 2000). Ants
tend to intensively forage on plants, especially when a food
source as honeydew or EFN is given (Blüthgen and Fiedler
2004). Pérez-Lachaud et al. (2006) gave a description of
the life cycle of Kapala iridicolor (Eucharitidae) from the
123
Neotropics, which oviposits on undeveloped flower buds of
Melampodium divaricatum (Asteraceae), possessing
extrafloral nectaries at the abaxial side of its leaves.
Observations of Carey et al. (2012) suggest that the parasitoid Orasema simulatrix (Eucharitidae) oviposits almost
exclusively in the near vicinity of the extrafloral nectaries
of Chilopsis linearis (Bigoniaceae), ensuring to gain access
to an ant colony. But so far, no attention has been paid to
the parasitoids‘ influence on the whole system and on
possible effects of the protective function of the ants for the
plants.
Up to now, studies on ant–plant protection mutualisms
mediated through a plant-derived food source have usually
only regarded direct interactions between ants and herbivorous insects (but see, e.g. Rudgers 2004; Pires and DelClaro 2014 and references therein). However, as our study
shows, interactions can be much more complex. To test
whether Leea manillensis is indeed involved in mutualistic
interactions with ants, we studied food production, ant
visitors, and their possible benefits for the plant. As a
further trophic level was involved in our study system, we
also needed to analyse the biology of the ant-parasitoid and
its interference with the ant–plant relationship. In this
study, we report about the parasitoid, its impact on the
fitness of Leea manillensis, and the entire ant–plant system
and show the effects propagated through three different
trophic levels.
Materials and methods
Study site
The study site was located in the midwestern Philippines on
the island Panay, within the ‘Northwest Panay Peninsula
Natural Park’ in the north of the island, an area of mountainous secondary and primary lowland rainforest around the
research station ‘Sibaliw’ (11490 9.7600 N, 121570 37.9500 E).
The station was established by the Philippine NGO ‘PhilinCon’ and Eberhard Curio in 1997 on an abandoned settlement area, with the youngest secondary forest being about
35 years old. The study was carried out in secondary as well
as primary forest habitats.
Study species
Leea manillensis (Leeaceae) is a large erect shrub with
height up to 6 m, terminal inflorescences, and large stipular
structures that enclose the next generation of shoots. It is
distributed widespread throughout the Philippines and parts
of Southeast Asia (Molina et al. 2013) and was formerly
included in L. guineensis (Ridsdale 1976). The species is
shade-tolerant, growing in the understory, as well as in
Eucharitid ant-parasitoid affects facultative ant-plant Leea manillensis…
forest gaps (and edges) in primary and secondary forests,
with especially high abundances in bright, young-aged
secondary growth forests. Photographs of Leea manillensis
are shown in Online Resource 1.
We checked the plants’ shoots for the occurrence of
extrafloral nectaries and food bodies, ants, or any other
visitors. Nectar sugar content was measured after 5-h
exclusion of ants from nectaries. Ant exclusion was
achieved via the application of grease at the basis of the
shoot. Droplets were taken with a glass capillary (KG01, A.
Hartenstein laboratory supplies: 100 9 0.9 mm, wall
thickness 0.1 mm), and sugar content in per cent was
measured using a hand-held refractometer (Hand-Held
Refractometer RHB-32ATC).
Parasitoid wasps
As we found an ant-parasitoid wasp from the genus
Chalcura (Eucharitidae) to have a strong influence on the
system, all the experimental shoots were regularly surveyed for wasps, planidia, and any abnormalities in growth
or habitus. Planidia were taken to the field station in vials
for further observations and photographs.
The parasitoid wasp was determined to genus level by
Stefan Schmidt at the Bavarian Zoological State Collection, Munich (ZSM), and voucher specimens of wasps
(females, males, planidia) were deposited at the ZSM.
The Eucharitidae (Chalcidoidea) is a small family with a
worldwide distribution and divided into three subfamilies
Oraseminae, Eucharitinae, and Gollumielinae. All of its
members, where the host is known, parasitize ant brood
(Pérez-Lachaud et al. 2006; Lachaud and Perez-Lachaud
2012). Species are found in all zoogeographical regions,
but most are confined to the tropics. Eucharitids attack ants
in at least 21 genera, distributed across the subfamilies
Formicinae, Myrmeciinae, Myrmicinae, and Ponerinae
(Heraty 1994). Some genera show a very broad host range,
with species found on several different genera, or even on
different subfamilies of ants. Most genera, however, are
restricted to a single genus of ants (Heraty 1994).
Experiments on herbivory
The study was conducted in 2011 during 3 months of the
rainy season from 15 August to 21 November. The initial
aim was to study the system Leea manillensis and its
defence mechanism against herbivory by producing
extrafloral nectar (EFN) therewith attracting ants. A total of
223 plants were found in the study area—primary and
secondary forests—and marked individually. Of these, 213
shoots were used for a short-term experiment on herbivory.
All of these plants were monitored regularly. With a shoot
expanding, the main experiment began, and plants were
monitored visually every 3 days for a period of 21 days.
We recorded the presence or absence of ants, ant species,
and abundance on the shoot, as well as the number of other
arthropods feeding on the plant tissue or at the EFN. After
the 21-day period, the shoots were cut, dried between
sheets of paper, and photographed (Canon Digital IXUS
50) on scale paper for further analyses related to leaf area
loss due to herbivory.
To analyse leaves for possible ant herbivore defence, leaf
area loss was measured for each shoot using Photoshop
(Adobe Photoshop CS5 Extended, version 12.0.4 9 64).
