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Ecology & Evolution BIOL 2306 Field course:

Pak Tam Chung Monday 13th – Thursday 16th October 2025
NOTE the trip is 4 days this year!!

Arrival: Students must arrive at the Po Leung Kuk Pak Tam Chung Holiday Camp by 9:00am on Monday
13th October! There will be a bus leaving HKU campus (Library extension road) to Pak Tam Chung at
7:30am on Monday and another returning to HKU campus on Thursday afternoon. Alternatively, the
camp is five minutes’ walk from the Pak Tam Chung bus stop, which is served by KMB bus 94 from Sai
Kung.

Please assemble in the upper Main Hall A above the canteen to register in your groups. You will leave your
belongings in the Hall until the rooms are available for occupation in the afternoon. Please arrive dressed for
fieldwork (with footwear and clothes you can get wet in, sunscreen etc), as you will NOT have the opportunity
to change before going out for the Ant Day!

Accommodation and Food: Students are in the group hostels. Your house number and dormitory are
indicated on the class Moodle site. Staff and demonstrators are in the family hostels. Meals will be in the
canteen, at 8.00 am for breakfast, 18:00 for dinner, and lunches to suit activities during the week.

Clothing: The weather in October is unpredictable but can be hot or cool, so you must bring a range of
appropriate clothing. FIELD TRIPS ARE RARELY CANCELLED BECAUSE OF BAD WEATHER! You
will need strong, comfortable shoes for walking and old trainers, sports shoes or something similar that you
can wear in the water. Thongs, flip-flops, or sandals are NOT ACCEPTABLE since you will cut your feet on
the sharp stones. Everything you wear in the field will get wet and muddy. Please bring your own hats, water
bottles and remember to bring sunscreen!

Equipment: You will need pens and pencils, a clipboard, a notebook, and a small rucksack to carry things
in the field. A camera (make sure it’s in a waterproof bag) and binoculars would also be VERY useful. Forceps
will be ESSENTIAL for collecting and handling specimens in the field, so you should bring AT LEAST one
pair. A CALCULATOR is also ESSENTIAL for data analysis and AT LEAST ONE MEMBER OF EACH
GROUP SHOULD BRING A NOTEBOOK/LAPTOP COMPUTER WITH EXCEL AND POWERPOINT
FOR PREPARING THE PRESENTATIONS; AND A USB DRIVE TO TRANSFER FILES.

Books: A series of Field Guides have been produced, which include guides to Hillsides and Sandy Shores,
and are reasonably priced ($60) and available from most Commercial Press bookstores and were designed for
this field course! Any student seriously interested in coastal ecology should also bring “The Sea Shore
Ecology of Hong Kong” by Morton & Morton (Hong Kong University Press). Similarly, for terrestrial
ecology, interested students should buy/bring “The Ecology and Biodiversity of Hong Kong” by Dudgeon &
Corlett. The Agriculture, Fisheries and Conservation Department (AFCD) also produce a series of booklets
on local ecology which are useful (especially the books on butterflies!). Remember that we will be looking
at a wide variety of animals and plants and so general textbooks will be useful.

Course rules: A field course of this size is only possible if everyone obeys the rules. NOBODY is allowed
to leave the camp without permission from the course coordinator Prof. Russell. This will only be given in
exceptional circumstances. You must also obey the Pak Tam Chung camp rules: especially minimizing noise
after 11.00 p.m. and littering - please avoid both! University Halls of Residence rules also apply.

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Field, lab and lecture sessions: In general, field trips are held to meet the best environmental conditions
for the habitats to be visited. The Sandy shore trip will be held in the morning this year (to use the low tides).
Laboratory sessions are timetabled around the fieldtrips SO CHECK YOUR HANDOUTS! Lectures will
usually be in the morning/evenings. Students are divided into SIX groups (see attached list), each led by a
staff member. ALL ACTIVITIES ARE COMPULSORY AND MAY BE EXAMINED IN THE
ASSESSMENT TEST.

ASSESSMENT TEST: The field course is assessed by a test which will include
questions on the field trips, demonstrations, lectures and seminars at the camp.
THE TEST IS ON 11:00am Monday 20 October in and the class will be split across
MBG07 and MB103
Field course groups
Students and staff are divided into six (6) groups which will stay together throughout the course. Groups will
also involve undergraduate students from the University of Johannesburg (UJ, South Africa) as part of an on-
going student exchange programme. During the camp groups will also be allocated scores for tasks performed
to decide which group is the best on the field course.

Field groups; staff members and demonstrators:

Baeckeans: Dr B Hau; Chloe, Yuen Nok Lam; Andre Ibanez
Army Ants: Prof L Ashton; Kit, Lee Kit Wah; Taneisha Barrett
Prickly Urchins: Prof BD Russell; Rainbow, Leung Wing Sum; Emily Jones
Black Kites: Prof R Greenfield; Uva, Fung Yu Yan; Samantha Klein
Horseshoe Crabs: Prof GA Williams; Ivan, Lam Pui Ying; Carolin Dahms
Atlas Moths: Prof T Bonebrake; Matt Hamer; Elaine, Chan Tsz Ying

Technical staff: Ms Oscar So; Mr Pak Hei Chan; and Mr Boris Yu. Note all technical staff are on hand to
attend to any medical incident.

In the case of a medical emergency Mr Pak Hei Chan (Tel: 3917 3808, Room: 9B) or
Ms Oscar So (Tel: 3917 3309, Room: 9F) are on call 24hrs a day and will arrange
necessary treatment / transport to the nearest hospital.
Background ecological information: Before clearance by humans, the vegetation in the field course
area, like much of Hong Kong, would have been covered in forest. Remnants of the original forest flora still
survive in ravines and steep cliffs, protected from fire and cutting in the past, but much of the original terrestrial
flora and fauna must now be extinct. The grasslands and shrublands, which cover most hillsides in Hong
Kong, are maintained by fire and, if protected, there is a fairly rapid succession to forest. Grasslands generally
cover the hilltops and are much more common in the east of the Sai Kung peninsula than in the west where
forest vegetation is dominant. In the lowlands of the Sai Kung peninsula there are large areas of abandoned
paddy fields. Each village usually has a small “Fung Shui wood” although these are now mostly badly
disturbed. In the 1950s Chinese red pine was planted on many Sai Kung hill slopes, especially in the west.
Many have been attacked by nematode parasites but as they die out, native understorey trees grow up to replace
them. In the absence of well-established forest, bird species are fewer than in comparable areas of Hong Kong.
Most of the larger animals are nocturnal and rarely seen, but porcupines, pangolins, masked palm civets,
leopard cats and wild boars have all been reported from the Sai Kung peninsula.
Near the field camp (~10 mins walk along the Tai Mong Tsai Road) is the Sheung Yiu village where
a walled village with a lime-making kiln are part of a folk museum. For more information about the Pak Tam
Chung area and the Sai Kung Peninsula, see the excellent Friends of the Earth booklet ‘The Sai Kung
Peninsula’ and visit the Government website of Sai Kung East Country Park.

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Monday 13th October - Does temperature determine who comes to
dinner? A case study with ants

OBJECTIVES: investigate the influence of abiotic factors on foraging activities of ant

communities and its impacts on species composition.

Learning outcomes:
At the end of the field trip, you should be able to:
• understand good experimental design and propose new, improved methods to answer specific
ecological questions;
• compile data, conduct preliminary analyses and interpret results;
• discuss the ecological significance of temperature on species behaviour, the importance of
microhabitats in shaping and structuring communities; and organisms’ adaptations to their
environment.

Introduction
Understanding the distribution of species in space and time is a major goal of ecology as it relates to many
fields such as biogeography, conservation, community ecology, etc. Abiotic factors such as temperature and
precipitation play a major role in regulating species distribution. For instance, dry-adapted species of plants
and animals (e.g. camels, succulent plants) will be found in arid environments, while cold-adapted species
(e.g. penguins, polar bears, marmots, snow gums) will be found at high latitudes or elevation. Environmental
conditions will thus filter for species with specific morphological and physiological adaptations that allow
them to maximize their foraging and reproductive activities and hence their overall performance.
As a consequence, habitats with different climatic conditions will support different communities of
animals and plants. This pattern appears obvious to us while travelling in regions further south or north than
Hong Kong (or if you watch any film). Yet, this same pattern applies within a single ecosystem, but at a
smaller scale (or even at a micro-scale). For instance, the temperature measured on the soil surface of a forest
under the shade of trees will generally be relatively similar to the ambient air temperature, while any surface
directly exposed to the sun (e.g. the top branches in the canopy, or soil surface exposed to solar radiation) can
be up to 20-30˚C higher. As a result, organisms living just a few meters apart can experience very different
environmental conditions and will thus need different adaptations to cope with their respective environments.
Arthropods, in particular insects, represent the majority of organisms in terrestrial ecosystems. Most
arthropods are less than a few centimetres or even millimetres long and are, therefore, very sensitive to
localized changes in microhabitats and the associated abiotic conditions. Over evolutionary time, this has led
to important adaptations of species to colonize and exploit the diversity of microhabitats. The external
phenotype of these organisms, therefore, reflect their ecologies and can be studied using a comparative
approach to investigate morphological or behavioural adaptations. An obvious example will be the study of
eye size in arthropods living on trees (arboreal species) or below ground (subterranean species). Where there
is light, for example within the canopy, vision is an important attribute either to detect predators or prey. As a
result, arboreal species have larger eyes than subterranean species which live in a dark environment and whose
eyes are often reduced or absent.
Ants are the dominant group of terrestrial arthropods and encountered in most ecosystems, from the cool
tundra of Canada to the warmest African or Australian deserts. With over 16,000 species of ants described
globally, they exhibit a wide range of morphological and behavioural diversity. This diversity reflects their
ecological success and is the result of the adaptations that ants have evolved to exploit most niches and
microhabitats.
The size (and mass) and colour of ants vary greatly with species ranging from 1 mm to over 35 mm, and
light yellow to jet black colouration. The size of organisms is an important ecological component as it
determines the scale at which this organism is living, the quantity of food needed for growth and reproduction

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(the larger, the more food intake is necessary), but also how the environment might constrain its activity. As
organisms get bigger their surface area to volume ratio decreases, thus larger organisms expose proportionally
less tissue to the environment and, as a result, are less prone to desiccation and heat up more slowly.
Furthermore, most body parts and organs of larger ants are raised higher from the substrate (soil surface) than
small ants and thus experience lower temperatures. For instance, for a surface exposed to solar radiation, the
temperature measured 3 mm above ground can be as much as 4˚C lower than the temperature measured on
the soil surface. Colouration also plays an important role in thermoregulation; dark coloured organisms are
unreflective and will heat up faster, while lighter organisms will be more reflective and thus heat up at a slower
rate.
As social organisms, individual ant foragers are continuously scouting and collecting food resources.
In order to maximize their success in resource exploitation, ants have evolved various foraging strategies. Ants
(and other social insects) can recruit nestmates to exploit food resources and protect them against competitors.
Recruitment is a behaviour in which a successful scout (an ant that finds food) will lay a pheromone trail
(pheromones are molecular markers carrying a specific signal) while going back to the nest to recruit more
foraging ants. The pheromone trail will provide information to other nestmates about the quality of the food
resource and induce a behavioural response. Newly recruited nestmates will follow the trail to the food
resource and will, in turn, lay new pheromones while returning to the nest with the food. Because pheromones
are molecular markers, they evaporate with time from the surface on which they have been deposited; and
increased temperature will accelerate the evaporation rate.
In this study we will measure the difference in temperature between two microhabitats and the effects of
temperature on the species composition, morphology (size and colour) and foraging behaviour of an ant
community. Four hypotheses with their respective predictions will be tested.