Leaf area loss was expressed as percentage of missing leaf
area as well as lost area in cm2. As leaves do not shrink by
more than 5 % during the process of drying, data from dried
leaves give a reliable estimate of missing leaf area (Heil
2004). A complete loss of the shoot due to herbivory, which
was a frequent incidence, was recorded as 100 % leaf area
loss. Shoots with only intermittent ant association were not
used for further experiments on herbivory.
Results
Leea manillensis and its associations with ants
The plant produced FB and EFN on its shoots that were
mainly visited by different ant species, but also by a variety
of other arthropods. Up to eight elongated extrafloral nectaries are found on the shoots’ stipule and two to six elongated extrafloral nectaries at the nodes above the stipule
(Online Resource 1). When the young leaves become
mature, the extrafloral nectaries stop nectar production. All
of the shoots possessed nectary glands. However, production
was very variable as many shoots did not produce FB or
EFN at all. Food bodies usually occurred only in the first
days after shoot expansion and predominantly around the
shoots’ stipule and on the internode above. We assume the
production of EFN and FB to be dependent on a combination of abiotic and biotic factors, but our study did not
emphasize any clear pattern of dependency in that respect.
Plant size did not play an important role, as we observed
even seedlings to produce EFN and attract ants. Plants
growing under low light conditions attracted ants as well as
plants growing in forest gaps (unpublished observations).
Out of 223 marked plants, a total of 80 plants were
visited by ants, whereas on 90, ants were never observed at
the shoots’ nectaries. On 53 plants, ants were only rarely
found at the nectaries. Only 36 % of the marked plants of
Leea manillensis showed a permanent association with
ants. On constantly ant-free plants also, no other visitors of
the extrafloral nectaries were observed, suggesting a general inactivity of the nectary glands. In contrast, on shoots
with regular ant occupation, droplets of EFN occurred on
123
the nectaries soon after ant exclusion. The average sugar
content of the EFN droplets after 5-h ant exclusion was
16.6 % (n = 13).
Ant activity was mainly concentrated on the biggest
nectary glands at the stipule and to a lesser extent at the
extrafloral nectaries positioned directly at the nodes.
Occasionally, ants were observed feeding on extrafloral
nectaries located at the underside of the leaves, but only on
fresh shoots or in combination with—and probably connected to—recent leaf damage.
Over the study period of 3 months, a total of 26 ant
species were observed visiting the extrafloral nectaries on
the shoots of Leea manillensis. Sixteen ant species out of
six genera showed a constant assemblage on the nectaries
(from shoot expansion until maturity of leaves), whereas
the rest visited the nectaries only temporarily. Most common, and assembling in the highest abundances with a
maximum number of 65 ants, was a single unidentified
species of the genus Dolichoderus. This species was only
found on the shoots of Leea in secondary forest. In
declining order in terms of the number of shoots occupied
during the study, we found the genera Technomyrmex (two
species), Crematogaster (five species), Tetramorium (three
species), Polyrhachis (four species,) and Camponotus (one
species). In general, about 10–20 ants assembled at the
extrafloral nectaries, except of big ants from the genera
Polyrhachis and Camponotus, where normally less than
five individuals were assembled. During heavy rain
showers, ants used to leave the shoots or seek shelter
beneath the plants’ leaves. In general, only one species
occupied a shoot at a time. Only rarely, two or three species
at a time were observed using the same food source on the
shoots. Ants of the genus Polyrhachis were the most
dominant, but also the genus Dolichoderus and species of
Crematogaster rank among the dominant species. In rare
cases of two or three ant species at a time, the dominant
species defended the nectaries against submissive species,
mostly small species of the genera Tetramorium and Crematogaster, which were frequently observed to sneak in.
Occasionally, the ant species composition on a plant
changed, sometimes even several times over the study
period. Most of the ant species associated with Leea
manillensis showed a diurnal as well as nocturnal activity
pattern. Species of the genus Polyrhachis showed a strictly
diurnal activity, while the single species of the genus
Camponotus was observed only at night-time. Some photographs of Leea manillensis and associated ant species are
shown in Online Resource 1.
Further arthropods
During the daily monitoring, besides ants and the parasitoid
wasp, a number of other arthropod species were frequently
123
observed on the shoots of Leea manillensis. Of these, the
most common visitors were weevils (Curculionidae), peacock flies (Tephritidae), a variety of Orthopterans, planthoppers (Fulgoroidea; Auchenorrhyncha), leaf beetles
(Chrysomelidae), and spiders (Aranaeae). Peacock flies
were only feeding at the EFN, spiders were also seen
hunting. The rest, besides feeding on the EFN, was mainly
observed damaging the young leaves. Ants used to tolerate
most of these species. In particular, weevils were tolerated
even at the extrafloral nectaries. Some photographs of
further arthropod species on Leea manillensis are shown in
Online Resource 4.
Ant association and herbivory
Two different kinds of shoot damage became apparent
during the study. On one hand, there were single feeding
incidents by a range of herbivores, sometimes lasting over
several days. This type of damage was mainly caused by
Orthopterans and leaf beetles with whole parts of the leaves
missing, or by weevils with leaves perforated. However,
these feeding events rarely caused damage exceeding 20 %
leaf area loss. In contrast, in 25 % of the shoots (54 of 213),
the loss of the whole shoot was observed caused by one or
more unidentified herbivores. In many cases, we realized
this to have occurred over night. Apparently, this kind of
damage had the most important impact on Leea manillensis.
Ant-free shoots suffered a significantly greater loss of
leaf area than ant-associated shoots (Fig. 1). The total
shoot loss, recorded as 100 % of leaf area lost, played a
significant role. 57 % of the plants not associated with ants
suffered from a total shoot loss, compared to only 8 % of
ant-associated plants. A re-analysis without the samples of
Fig. 1 Loss of per cent leaf area in ant-associated (n = 89) versus
ant-free shoots (=91) of Leea manillensis. Differences are significant
Mann–Whitney U test: p \ 0.001. 100 % leaf area loss corresponds
to a total shoot loss
complete shoot loss revealed no differences in general
damage in both groups (Fig. 2).
The parasitoid wasp of the genus Chalcura:
observations and experimental results
Oviposition
Female wasps of the genus Chalcura were frequently
observed to place their eggs into the stipule of the plants
shoots close to the stipules’ extrafloral nectaries. These
large stipular structures enclose and protect the next generation of leaves in an early developmental stage. Whenever the arrival of a female wasp was observed, the wasps
never directly landed on the stipule, but on a leaf close to it.
After a period of a few minutes, meanwhile probably
observing the shoot, the wasps directly walked or flew to
the stipule.
During these observed ovipositions, which lasted about
1 h up to maximum of 2 h, the female wasp placed masses
of eggs directly into the closed stipule without changing the
position. During the 3 months of monitoring Leea manillensis, a total of 136 females were observed to oviposit into
the stipules of 66 plants, indicating single plants to be
utilized repeatedly and often simultaneously by two, three,
or even four females (Online Resource 2). On 25 out of 66
plants over a period of a few days up to 2 weeks, wasps
repeatedly were observed to oviposit into the plants stipule.
Experimentally opening those stipules, we found the
undeveloped young leaves to be embedded in masses of
eggs (Online Resource 2).
Out of 136 observations of oviposition, 128 took place
on ant-free shoots. Only eight wasps were observed on ant-
Fig. 2 Loss of per cent leaf area in ant-associated versus ant-free
shoots. Data from Fig. 1 re-analysed, but cases with complete shoot
loss excluded. Then, no longer resulting in difference in leaf damage
(Mann–Whitney U test: p = 0484.)
occupied shoots. Four of them were either not noticed or
ignored, one wasp was killed and three were chased away
by assembling ants. Occasionally, female wasps were
observed to feed at the extrafloral nectaries.
Infestation with planidia and loss of leaf area
Out of 213 experimental shoots, a total of 53 % were
infested with planidia (n = 114). The parasitoid wasp had
a massive effect on Leea manillensis, its association with
ants, its shoot development and survival. Only 19 % of
shoots infected with planidia were visited by ants, compared to 68 % of uninfected shoots (Fig. 4). Shoots infested by planidia had a significantly higher loss of leaf area
than planidia-free shoots. Again, the total loss of the shoot
due to herbivory was responsible for most of the differences between the two groups. In planidia-infested shoots,
the incidence of a complete shoot loss was almost 10 times
higher than in those not infested, with 38 % compared to
only 4 % (Figs. 3 and 4).
Observations on the wasp biology and its influence
on ants
Shoot development was strongly affected on planidia-infested plants. In 38 % of the observed plants, the planidia
hatched synchronically with the shoot expanding, with no
visible harm to the young leaves. In 62 % of infested
shoots, the mass eclosion of planidia apparently affected
the stipules’ tissue. In these cases and probably depending
on shoot size, its stage of development and the number of
planidia hatching inside the shoots suffered from different
kinds of shoot damage (Online Resource 3).
Fig. 3 Loss of per cent leaf area in Leea manillensis. Comparison
between shoots infested with planidia (n = 114) and not infested
shoots (n = 99). Mann–Whitney U test: p \ 0.01. 100 % leaf area
loss corresponds to a complete shoot loss
123
Discussion
Ants, EFN, and herbivory
Fig. 4 Comparison of the two samples ‘planidia-infested’ (n = 114)
and ‘not infested’ shoots (n = 99) of Leea manillensis in terms of
a their average leaf area loss caused by herbivory, b their percentage
of complete shoot loss caused by herbivory, and c their percentage of
shoots with ant association
Emerging planidia had a body length of about 0.13 mm
and were found sitting in a typical lurking erected position
on the expanding shoot, most often in high abundances,
concentrating on the area around the stipule and its nectaries (see Online Resource 2). In this early stage, often
dead ants were found hanging on strands attached to the
shoot. Those ants always had planidia on them. This
observation was made for most of the common ant visitors.
In the following days of observation, the number of
planidia on the shoot declined, either due to them getting
carried away by foraging ants or other visitors, or by
actively spreading over the whole plant by moving along
strands. The infestation with planidia caused a complete
loss of ant association after a short time. Ants then actively
avoided the shoot.
Planidia taken to the field station in vials survived up to
3 weeks without food intake. Ants placed into the vials
immediately showed an intensified, but apparently inefficient cleaning behaviour. After a short time, first locomotive dysfunctions were observed, followed by
immobility connected with a typical twitch of legs and
antennae. In the final state, ants did not show any vital
signs at all. With fresh planidia and small ant species of
the genera Crematogaster, Technomyrmex, or Dolichoderus, this state was reached in 10–30 min. In big ant
species like Polyrhachis, it took up to 1 h. The planidia
were preferentially situated at the soft parts of the ants
between the segments, probably feeding and apparently
gaining body mass. Apart from that, there were always
some larvae showing the typical lurking position on the
ant’s bodies. Photographs of the ant-parasitoid wasp and
effects on Leea manillensis and associated ants are given
in Online Resources 2 and 3.
123
Our results show that Leea manillensis consists of a highly
variable system, shaped by a number of factors. We could
confirm that it has mutualistic interactions with ants
attracted by extrafloral nectaries and food bodies. Ants
provided a certain degree of protection. However, the
indirect defence mechanism via ants turned out to be very
fragile, dependent on different conditions that influenced
the magnitude of the beneficial effect of ants on plants (see
also Rico-Gray et al. 2012). We found no evidence of any
specific relationship between Leea and the attracted ants.