1) Tree canopy will shade the ground surface and reduce temperature as compared to more open areas
directly exposed to solar radiation. As a consequence, we predict that baits placed in the shade will
experience cooler temperatures than baits exposed to direct solar radiation.
2) Due to surface area/volume ratio, larger organisms have increased tolerance for higher temperatures than
smaller ones. Thus, we predict that larger ants should be able to tolerate higher temperature than smaller
ants and, consequently, be more common in habitats exposed to solar radiation.
3) Dark-coloured organisms are expected to heat up more quickly than light coloured individuals. Thus, as
temperature increases, dark-coloured species should be more limited by temperature than light coloured
species in their foraging activities. Here, we predict a higher frequency of light-coloured species on baits
exposed to solar radiation than on baits in the shade.
4) Some ant species use pheromone trails to recruit nestmates to collect food resources, while other species
forage as individuals (no recruitment involved) and rely mostly on visual cues for orientation. As
temperature increases, the lifetime of the pheromone trails decreases, reducing the efficiency of this
mechanism as a foraging strategy. We hypothesise that with increased temperature, efficiency of
recruitment through pheromone trails will be reduced. Therefore, we predict that for high temperatures
(baits exposed to the sun), species using individual foraging will be less impacted than species which use
pheromone trails and, thus, will occur more frequently on baits exposed to the solar radiation than species
using pheromone trails.

Simply put, ants foraging in open habitats, with high soil surface temperatures due to direct solar radiation,
should be large, light-coloured species with individual foraging strategies; while nearby, shaded
environments should favour smaller, dark-coloured species using mass foraging strategies characterized with
high recruitment of nestmates through pheromone trails.

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Study sites
Each group will be assigned to a section of the grounds surrounding PTC for this exercise.

Field work:
1. Each group will be assigned a site location, and students will work individually, collecting data for two
baits, one in the sun (well exposed to solar radiation) and one in the shade. Ensure that baits are separated
from each other by at least 4 meters.
2. With the help of your demonstrators, each student will identify appropriate locations to set baits; one in a
sunny location and one in the shade. Ensure that each location will experience their treatments (i.e., be in
the sun or shade) for the entire sampling period (one hour).
3. Place the white disk (white plastic lid) on the ground and ensure that all the edges of the disk are in contact
with the ground (press it down if necessary). Place your bait (the sausage slice) in the centre of the
observation disk. This will allow you to quantify the ants both on and immediately around the bait (the
entire disk).
4. Record ground temperature at four points around each bait (one person per group is in charge to measure
temperature). Each point will be about 5 cm from the edge of the baiting station and temperatures should
be recorded at both the start and end of the experiment. This is a continuous variable which is expressed
in ˚C.
5. For each treatment, note the time at which you place the bait. Collect data for each bait every 10 minutes
for an hour (i.e., 0, 10, 20, 30, 40, 50, 60 minutes). NOTE: you will need to start the baits of the two
treatments approximately 5 minutes apart so that the data are offset.
At each time note:
a. the absence (0) or the number of ants in contact with the bait and on the white disk top (not
outside the disc). Ants can recruit a lot of nestmates in a short period of time, so use your phone
to take a picture to later count all the ants present (remember to note which time period and
treatment your picture corresponds to and that the picture is not blurry!). This is a discrete
variable.
b. whether the different species that you score are foraging as individuals or trail-following.
6. At the end of your observation period (after 1 hour),
a. Prepare a label indicating your name, group number and if your lid was in the shade or in the
sun and place the label within the Ziploc bag.
b. Then quickly pick up the lid with the bait and as many ants as possible on it and place it within
the Ziploc bag. NOTE: you need to use a separate Ziploc bag for each of your two baiting
stations!
7. Place all your labelled Ziploc bags in the cooler carried by one of the demonstrators.
8. Optional: Note or record any competitive interactions (aggression, avoidance) that you observe either at
the intra- or interspecific level (this could be useful for the presentation!).

In the laboratory (class):

1. As soon as you arrive back to the camp, place all your labelled Ziploc bags in a cooler with dry ice for
20 minutes.
2. For each baiting station, transfer the ants collected in the Ziploc bag into a vial with ethanol.
Remember to add a label in the vial with the same information as the one in the Ziploc bag. Leave
the ants in the vial for at least 10 minutes, then randomly collect 5-10 individual ants.
3. Measure the size of at least 5 individuals (but no more than 10) that you collected from each baiting
station. This is a continuous variable with units expressed in millimetres (mm).
4. Then determine the colour of individuals for each baiting station (see Figure 1). Dry the ants and then
determine which point of the scale best represents the colour of your specimens and score them as
either light colour or dark colour. This is a categorical variable.

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Figure 1. Colour scale for the determination of light vs dark colour for the ants collected.

Data presentation (AS A GROUP):

Prepare a 6 minute presentation (7 ppt. slides maximum) to present:
1) The mean temperature of the two treatments (sun and shade) and the proportion of the baits exploited by
the ants for both treatments;
2) The mean size of ants of the two different treatments;
3) The respective proportions of light and dark coloured ants for both treatments;
4) The respective proportions of solitary and trail-foraging ants for both treatments;
5) The mean abundance of ants on baits for both treatments over time.
6) Discuss your results in the light of your knowledge and the literature available to explain how ground-
dwelling ant communities are organized.
7) Think about other factors that might have interfered with / influenced the experiment and provide any
other information you feel is interesting or important.

Tip: Think about the scale relevant to the studied organisms, not the scale at which you experience the
environment – think like an ant!

PLEASE ensure you are ready for the presentations by 18:00!

Prof Louise Ashton & Prof Bayden Russell

(October 2025)

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DATA SHEET FOR FIELDWORK

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th
Tuesday 14 October - Sandy Shore at Starfish Bay
Low tide – 0.86 m @ 11:00

OBJECTIVES: to investigate the diversity and abundance of animals along the ecological
gradients of a sandy shore, and the adaptations important to life in sand.
Learning outcomes:
At the end of the field trip, you should be able to:
• use a vertical transect sampling technique to quantify the distribution pattern of sandy shore
organisms;
• draw graphs and present information to interpret these zonation patterns;
• interpret the morphological and behavioural adaptations of sandy shore organisms.

Introduction to sandy shores in Hong Kong

Sandy shores can be divided into two major habitat types depending on the strength of wave action the
shore receives. Open sandy shores often receive greater wave action resulting in the sand substratum being
composed of larger particles and being highly mobile and, hence, are also called mobile sandy shores.
Protected sandy shores can be found in sheltered locations where wave action is weaker allowing for the
deposition of finer sediment particles. The more stable substratum of protected sandy shores experience
relatively little disturbance and consequently have higher species diversity than open sandy shores.
A sandy shore is a three-dimensional habitat. Organisms can live both on the surface of the substratum
and also within the sand by burrowing. The fauna of sandy shores is conventionally grouped into three
categories:
i) Surface dwellers, which are collectively referred to as epifauna. Most epifauna are highly mobile
crustaceans (e.g. crabs) and the less mobile gastropods (e.g. mud snails). They are usually scavengers,
feeding on macroscopic detritus or particles sifted from the sand.
ii) Burrowers, including both shallow and deep burrowers, are collectively known as infauna. Most
infauna are bivalves and worms.
iii) Temporary burrowers, which may be surface dwellers or burrowers during different phases of the
tidal cycle. Surface dwellers are less abundant than infaunal species.