Meng et al. 2011 describe similar conditions in the species
Leea glabra (Leeaceae) growing in South China. The broad
range of ant species observed to visit the extrafloral nectaries is consistent with a pattern typically found in many
facultative ant–plant associations and indicating a low
specificity (Fiala et al. 1994; Kessler and Heil 2011). Not
only did we find a broad range of ant species, but also a
high intraspecific variability in the abundance of ants
assembling at the extrafloral nectaries. Only 36 % of the
observed shoots showed a permanent presence of ants
perhaps due to varying EFN production of the plants. In
general, variability is quite common in facultative ant–
plant systems (Horvitz and Schemske 1984), and different
abiotic and biotic factors may have massive effects on
species interactions (Thompson 1999), directly influencing
the outcome of ant–plant–herbivore mutual effects and
determining the benefit for the plant (e.g. Ness et al. 2006;
Rosumek et al. 2009; Pires and Del-Claro 2014).
Nonetheless, myrmecophily in plants evidentially is an
adaptive mechanism of defence (e.g. Heil and McKey
2003). In this study, we are able to show that ant attraction
by the production of EFN on shoots significantly increased
the chances of shoot survival of Leea manillensis. Certainly, herbivory pressure is an important parameter for
Leea and a critical selective factor as reported for many
plant species (e.g. Vasconcelos 1991; Karban and Agrawal
2002). While the average herbivore damage on the shoots,
with \20 % leaf area loss, was probably tolerable for the
plant, the phenomenon of complete shoot loss played a
considerable role: 57 % of the shoots without notable ant
occupation suffered a total shoot loss, in contrast to only
8 % in ant-occupied shoots. In Leea manillensis, the large
stipule offers the biggest nectaries and is the part of the
shoot with the highest rate of ant visitation. The optimal
defence theory (McKey 1974) predicts that a plant should
optimize its defensive investment according to the organs’
current and future value and its likelihood of attack (reviewed in Heil 2008). In Leea, the defensive investment
and its focus on the shoots’ stipule corresponds with its
high risk of attack. Beyond that, the successful attraction of
ants to the stipules offered protection to an even more
important enemy, not only directly harming the shoots, but
also indirectly by affecting the defensive function of the
ants for Leea manillensis.
Influence of the parasitoid wasp
The ant-parasitoid wasp focused on the stipules of Leea
manillensis for egg deposition. A broad range of plant hosts
and different types of plant tissue have been reported as a
depot for eggs in other Eucharitid ant-parasitoids: seeds,
flower buds, flower stems or young fruits, leaves or young
shoots (Heraty 1985, 1994; Varone and Briano 2009; Carey
et al. 2012). In Leea manillensis, the stipule that encloses
the next generation of shoot is the spatial focus of investment into defensive mechanisms via EFN and FB. Obviously not general herbivores are the most damaging threats
for Leea manillensis in the study area, but the specialized
ant-parasitoid.
The hatching planidia had a massive impact on the
whole ant–plant–herbivore system. In our study, about
53 % of the plants’ shoots were observed to be infested
with planidia. All of the common ant visitors at the
extrafloral nectaries strictly avoided the shoots after the
eclosion of the wasps’ larvae. That might explain why we
noticed planidia to leave the shoot and disperse. The
avoidance of shoots and the observed aggressive behaviour of ants towards adult wasps indicate that the ants
were able to recognize this parasitoid as a threat. On
81 % of the planidia-infested shoots, ants were never
observed to visit the extrafloral nectaries. Our observations suggest that this avoidance finally led to a cessation
of EFN secretion in most of the shoots. Hence, the
avoidance of planidia directly resulted in a lasting lack of
ants on the shoots, which then caused an almost 10-fold
increase in complete shoot losses compared to the unaffected shoots.
On the other hand, the presence of ants had a great effect
on the behaviour of the parasitoid wasp. Low presence of
ants enhanced the risk of egg deposition on the plant, often
leading to a vicious circle hard to overcome. Successful
ovipositions into the stipule in the presence of ants were
very rare. Half of all observed attempts in the presence of
ants were interrupted due to very aggressive ant behaviour.
In the Eucharitidae, a high host specificity is assumed as
well as a certain amount of co-evolution with a particular
ant host subfamily (Heraty 2002). We found no evidence
that the parasitoid wasp on Leea manillensis had its focus
on a specific ant species, genus, or subfamily. In the contrary, we observed a broad range of ant species to get
attacked and killed by the planidia, but have no information
about the effect of the parasitoid on the ant colony level. As
far as it is known, hatching planidia are carried into the
colony via foraging ants (Pérez-Lachaud et al. 2006;
Lachaud and Perez-Lachaud 2012). On Leea, the number
of larvae on the shoots declined over time, and we
observed planidia to spread over the whole plant along
strands. We assume but do not know whether they were
carried away by ant foragers. So far, the attack of adult ants
has not been reported for the family Eucharitidae. In our
experiments with planidia and different ant species, all of
the ants died quite fast. On Leea, we found foraging ants
killed and attached to leaves and stems. Ants tend to keep
foraging trails free of dead nestmates to avoid disease
transmission (e.g. Wilson et al. 1958). Besides lurking on
the shoot and other plant parts, lurking on corpses might be
another way to ensure a contact with foraging ants. Access
to ant colonies might also occur via alternate hosts (Heraty
et al. 2004). As we observed the planidia to attack all kinds
of insects in the vial experiments, they might indeed use a
number of different taxa of insect prey as a transport
vehicle. However, the killing of potential host ants foraging
on the plant does not fit to what is known so far about
phoretic behaviour of the Eucharitidae. At this point, we
could only speculate what strategies planidia might use to
gain access to the ant host colony.
A growing number of studies provide support for topdown influence on ant community, coexistence, and
structure in the form of behavioural responses to parasitoids, although they were yet only reported from dipteran
parasitoids (e.g. Feener 1981; Morrison 2000; LeBrun
2005, Feener 2000). Parasitoids not only pose the direct
threat of mortality but also provoke a behavioural response
of ants, consisting of an alteration of their foraging behaviour. This interaction modification (Wootton 1994) not
only affects the defensive function of the ants for the
plants, but may also change interactions among competing
ant species, in which the presence of the parasitoid indirectly affects the abundance of species. Further investigations should focus on the effects of this parasitoid on
different ant species visiting the plant. The unexpectedly
strong influence of this parasitoid on the ants and plant in
our study shows for a first time that top-down processes
might be very important for different trophic levels in
facultative ant–plant mutualisms.
Acknowledgments The study was generously supported by the
DAAD (German Academic Exchange Service). We are very grateful
to the Philippine NGO PhilinCon and the staff of the research station
Sibaliw for their support, especially Benjamin S. Tacud jr. and Junmar E. Jamangal. We thank the PASU Rhodel B. Lababit (DENR) for
the generous permission to work at the NW Panay Natural Park. We
also thank Stefan Schmidt (ZMS) for the determination of the parasitoid wasp.
123
Compliance with ethical standards
Conflict of interest
of interest.
The authors declare that they have no conflict
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123
Appendix 5
Flight Styles in some Passerines
By Eberhard Curio1
Abstract. – Semi-quantitative observations of the circumstances account under which
bounding flight and steady (flapping) flight are performed by two forest-dwelling passerines
of two different lineages (Pycnonotidae, Philippine Bulbul Hypsipetes philippinus; Paridae,
Great Tit Parus major) in free flight. Bounding flight is one of two intermittent flight styles,
to the exclusion of undulating flight, in passerines weighing ≤100g. A new bounding flightspeed relationship is apt to shed light on the power output of bounding flight. In line the
function found in most flying animals for metabolic energy expenditure against speed the
relationship support a flattened J-curve as compared to U-shaped curve found in some birds;
this is based on the monotony of the parameters of bounding as a function of speed. Whether
bounding flight is saving energy as proposed by the work of Rayner et al., and of Tobalske in
tunnel wind experiments, would depend to the kinematics of flight in free-flying birds such as
the Bulbul and the Tit. – Further observations demonstrate that both steady flight and
bounding flight are used in foraging and in chases of two birds of the same species, whilst
‘mock’ prey capture and ‘group acrobatics’ in the Bulbul are tentatively assigned a training
role serving prey capture and predator evasion, respectively. – Observations of Tree Sparrows
(Passer montanus) support the idea of flapping flight changing abruptly into the still phase of
bounding flight and is therefore consistent with ’body morphing’ in the Bulbul.
Keywords: Flight style, passerines, bounding flight, functional significance, Philippines,
training flight.
Eberhard Curio, Conservation Biology Unit, Ruhr University Bochum, D-44780 Bochum,
Germany; Tel. No. +49-(0)234-3222858. Email: [email protected]
Introduction
Of two intermittent flight modes, the undulating flight and the bounding flight (Welty &
Baptista 1988, Moore & Birkhead 1991), only the latter is exercised by the three passerines
closely observed in the field. Their body masses fall well within the range ≤100g for the two
flight modes to be expected on theoretical grounds (Rayner 1985, Rayner et al. 2001,
Tobalske 1995, 2001).
Speed of flight as an all-important determinant associated the number of flight parameters.
For two of the three species, namely the Philippine Bulbul (Hypsipetes philippinus) (B.) and
the Great Tit (Parus major) (T.), correlations of speed with other flight parameters are
observed that can shed light to the energetics of flight. Based on their various models of
bounding flight in the wind tunnel (Tobalske 1995, 2001) and mathematical models (Rayner
1985, Rayner et al. 2001) the semi-quantitative observations of free-flying passerines are apt
to contribute to the functional explanations of flight.
They can account for details of bounding, e. g. depth and width of the ‘valleys’, they lack the
a quantitative assessment of the number of wing beats per flapping cycle but they can provide
estimate of in relative terms of ‘bigger’ and ‘smaller’. This is based of energy saving although
1
This paper in publication No. 93 of the ‘Panay Eco-Social Conservation Project’
(www.panaycon.org), the successor of PESCP.
the interpretation of the models is marred by conceptual pitfalls enumerated the end of the
Discussion.
In a third species, the Tree Sparrow (Passer montanus), observations of birds engaged in
climbing flight shed to light on the phenomenon of ‘body morphing’ (Curio & Tacud 2009).
Method
The observational methods employed were ad libitum sampling of flight events (Martin &
Bateson 1990). A flight event was defined as locomotion with the help of wings while hops
bridging two short distances in the canopy were not. – Observations were recorded
immediately they are seen, preferentially by drawing sketches with explanatory notes. A
number those sketches had found in their way into the present paper.
Flight Styles
The Speed-Bounding Relationship. Bulbuls (B.) and Great Tits (T.) perform bounding flight
or flap-bounding when traversing open terrain. Only rarely did they excise steady flight with
continuous wing flapping. Only above a distance of ca 2m do who engage in bounding (Fig.