Life in sandy shores is distributed according to ecological gradients. On the rocky shore we investigated
vertical tidal gradients and wave exposure as two ecological gradients. As a sandy shore is three-dimensional,
ecological gradients, therefore, exist in three directions:

Vertical gradients are the most obvious on sandy shores and are caused by the tidal cycles. Sand in the
lower-intertidal zone only dries out at the surface during low tides. In the upper intertidal zone, sand may be
drier for longer periods and to a greater depth, as this area spends more time uncovered by the tides and
exposed to air. Exceptions include pools of seawater trapped in beach depressions although evaporation may
create hypersaline conditions. We therefore expect organisms inhabiting upper intertidal areas to be better
adapted to cope with water stress and desiccation.

Horizontal gradients are created by local topography and hydrography, such as the presence of a stream, or
variation in exposure to wave action.

Depth gradients within a sandy shore are created by oxygen availability and depend to a large extent on the
particle size of sand. Generally, oxygen availability decreases with increasing depth. This can be seen as a
blackening of sand due to the action of anaerobic bacteria (depositing sulphides, often giving a ‘bad eggs’
smell). Smaller sand grain sizes support lower oxygen content of pore water (the water between sand
particles). Since smaller sand grain sizes are associated with lower intertidal areas, we expect the black

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anaerobic layer to occur at a shallower depth than in upper intertidal regions where larger grain sizes
predominate. Dry sand is aerated and so does not experience an anaerobic layer.

The Study Site: Starfish Bay

The site at Starfish Bay is a protected sandy shore. The shore, being flat and well sheltered from the
impact of strong oceanic waves, has a wide intertidal zone. At the top of the shore is a steeply eroded bank
fringed by grasses, creepers and shrubs. On the seaward side of the bank, in the coarse sand, can be seen the
burrows (~ 5cm diameter) of the ghost crabs (Ocypode sp.). Lower down the shore, the surface is dominated
by the mud snails including Batillaria, Pirenella and Cerithidea spp. Fiddler crabs, sand-bubbler crabs
(Scopimera intermedia) and soldier crabs (Mictyris brevidactylus) can also be seen moving swiftly over the
sand surface during quiet, undisturbed conditions. If you are quiet, you may even see the impressive sight of
soldier crabs moving swiftly across the beach in troops. Evidence of infaunal species can be seen from the
many siphon holes and sand-casts on the surface. If you look carefully at the wet sand near the water line,
you may also see the feeding proboscis of infaunal animals sweeping the surface in a spiral motion. A common
burrowing animal in this part of the shore is the innkeeper worm (Ochestostoma erythrogrammon), which has
a red-coloured body and a translucent, extendable proboscis. Try digging through the sand surface, you will
see that whereas the sand is yellow and dry in the upper beach, it is wetter and becomes black just below the
surface in the lower beach area.
You will find that during low tides, most animals on sandy shores are infauna, since living in burrows
has the benefit of protection from wave action, predation and desiccation. Life within the sand, however, also
poses challenges in finding food, gaining enough oxygen and also finding a reproductive partner. On the
shore, you should try to observe some common infauna (e.g. bivalves or worms) to determine what adaptations
these animals have for overcoming the difficulties of living within the sand.

Field Work - Distribution and Abundance of Sandy Shore Organisms:

1. Each group will be assigned a transect location at intervals along the beach.
2. With the help of your demonstrators, use the 50 m tape measure to locate your transect along the vertical
gradient of the shore. Start from the strandline at the top of the shore near the vegetation and move down
the beach to the low water mark. Note the height of the last high tide along your transect (i.e. the
strandline). Remember to record the location of your transect in relation to others on the beach.
3. Two students will work out the profile of the shore using the Cross-staff and metre ruler – they will do
this for every 1 m interval down the beach. START AT THE LOW WATER MARK AND WORK
UPWARDS (as the tide will be rising).
4. Divide the total length of the transect sequentially into sampling stations (3 m intervals along the transect;
your demonstrators will help you decide this interval) and place a quadrat at each station. You will then
form pairs (1 pair per station) and investigate the species diversity and abundance of animals within your

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quadrat. REMEMBER TO RECORD THE EXACT LOCATION (e.g. x m) OF YOUR STATION
ALONG THE TRANSECT.
(i) Firstly, record the surface epifauna. Remember that epifaunal species are mobile, so enclose the
quadrat area with a ‘wall’ of sand (to prevent them running away, or other animals moving into your
quadrat). Some epifaunal animals will burrow rapidly when disturbed, look out for this when you begin
your recording. Use the field identification guide provided (and ask staff/demonstrators if you get stuck!)
to identify your animals. Release them beyond the quadrat afterwards so they will not be re-encountered.
If there are any animals you are unable to identify, take a few photographs with a reference scale (e.g. a
ruler) and ask a demonstrator on return to the classroom. Make a careful note of the abundance of each
species (i.e. how many individuals of each species).
(ii) Dig up the sand in your quadrat to a depth of ~ 20 cm. Note the depth where the anoxic layer
appears (i.e. where there is blackening of the sand). Also note the particle size of the sand in your station.
Place the dug-up sand on the plastic sheet and very carefully sieve through it using the plastic sieves
provided and record the organisms present as infauna and their abundance (you may also see siphon tubes
as further evidence of infaunal animals). If there are any animals you are unable to identify, take a few
photographs with a reference scale (e.g. a ruler) and ask a demonstrator on return to the classroom.
Release the others beyond the quadrat so they will not be re-encountered. This will give you an
opportunity to observe escape-responses for some species (e.g. Mictyris).
5. When you have finished your sampling, please refill your holes with the excavated sand and replace any
animals you have been studying GENTLY back where you found them. Make sure you collect all the
equipment for transport back to the camp. Dispose of any rubbish (empty water bottles, candy wrappers
etc.) properly.