1, lower panel). Based on crude measurements, the following relationship is new: as the
distance covered increases as the bird is lengthening bounding and simultaneously the
amplitude of the bounds decreases. At the highest speeds the amplitude becomes so shallow
that is hardly noticeable (Fig. 1, upper panel). In the B., the flight at high speeds occurred
above the canopy, seldom through it. There are were mostly two birds flying following
linearly each other, over long distances from the appearance in the canopy to their vanishing
out of sight. Similarly the T. were observed to fly at the highest speed when commuting
between an isolated nest tree and the nearest cover (Fig. 1, upper panel). The line of flight
covered the shortest distance between the forest and the nest, or similarly when crossing a
street.
The number of wing beats per flapping cycle decreases at the highest speeds while the
average wing beat frequency increases. This suggestion is consonant the Tobalske’s (1995)
observation of free-flying European Starlings (Sturnus vulgaris) in the wind tunnel.
Horizontal flight involves bounding to a large extent. Short hops in all directions characterize
movements in the canopy. When these moves go over longer distances they encompass
bounding in upwards flights not exceeding a certain angle (see Fig. 2 below). Downward
flights never employ bounding; they rather employ steady flights, except for the last stretch
ending in a stall (‘Unterfliegungslandung’ Lorenz 1933). This upward movement is
functioning as a brake at the end of a downward sweep.
Moving within and outside of the Canopy. – As said, flights between trees and within
closed canopy requiring the bird to reach out beyond (short) hops employ bounding.
Bounding is performed at all angles below bounded by a climbing angle in steady flight
involving climbs (Fig. 2). The border line between bounding flight upwards and climbing can
be approximately inferred from the angles subtended from the vertical (Fig. 2); the bounding
angle is measured by taking the line connecting the peaks of all ‘valleys’ against the vertical.
The angle of climbing is measured similarly by translating the distances of the sketches into
real-world measurements. In a crude approximation the bounding angle and the climbing
angle when subtended from the vertical measured about 45° and 5°, respectively. If bounding
is believed to save energy (Rayner 1985) this style is superior to steady flight involved in
climbing. - At rare occasions climbing flights occur at larger angles in the distance is
considerable and the bird is traversing open ground as compared to the relatively short climbs
in the canopy. This is probably driven by fear from accipitrine hawks. And tits had been
shown at bound at steep angles close to 5° (Moore & Birkhead 1991). Hence, the two flight
styles may overlap in bounding/climbing angle.
Sometimes a bird is performing a steady flight over longer distances. This occurs in the
pursuit of slow flying insects, or, two birds are following each other (Fig. 3). Why is the
steady flight in chases replacing bounding? The flights suggest that they are birds use chasing
when at home. Their behaviour is superficially different from the extended chases over long
distances almost always above the canopy under over open terrain. A pair of T. the behaved
of a to the birds expecting predation (Fig. 1).
Bulbul-specific Flight-styles. – Unique chases of 3-6 individuals in a group start invariably
high up in the canopy and end low above the ground, or, when horizontally oriented, in
another part of the canopy (Fig. 4 a). Chases start and end abruptly, going over about 15-25
m. The flight-style is steady as sketched in one of several birds of a group (Fig. 4 a). There is
no sign of hostility, or any calling. The group acrobatics rise at fruiting trees that are
harvested. – The one other flight-style the driven by foraging (Fig. 4 b). The B. performs,
always in steady flight is that allow maneuverability, loops in flycatcher-style to return back
to the vantage point. It is always clear whether an insect has been captured on the wing. The
hawking-style of flight is certain in the T. as well and when conspicuously foraging on moths.
– A problematic flight-style is seen during ‘mock’ prey capture on the wing (Fig. 4 c). In
steady flight is following a each twist of a ‘perceived’ insect on the wing. The twisted flight
and the steady flight-style suggest the bird is following a real or virtual insect, but other
interpretations cannot be ruled out (see Discussion).
Like many passerines B. and T. can hover. The cherry-like fruit of Ficus benjamina is largely
plucked while hovering.
Tree Sparrows adjust the Duration of Climbing. – The body-morphing hypothesis hold the
bounds of the bounding flight-style can be give way the still phase with wings folded
(sleeked) against the body at a point of time. This is important for passing to narrow openings
in dense foliage (Curio & Tacud 2009). Consonant with this timing is the another flight-style
in climbing to used to overcome an obstacle. Tree Sparrows (Passer montanus) adjust the
duration of a climb in steady flight to pass over a vertical fence regardless of the start point of
the climb, or, regardless of the climb angle (Fig. 5). This performance shared the bodymorphing the abrupt change from flapping flight to a still phase during a descent to the
ground.
Discussion
The Mechanism and Function of intermittent Flight. – Of the two types of intermittent
flight styles, the undulating flight and the bounding flight, the B. and the T. perform only the
latter. The eases the discussion that will consider the bounding flight (= flap bounding) and
the steady flight with continuous flapping.
The thrust of the present paper is the demonstration of the monotonous, multi-dimensional
relationship between bounding flight and speed (Fig. 1). The European Starling (Sturnus
vulgaris) and the Budgerigar (Melopsittacus undulatus) perform two flight-styles, i. e.
undulating flight and bounding flight. Authors (Rayner et al. 2001, Tobalske 2001, 1995) had
unanimously asserted the flap-bounding replaces flap-gliding only at high speed. The
relationship of bounding and speed in two unrelated passerines, in the B. and the T., has leads
to serious repercussions on the power output as the function of flight speed.