Interpreting Your Data

Back at the camp, each group will prepare a poster. There will be an introductory talk on how to present your
posters and graphs at 15:00, after which the demonstrators will help you in organizing your data for the poster
and the identification of any species brought back for closer examination. Your poster should be prepared for
a scientific audience, clear and informative, and include:
i. A graph to show the beach profile along your transect
ii. A list (and perhaps drawings) of all the animals you encountered
iii. Graphs to illustrate the abundance of KEY species along your transect.
iv. Any other information you feel is interesting or important.

PLEASE ensure your posters are put up for DISPLAY

Before 18:00 on Tuesday evening
GROUPS will present their results at 18:30

Prof Gray A Williams

(September 2025)

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Common sandy shore species (* = you are expected to know these species):

Latin Name Common Name Habitat Feeding type

Arthropoda: Chelicerata
Tachypleus tridentatus Horseshoe crab Surface Dweller Predator
Arthropoda: Crustacea
Ocypode ceratophthalmus Ghost crab Deep Burrower Predator
Scopimera intermedia* Sand-bubbler crab Shallow Burrower Mud Feeder
Mictyris brevidactylus* Soldier crab Temporary Burrower Mud Feeder
Tmethypocoelis ceratophora Buddha crab Temporary Burrower Mud Feeder
Gelasimus borealis Fiddler crab Deep Burrower Mud Feeder
Clibanarius longitarsus* Hermit crab Surface Dweller Scavenger
Callianassa japonica Mud-shrimp Shallow Burrower Filter Feeder
Upogebia major Mud-shrimp Shallow Burrower Filter Feeder
Alpheus brevicristatus Pistol shrimp Shallow Burrower
Mollusca: Bivalvia
Anadara granosa Bloody cockle Shallow Burrower Filter Feeder
Coecella chinensis* Shallow Burrower Filter Feeder
Dosinia japonica Clam Shallow Burrower Filter Feeder
Gafrarium pectinatum Clam Shallow Burrower Filter Feeder
Tapes philippinarum Cockle Shallow Burrower Filter Feeder
Marcia hiantina Clam Shallow Burrower Filter Feeder
Anomalodiscus squamosus False cockle Shallow Burrower Filter Feeder
Mollusca: Gastropoda
Batillaria zonalis* Mud snail Surface Dweller Mud Feeder
Cerithidea moerchii* Mud snail Surface Dweller Mud Feeder
Pirenella asiatica Mud snail Surface Dweller Mud Feeder
Clithon oualaniense* Polymorphic shell Surface Dweller Detritus Feeder
Reticunassa festivus* Nassa shell Surface Dweller Scavenger
Polinices sp. Moon-shell Shallow Burrower Bivalve Predator
Natica sp. Moon-shell Shallow Burrower Bivalve Predator
Philine orientalis Shallow Burrower Bivalve Predator
Annelida
Ceratonereis sp. Mobile burrower Detritus Feeder
Cirratulus cirratus Tube Dweller
Perinereis cultrifera Peacock worm Mobile Burrower Predator

Sipuncula
Sipunculus nudus* Peanut worm Deep Burrower Detritus Feeder
Echiura
Ochetostoma Innkeeper worm Deep Burrower Detritus Feeder
erythrogrammon*
Nemertea
Lineus sp. Ribbon worm Deep Burrower Detritus Feeder
Echinodermata
Archaster typicus* Starfish Surface Dweller Detritus Feeder
Salmacis sphaeroides Sea urchin Surface Dweller Detritus Feeder
Brachiopoda
Lingula anatine Lamp shell Burrower Filter Feeder

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th
Wednesday 15 – Hillside shrubland
OBJECTIVES: to learn to recognize common hillside plant species; to study ecological
succession in a degraded hillside habitat.
Learning outcomes:
At the end of the field trip, you should be able to:
• Apply plant identification skills in identifying some of the common tree and shrub species of Hong Kong
• Describe the succession process in Hong Kong’s hillside and the key plant species at different stages of
succession
• Calculate the above-ground biomass of trees and forests in Hong Kong

Hong Kong Shrubland: an Introduction

The topography of Hong Kong is very rugged and over 800 km2 of the land area is considered hilly.
Hillsides are thus the most extensive terrestrial feature in Hong Kong. The natural vegetation on these
hillsides would be tropical rainforest, rich in wildlife and with trees 25-35 m tall. None of the original
forest survives today, except as tiny, disturbed remnants in some remote and steep ravines that have
escaped complete destruction by Hong Kong’s increasing human population. Today, the most common
hillside habitats are grassland (about 15 % of the total land area), shrubland (21 %), secondary forest
(24 %) and artificial plantation (5 %) (Kwong et al. 2022). This forest is called “secondary” to
distinguish it from the original, or “primary”, forest. Most of the original forest was probably cleared
for farming, as shown by the presence of abandoned terraces for rice or tea cultivation on many hillsides.
When a previously cultivated area is abandoned, the site is quickly occupied by grasses. If there is no
further disturbance, this grassland is invaded by shrubs and trees, which shade out the grasses, replacing
them by shrubland and then forest. This process of vegetation recovery following disturbance is known
as ecological succession.
The large areas of grassland and shrubland in Hong Kong today are the result of this process of
succession to forest being slowed or prevented by human disturbances. Until the last few decades, the
harvesting of plants (mostly grasses, ferns and small shrubs) for domestic fuel was a major factor
preventing forest succession on uncultivated hillsides in Hong Kong. This harvesting of hillside biomass
has now stopped, and hill fires have become the major disturbance factor. There are probably no natural
fires in Hong Kong: all are started deliberately or accidentally by people. The worst times for hill fires
are when large numbers of people visit ancestral graves on hillsides during dry weather in Ching Ming
and Chung Yeung festivals and burn joss sticks and paper as offerings to the ancestors. There can be
more than 100 accidental hill fires in a single day during these festivals. Grassland dominates in
frequently burned areas, because the common grasses can quickly re-grow from their base after fire.
Ten years without fire is enough for the grassland to be replaced by shrubland. Secondary forest,
dominated by Machilus species, 10-15 m in height, can develop in a further 20-40 years, but forest
succession is apparently much slower on sites which are remote from tree seed sources or where the soil
has been highly degraded.
There is no sharp discontinuity between shrubland and secondary forest, and all the intermediate
stages can be seen on Hong Kong’s hillsides. In the early stages of shrubland formation, the dominant
woody species are low-growing, light-demanding, multi-stemmed shrubs, such as Baeckea frutescens,
Rhodomyrtus tomentosa, Rhaphiolepis indica and Eurya species. Taller shrubland (> 2 m and 5-20 years
old) includes more tree species, with a single main stem or trunk, such as Schefflera heptaphylla,
Mallotus paniculatus, Melicope pteleifolia, Zanthoxylum avicennae and Machilus species, in addition to
many shrub species. Eventually, the trees form a complete canopy, which shades out most of the lower-
growing shrubs.
The major controlling factors on the rate of succession are: fires, which favour grasses and the fire-
tolerant shrubs; seed dispersal, mostly by birds, which brings new shrub and tree species to the site; and

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competition between plants for light. Only plant species that are tall enough to reach the light, or shade-
tolerant enough to survive below the tall species, can survive to the secondary forest stage.
Many studies have been conducted in Hong Kong to understand the mechanisms of forest
succession and identify natural and artificial barriers to forest succession. The aim was to restore native
forest for biodiversity and ecosystem services. In more recent years, under the impact of global climate
change, and the fact that trees store carbon in woods, there are many studies trying to quantify the amount
of carbon stored in trees and hence forests. Reforestation has been widely seen as a major nature-based
solution in capturing atmospheric carbon and mitigating global climate change. Most studies focus on
estimating the above ground biomass (AGB) using allometric models. To do so, tree surveys are needed
in obtaining the necessary data on tree species, diameter at breast height (DBH), total tree height and
wood specific density.

References
Kwong, I.H.Y., Wong, F.K.K., Fung, T., Liu, E.K.Y., Lee, R.H. and Ng, T.P.T. (2022) A Multi-Stage
Approach Combining Very High-Resolution Satellite Image, GIS Database and Post-classification
Modification Rules for Habitat Mapping in Hong Kong. Remote Sens. 2022, 14, 67.
[Link]

What are we going to do? Field work

Groups will combine to work in pairs (i.e. Groups 1&2, 3&4 and 5&6) and will be assigned a site in the
countryside where they will set up a monitoring quadrat of 10m by 10m. Each pair of groups will have
six 10 m long ropes as transect lines to establish the quadrat. After arriving at their site, each pair of
groups should set up a 10 m x 10 m square quadrat using FOUR10 m long ropes. Then, use the other
two 10 m long ropes to dissect the quadrat into four 5 m x 5 m sub-quadrats. Each group will then be
divided into TWO subgroups, each with 5-6 people. Each Subgroup will work in one of the four 5 x 5m
areas of the quadrat. Each sub-group will have a large plastic bag for keeping plant samples, some tags,
a marker pen and Sellotape to label plant samples and tags, a 1.2 m long measuring pole plus a meter
ruler, and dial callipers. Field record sheets will be provided for taking data.

Before taking any data, observe the plant species in your 5 x 5m quadrat and ask your demonstrators for
help with identification. Then, for all free-standing woody plant species that are taller than 1.2 m (i.e.
trees and shrubs, but not climbers, grasses, ferns and herbs) in your quadrat measure their diameters at
1.2 m (i.e. DBH-diameter at breast height) and put them on the record sheet provided. Do not measure
plants if DBH is <1 cm, even if it is taller than 1.2 m. Then, use the marker pen to mark the point of
DBH measurement on the measured plant to avoid double counting. For plants that have more than 1
stem that is taller than 1.2 m, measure the biggest stem. Lastly, measure the height of the tallest stem of
a plant using the measuring pole and the meter ruler. More details will be given at the field course on
the data to be taken. Unknown species should be assigned a number (species 1, 2 etc.) and a specimen
(a twig with several leaves) should be collected and labelled for identification later in the laboratory.

Back in the laboratory, identify all the unknown plant species collected with the help of demonstrators.
A briefing on how to compile the data will be given. The method to estimate AGB will also be given.

You will very likely encounter the following plant species on that day. You may also consult the Hong
Kong Field Guides Series 3: Hillsides and other references prior to the field camp.

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Species Name Growth form Species name Growth form

Adinandra millettii * Shrub Ilex pubescens Shrub
Antidesma japonicum Shrub Litsea rotundifolia * Shrub
Antidesma venosum Shrub Machilus chekiangensis Tree
Antirhea chinensis Tree Maesa perlarius Shrub
Aporusa dioica Tree Melastoma candidum Shrub
Baeckea frutescens * Shrub Melastoma sanguineum * Shrub
Breynia fruticosa * Shrub Ormosia semicastrata Tree
Bridelia tomentosa Tree Pinus elliottii # Tree
Cinnamomun parthenoxylon Tree Pittosporum glabratum Shrub
Cratoxylum cochinchinense * Tree Psychotria asiatica * Shrub
Diospyros morrisiana * Tree Rhaphiolepis indica * Shrub
Diplospora dubia Shrub Rhodomyrtus tomentosa * Shrub
Eurya nitida * Shrub Rhus chinensis Tree
Ficus variolosa * Shrub Rhus hypoleuca Tree
Garcinia oblongifolia Tree Rhus succedanea * Tree
Gardenia jasminoides * Shrub Sapium discolor * Tree
Glochidion eriocarpum Shrub Schefflera heptaphylla Tree
Glochidion wrightii * Shrub Wikstroemia indica Shrub
Homalium cochinchinensis * Shrub Zanthoxylum avicennae Tree
* Very common species on the studied hillside, you are expected to know these
# Planted exotic species

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Hong Kong Hillside shrubland
Field Record Sheet
Group No.: _______ Date: ______________ Sub-quadrat: _____ Recorded by: ____________

No. Species DBH (mm) Note1

Dr Billy Hau
(October 2025)

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Thursday 16th October - Butterfly Count

OBJECTIVES: To learn to recognize some Hong Kong butterflies; to investigate patterns

of relative abundance of organisms; to become used to making field observations and
using scientific names.
Learning outcomes:
At the end of the Butterfly count you should be able to:
• learn to sample, handle and recognize Hong Kong butterflies to family and species;
• quantify patterns of relative abundance of species to determine which species are common/rare;
• suggest possible explanations for the underlying ecological causes of these patterns.