Based on a mathematical model Rayner (1985) proposed the birds, with body masses of
≤100g (Tobalske 2001), bounding save energy only at high speeds. This would in line with
the observation of the replacement of undulating by bounding mentioned above. What that
happens a lower speeds? One it few consonant that bounding would saves energy by lifting
the body and tail to extent to it reduces the costs of transport in the bird regardless of speed
(Tobalske 2001), would not settle the issue. In tunnel wind experiments the flap frequency
increased with speed. At the same time the energy output changed in a U-shaped curve as
speed increases both in the European Starling and the Budgerigar (Rayner 1985, Tobalske
2001). The crucial observation come the present findings that the monotony of the speedbounding relationship. The monotonous change of the width and depth of the valleys, and the
wing beats of the flapping cycle (Fig. 1), as a function of speed, argue against a U-curve of
the energy output with flight parameters. This leaves the flattened J-curve as the logical
alternative.
The basic tenet of Rayner (1985) that intermittent flight-styles have been evolved as energysaving mechanisms not either refuted not supported by the results. Support has from
electromyographic experiments by Tobalske (1995). He found in the Starling that in the nonflapping phase certain muscles (M. pectoralis, M. supracoracoideus) are resting. In this way,
muscles would be optimally functioning in that energy could be saved. This rests on the
assumption that during the flapping phase the saving not offset by an increase of power output
thereafter. Thus, the support is not equivocal.
Other Flight-styles of Bulbuls. – Three other flight-styles demand a high level of control and
employ therefore continuous flapping (Fig. 4 a). One style is unique that is involves a highly
coordinated maneuver of at least 3-6 individuals chasing each other and ever colliding in the
chase. The chases downward kept close to a mature tree, or, beneath the canopy when
horizontally oriented. (Quite often two birds were closely chasing in each other through the
canopy, but then calling characteristically.) – Rarely a single bird was seen following a
twisted path if with to capture an insect of the wing (Fig. 4 c). The flight-style was steady
flapping. It cannot be excluded that chases were ending in success. However, firstly, the path
followed ended that it begun, in steady flight; secondly, there was not visible swallowing of
an insect. Therefore I classify the group acrobatics and the ‘mock’ prey capture tentatively as
training to avoid aerial predators and exercising foraging, respectively.
Conceptual Difficulties. – The bird flight is a multi-purpose method of locomotion. A
functional interpretation is therefore fraught with conceptual difficulties. That bird flight is
adaptive and, hence, can be investigated by methods that appear stringent as ideas underlying
causation of traits (Curio 1973; Endler 1986). The levels of explanation, the mechanistic
(proximate) and the functional (ultimate), are sharply is to be discriminated.
Bounding flight was functionally explained by Rayner (1985) he suggested to saving of
energy at very high speeds (functional explanation), leaving open the question happens as
lower speeds. In the same paper this is concluded that bounding to due to ‘conflicting
constraints imposed by physiology and mechanics of flying birds’ (proximate explanation).
The naturalistic observation about bounding as presented here support the model in regard to
a functional explanation.
In a second paper Rayner et al. (2001) stuck to the energy saving hypothesis, in couched in
new terms, but mixed it with terms of proximate explanations. Based on different arguments
the authors concede the bounding to be have evolved functionally in response the ‘flight
performance optimization’, for instance against pressure of predation; this is a sub-goal of
optimization. To fully understand bounding one needs to study it functionally and
proximately. The study of function has taught us to look for more than just one function: a
behaviour may serve multiple goals (Curio 1973). Our understanding of the mechanics of
intermittent flight has been tremendously promoted through models and experiments of
Rayner et al. (2001) and Tobalske (1995, 2001) but has it lacks direct predictions springing
from alternative hypotheses. Progress will be made by 1) not mixing conceptually proximate
and ultimate explanations; or, 2) formulating hypotheses thought to be exclusively designed
while a hypothesis while in fact it
embodies of the another; or, 3) by looking for support of
a hypothesis is one species by it testing with a different species.
‘Body-morphing’ further supported. – The flight maneuver consists of the precise timing
of folding the wings when passing through a bird net with impunity (Curio & Tacud 2009).
‘Body-morphing’ rests on the assumption that the flapping phase the interrupted at any
moment to give way to the still phase. A corroborating observation is found in the climbs of
Tree Sparrows over a fence (Fig. 5): regardless where the climbing with steady flight started
and reached the summit of the ascent, can they stop any moment before descending of the
other side of the fence. This observation falsifies the idea that flapping can be performed only
fixed time intervals.
Zusammenfassung. – Sperlingsvögel (Passeriformes) vollführen einen Bogenflug, einen von
zwei Flugstilen, bei dem aktive Flatterphasen mit passiven Phasen bei angelegten Flügeln
einander zu ablösen, und einen stetigen Flug (steady or flapping flight). Wie die von Theorie
voraus gesagt, eignet der Bogenflug Vögeln von weniger als 100 g Körpergewicht. Semiquantitative Beobachtungen lassen die Umstände beschreiben, unter denen bei zwei frei
lebenden Arten
(Pyconotidae, Philippinen-Bülbül Hypsitetes philippinus; Paridae,
Kohlmeise Parus major) der eine oder der andere Flugstil auftritt. Eine neue Beziehung
zwischen Eigenschaften des Bogenflugs, der Fluggeschwindigkeit, und zurücklegenden
Distanz wird graphisch geschrieben und lässt Aussagen über den Kraftaufwand des
Bogenflugs zu. Entsprechend den Befunden bei den meisten fliegenden Tierarten gehorcht
der metabolische Energieaufwand als Funktion der Fluggeschwindigkeit einer flachen JKurve im Vergleich zu einer U-Kurve wie bei den anderen Vögeln; dies erhellt aus der
Monotonie des Bogenflug-Geschwindigkeits-Relation. Ob der Bogenflug Energie spart, wie
Rayner et al. aufgrund mathematischer Modelle voraus sagen, und solchen von Tobalske in
Windtunnel-Versuchen, würde von der Kinematik des Fluges bei freifliegenden Vögeln wie
dem Bülbül und der Kohlmeise abhängen. Begriffliche Schwierigkeiten der Erklärung
aufgrund funktionaler oder kausaler Betrachtung besonders von Modellberechnungen oder
Versuchen werden aufgezeigt. – Weitere Beobachtungen zeigen sowohl Bogenflug oder auch
stetiger Flug bei der Nahrungssuche, auch bei Jagden zweier Vögel derselben Art (Bülbül)
wie auch der ‚Scheinjagd‘ nach Fluginsekten sowie bei ‚Gruppen-Flugspielen‘ des Bülbüls
werden geschrieben. ‚Scheinjagden‘ und ‚Gruppen-Flugspielen‘ werden hypothetisch als
Flüge zu zum Training des Insektenfanges bzw. des Feindschutzes funktional gedeutet. –
Beobachtungen von Feldsperlingen (Passer montanus) stützen die Hypothese, dass der
stetiger Flug unmittelbar in die passive Phase des Bogenflugs übergehen kann und damit das
Modell des ‚body-morphing‘ beim Durchfliegen enger Hindernisse stützt.