The ecological problem

The Earth is home to an estimated 8-10 million species, some of which are common and others that are rare.
What is the pattern of commonness and rarity? Are most species common, or are most rare? Or are they all
equally common (or rare)? What is the general pattern? Before we can answer that question, we have to
decide what we mean by ‘common’ and ‘rare’. A species can be widespread and present at many places but
occur at low densities only, whereas another species might occur at very high densities but occur in only a
few places. Which is common and which is rare? For simplicity, since we are studying only one place
during this day of the field course, the term ‘common’ will be used to refer to species that are abundant in
that place, and ‘rare’ will be applied to species that are present in relatively low numbers. So we can
rephrase our question to ask: what is the pattern of relative abundance of organisms in nature? Describing a
pattern is an important first step in any ecological investigation because it is only once we have described a
pattern adequately that we can begin to investigate the factors that cause it. So the process of description of
a pattern will always precede the process of finding an explanation for it. Of course, once we have an
explanation, we can use it to make a prediction (that can be tested) about the occurrence of such patterns in
other places (or at other times).
Why are we interested in patterns of relative abundance? The reason is that species are important to
humans precisely because of their commonness or rarity: for example, we want to protect pandas or tigers,
and we want to limit the numbers of rats, mosquitoes and cockroaches. Accordingly, we try to manage the
environment in a way that will increase the numbers of animals that are endangered (and hence rare) and
decrease the populations of nuisance species that are abundant. If we do not manage the environment
properly, or we mismanage it, then we tend to reduce the abundance of many species – perhaps endangering
them – but, at the same time, we may inadvertently make conditions more favourable for the few species
that thrive in human-modified environments.
During this day of the field course we are going to describe the relative abundance of one species-
rich group of organisms in the vicinity of the Pak Tam Chung and use them as a ‘model system’ to
investigate species-abundance relationships. By doing this you will gain some familiarity with the study
organisms and learn to recognise some of the more frequently-encountered species. Once we have
determined the overall pattern of rarity and abundance, then we can begin to think about possible
explanations for this pattern.

The study organisms

Plants, birds and mammals are the best-known and best-studied components of terrestrial biodiversity, but
they make up only a tiny fraction of the total number of species: probably less than 1%. Most species are
invertebrates, and most of them are insects. In contrast to the plants, birds and mammals, where most species
have already been described and named by scientists, the insects are still largely unknown. A million or so
insects have been given scientific names, but 80-90% of the global total of insect species probably remains
undescribed. The butterflies (order Lepidoptera) are an exception to this generalization. Because they are

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active during daylight, conspicuous and attractive they have been relatively well-studied; it is likely that
over than 90% of the global total of species (estimated at ~17,500, most occurring in the tropics) have been
described. This makes them an excellent ‘model system’ for illustrating the patterns of diversity in terrestrial
animals.
Despite centuries of sustained human impact, Hong Kong is home to ~240 butterfly species. (This is
a lot: UK, which is much larger, has only 62 butterfly species.) There are 9 families (see Table 1 below);
note that some scientists arrange the families in a slightly different way to Table 1. Each family includes a
number of common species that you are likely to observe around Pak Tam Chung or elsewhere in Hong
Kong.
• The Hesperiidae (56 species in Hong Kong; 8 common) can be recognized by their broad heads,
stout bodies, relatively small wings and, with a few exceptions, their dull colours. They look less like
‘typical’ butterflies than members of the other families (which are grouped together in the
superfamily Papillionaceae). When resting, hesperiids often hold their fore wings and hind wings at
different angles.
• The Papilionidae (21 species; 10 common) are mostly large and colourful butterflies (often
combining black with a brighter colour); some have a long 'tail' extending from the rear of each hind
wing.
• The Pieridae (22 species; 7 common) are medium-sized butterflies without ‘tails’; most are basically
white, yellow or orange, sometimes with black, brown or red patterns.
• The Lycaenidae (51 species; 11 common) are small (most < 2cm) to very small, and often have one
or more pairs of tiny filamentous ‘tails’ on the hind wings; but many species have blue upper
surfaces of the wings and greyish undersides.
• The Nymphalidae (86 species; 38 common) which includes the Satyrinae (18 species; 7 common,
usually medium sized, brown butterflies with ‘eye-spots’ on the wings) and the Danainae (13
species; 6 common, usually medium sized to large and can be very conspicuous), range from
medium to fairly large in size; many of them have bright colours and complex patterns.
• The Riodinidae (3 species; 2 common) are small reddish brown butterflies.

The Hong Kong Lepidopterists’ Society maintains an internet presence at [Link] and their site
contains good images of most local butterflies. Please visit it (click ‘Hong Kong Butterflies’ on the menu
bar) if you would like to prepare yourself for this field trip. Two field guides that you may find useful are
listed below.
Note that the butterflies are only a small fraction of the species in the order Lepidoptera – the
remaining species are known as moths (>2200 species). Most moths are nocturnal, but Hong Kong has
several species of day-flying moths that can be confused with butterflies if you do not look carefully. Moths
can usually be distinguished by their tendency to rest with the wings held horizontally, whereas most
butterflies rest with both pairs of wings erect above the body (but see the note about Hesperiidae above).
The antennae of moths are thread-like or feathery while those of butterflies are filamentous with a swollen,
club-like tip; moths also have relatively robust or thick bodies.

The methods and the data output

You will work in pairs within your groups: each pair will be given an identification sheet and a butterfly net.
Use the sheet to identify all of the butterflies that you see and enter the number of individuals of each
species seen on the sheet provided. Count only individuals that you are sure are the same as the species on
the ID sheet; if you are unsure about a species ID, either leave it out (i.e. do not count it) or include it by
counting it as a member of a similar-looking species that you have already seen (this is to avoid ‘false
positives’ and inclusion of new species that are incorrectly identified). You will be provided with a butterfly
net that can be used to capture those specimens that you cannot identify from a distance. Handle them
carefully and release them unharmed after you have examined them. You may encounter species that are
not included on the ID sheet; make a sketch of their appearance noting the colour and shape of wing
markings, or simply take a photo on your phone and we will try to ID them in the laboratory.

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At the end of the field session, add up the number of individuals of each species seen by each pair in
the group to provide a group total, work out the relative abundance of each (i.e. % each contributes to the
grand total) then rank the species by their relative abundance (from most abundant to least abundant). This
final list (species name, total for each and % in rank order) will be your group output from the fieldwork.
Once you have the list, the group can look at the pattern of relative abundance of butterflies and think about
reasons that may account for it. Be prepared to discuss your observations and explanations with the rest of
the class.

Reference books
Lo, P.Y.F. & Hui, W.L. (2004). Hong Kong Butterflies. Friends of the Country Parks, Hong Kong.
Yiu, V. (2004). Hong Kong Butterfly Watching Field Guide. Hong Kong Discovery, Hong Kong. (Only
available in Chinese: 饒戈, 2004. 香港觀蝶圖鑑觀蝶地圖. 野外動向, 香港)

Table 1. Common butterflies likely to be seen around Pak Tam Chung. This list is not exhaustive, but it
includes all species on the ID sheets. You do not need to know all of the species! But you should be
familiar with all of the butterfly families.

Scientific name Chinese name No. individuals seen (& comments, if any)

Papilionidae
Graphium agamemnon................................. 統帥青鳳蝶
Graphium antiphates ................................... 綠鳳蝶
Graphium doson............................................ 木蘭青鳳蝶
Graphium sarpedon ..................................... 青鳳蝶
Pachliopta aristolochiae .............................. 紅珠鳳蝶
Papilio clytia ................................................ 斑鳳蝶
Papilio helenus ............................................ 玉斑鳳蝶
Papilio memnon ........................................... 美鳳蝶
Papilio paris ................................................ 巴黎翠鳳蝶
Papilio polytes ............................................. 玉帶鳳蝶
Papilio protenor .......................................... 藍鳳蝶

Pieridae
Catopsilia pomona ....................................... 遷粉蝶
Delias pasithoe ............................................. 報喜斑粉蝶
Eurema blanda …......................................... 檗黃粉蝶
Eurema hecabe….......................................... 寬邊黃粉蝶
Pieris canidia ............................................... 東方菜粉蝶

Danainae (Nymphalidae)
Danaus chrysippus …................................... 金斑蝶
Danaus genutia …........................................ 虎斑蝶
Euploea core ……......................................... 幻紫斑蝶
Euploea midamus ......................................... 藍點紫斑蝶

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Ideopsis similis …......................................... 擬旖斑蝶
Parantica aglea …........................................ 絹斑蝶
Tirumala limniace ........................................ 青斑蝶

Riodinidae
Abisara echerius .......................................... 蛇目褐蜆蝶
Zemeros flegyas …........................................ 波蜆蝶

Hesperiidae
Astictopterus jama ....................................... 腌翅弄蝶
Borbo cinnara .............................................. 秈弄蝶
Iambrix salsala ............................................. 雅弄蝶
Odontoptilum angulata ................................ 角翅弄蝶
Polytremis lubricans .................................... 黃紋孔弄蝶
Suastus gremius ........................................... 素弄蝶
Tagiades litigiosa ......................................... 沾邊裙弄蝶
Udaspes folus ............................................... 薑弄蝶

Nymphalidae
Argynnis hyperbius ...................................... 斐豹蛺蝶
Ariadne ariadne ........................................... 波蛺蝶
Athyma nefte ................................................ 相思帶蛺蝶
Athyma selenophora ..................................... 新月帶蛺蝶
Charaxes bernardus ..................................... 白帶螯蛺蝶
Cupha erymanthis ........................................ 黃襟蛺蝶
Cyrestis thyodamas ...................................... 網絲蛺蝶
Euthalia lubentina ....................................... 紅斑翠蛺蝶
Hestina assimlis ........................................... 黑脈蛺蝶
Hypolimnas bolina ....................................... 幻紫斑蛺蝶
Hypolimnas misippus ................................... 金斑蛺蝶
Junonia almana ........................................... 美眼蛺蝶
Junonia atlites ............................................. 波紋眼蛺蝶
Junonia iphita .............................................. 鉤翅眼蛺蝶
Junonia lemonias ......................................... 蛇眼蛺蝶
Lexias pardalis ............................................. 小豹律蛺蝶
Limenitis sulpitia .......................................... 殘鍔線蛺蝶
Neptis clinia ................................................. 珂環蛺蝶
Neptis hylas ................................................. 中環蛺蝶
Pantoporia hordonia .................................... 金蟠蛺蝶
Phaedyma columella .................................... 柱菲蛺蝶
Rohana parisatis .......................................... 羅蛺蝶

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Lycaenidae
Acytolepis puspa …...................................... 鈕灰蝶
Artipe eryx …................................................ 綠灰蝶
Chilades lajus …........................................... 紫灰蝶
Cigaritis lohita ............................................. 銀線灰蝶
Everes lacturnus …....................................... 長尾藍灰蝶
Heliophorus epicles ….................................. 彩灰蝶
Iraota timoleon …......................................... 鐵木異灰蝶
Jamides bochus ............................................ 雅灰蝶
Luthrodes pandava …................................... 曲紋紫灰蝶
Nacaduba kurava …...................................... 古樓娜灰蝶
Prosotas dubiosa …...................................... 疑波灰蝶
Pseudozizeeria maha .................................... 酢漿灰蝶
Zizula hylax .................................................. 長腹灰蝶

Satyrinae (Nymphalidae)
Discophora sondaica ................................... 鳳眼方環蝶
Faunus eumeus ............................................. 串珠環蝶
Lethe confusa ............................................... 白帶黛眼蝶
Melanitis phedima ........................................ 睇暮眼蝶
Mycalesis mineus ......................................... 小眉眼蝶
Mycalesis mucianus ..................................... 平頂眉眼蝶
Ypthima baldus ............................................. 矍眼蝶

Note: while there will be no need for you to injure or kill any butterflies during this exercise, you should be
aware that Troides helena is protected in Hong Kong under the Wild Animals Protection Ordinance (cap.
170); in addition, Troides aeacus and Troides helena are included under the Protection of Endangered
Species of Animals and Plants Ordinance (Cap. 586).

Prof Tim Bonebrake

(October 2025)

Ecology & Evolution Field course 2025-26