Acknowledgements
This study of the PanayCon was conducted under the aegis of a Memorandum of Agreement
between DENR (Dept. Environment and Natural Resources, Quezon City) and PhilinCon and
the support by PAWB/DENR Director Dr. M. Lim, DENR Reg. VI Regional Executive
Director Julian Amador, his successor RED Jim O. Sampulna, and DENR Park
Superintendent Rhodel Lababit is gratefully acknowledged and is as the collaboration the
University of the Philippines, President Hon. Alfredo E. Pascual and Vice-Chancellor of
Administration Prof. Maragtas Amante. Vital support came from the Frankfurt Zoological
Society (CEO Dr. C. Schenck, Asia Superintendent Dr. A. Müllner), the Bird Protection
Committee (President Prof. Dr. E. Schneider), the Bird Research and Conservation
Foundation (President E. C.), the Erwin-Warth-Stiftung (President Hilde Stühlinger), Rettet
den Regenwald [Save the Rainforest] (R. Behrend), A. M. de Dios, Oskar Lopez and other
donors. Further assistance was rendered by PanayCon staff Dr. E. Sanchez Jr. (President
PhilinCon), and most circumspectly by J. Tacud, E. Geronimo and J. Jamangal at the
Research Station ‘Sibaliw’.
References
Curio E (1973) Towards a methodology of teleology. Experientia 29: 1045-1058
Curio E & Tacud J (2009) ‘Body morphing’ enables forest dwelling birds to master dense
foliage with impunity: a novel avian flight performance. (Ökol Vögel) Ecol Birds 31: 1-12
Endler J A (1996) Natural selection in the wild. Princeton Univ Press, Princeton, New Jersey
NJ
Martin P & Bateson P (1990) Measuring behaviour. Cambridge Univ Press, Cambridge, New
York, Port Chester, Melbourne, Sydney. 4th ed.
Moore M & Birkhead T (1991) The Cambridge Encyclopedia of Ornithology. Univ Press
Cambridge, Cambridge
Lorenz K (1933) Beobachtetes über das Fliegen der Vögel und die Beziehungen der Flügelund Steuerform zur Art des Fluges. J Ornithol 81: 107-236
Rayner J M V (1985) Bounding and undulating flight in birds. J theor Biol 117: 47-7
Rayner J M V, Viscardi P W, Ward S & Speakman J R (2001) Aerodynamics and energetics
of intermittent flight in birds. Amer Zool 41: 188-204
Tobalske B W (1995) Neuromuscular control and kinematics of intermittent flight in the
European Starling (Sturnus vulgaris). J Experimental Biology 198: 1259-2273
Tobalske B W (2001) Morphology, velocity, and intermittent flight in birds. Amer Zool 41:
177-187
Welty J C & Baptista L (1988) The Life of Birds. Saunders, New York. 4th ed.
Legends
Fig. 1. Bounding or bound-gliding related to amplitude and distance flown in the Philippine
Bulbul (B.) and the Great Tit (T.). Flight speed is related to distance and increases from
bottom to top. – A pair of T. commuting over 30m between the nest box and the nearest
forest. – Long distance flight in the B. over non-familiar home range.
Note: Distance flown with small deviations to the right or left superimposed on the flight
path. Dashes along the flight path symbolize schematically single wing flaps in this and the
following figures, with inter-dash intervals indicating speed, not necessarily wing beat
frequency (that in the European Starling wing flaps increase linearly with speed [Tobalske
2005]).
Fig. 2. Movements within and between portions within the familiar home range in the B.
Conditions for bounding vs steady flight (steep climb), and of almost any steady flight over
the longer distances, also within familiar home range.
Fig. 3. Continuous flapping flight at highest speed to capture an insect on the wing, or, two of
B. chasing each other, following the same flight path.
Fig. 4. Chases (‘acrobatics’) of 3-6 B. in steady flight, with path varying from horizontal vs
vertical, different observations (a). – Chasing insects on the wing (‘hawking’) (b). ‘Hawking’
with twists all a long the full length to the flight path (c).
Fig. 5. Climb by Tree Sparrow Passer montanus: ascent to overcome a fence from various
distances, then descending with bounding flight on the other side of the fence. The ascent
starts from any distance from the obstacle while the landing point (the same in the case) stays
the same.
Abb. 1
Abb. 2
Abb. 3
Abb. 4
Abb. 5

Annual Report - Panay Eco-Social Conservation Project

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