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Lim Lee Hong, Susan (1952–2014) — monogenean systematist and Commissioner 2006–2014



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Lim Lee Hong, Susan (1952–2014) — monogenean systematist and
Commissioner 2006–2014
David I. Gibson & Peter K. L. Ng
Professor Lim Lee Hong, Susan, better known to her international colleagues and
friends as Susan Lim, or just ‘Susan’, died on 2nd August 2014 in Petaling Jaya,
Selangor, Malaysia, after losing a long fight with cancer. She was a very active
parasitologist, a full professor in the Institute of Biological Sciences at the University
of Malaya in Kuala Lumpur since 2003 and a Member of the International
Commission on Zoological Nomenclature since 2006.
Susan was the leading specialist in Malaysia and Southeast Asia on a group of
parasitic flatworms called the Monogenea. Most monogeneans are ectoparasites of
fishes; they are a relatively large group with about 5,000 described species. Some
monogeneans are of significant economic importance because, when they occur in
huge numbers, they can have a serious pathogenic impact on fishes, especially food
fishes cultured in farms. In a career spanning some 35 years, Susan established herself
as one of the top scholars in her field.
217Bulletin of Zoological Nomenclature 71(4) December 2014
Born on St Valentine’s Day in 1952 at Seremban in the State of Negeri Sembilan,
Susan was the second of three daughters of the owner of an oil palm and rubber
business. Educated by nuns in a Roman Catholic school, by the age of six she could
neither speak nor understand a word of English and failed all subjects except
arithmetic. Eventually, having moved to an all-boys school (causing quite a stir in
those days) for her Advanced Level studies, she nevertheless acquitted herself
exceedingly well. In 1971, she obtained a deserved place at the University of Malaya
in Kuala Lumpur to study zoology, eventually graduating with an honours degree. In
those days it was still quite dicult for an ethnic Chinese (and a woman at that!) to
obtain training abroad, so Susan remained at the University of Malaya for her MSc
and PhD, funding her studies as a careers tutor (1976–89). In 1978, she was awarded
a UNESCO scholarship to work on monogeneans for three months with Dr Kálmán
Molnár in Budapest, Hungary, and in 1982, she obtained a fellowship from the
USSR Academy of Sciences to spend three months in St Petersburg, Russia, working
with Prof. Oleg Bauer and Dr Alec Gusev; the latter was, at that time, the leading
world expert on monogeneans. Trained and inspired to study these parasites, she
started her PhD (Distribution and Diversity of Monogeneans in Freshwater Fishes of
Peninsular Malaysia) in 1980 under the supervision of Prof. Jose I. Furtado in the
then Department of Zoology. During this period, she also embarked on a new
venture – she married her colleague George Liew and was later blessed with a
daughter and a son.
Through the 1980s, Susan published actively, describing many monogeneans and
establishing herself as a key player in the field. In 1987, she completed her PhD,
remaining on the university staas a zoology tutor. However, her academic prowess
was such that she was promoted to a lecturer in 1989, after which she never looked
back, becoming a full professor by 2003. Although Susan continued her work on
freshwater monogeneans, she gradually transferred her attention to the marine fauna.
Publishing regularly in good international journals, she became well known inter-
nationally from her papers and from her active attendance at and participation in
international meetings. These tended to be specialist meetings, such as the Inter-
national Symposium on Ichthyoparasitology and the International Symposium on
Monogenea, to which she always contributed presentations of her work.
Initially, as Susan was working in a region where the fauna was little known, her
research was mostly at the alpha-taxonomic level. She described more than 100 new
species, several new genera and a new family. Taking into account these and her
specific re-assignments (together more than 200 taxa), she became the sixth most
prolific monogenean worker ever (and the foremost female worker). As her expertise
developed, she undertook major generic revisions with a wider geographical rel-
evance to workers throughout Asia and around the Indian and Pacific Oceans. These
included papers on Hamatopeduncularia,Thaparocleidus,Calydiscoides (one species
of which has subsequently been named C. limae), Triacanthinella,Neohaliotrema and
Neocalceostoma. These were followed by even more general revisionary works and
reviews, such as ‘Sundaic monogeneans and Gondwana’, ‘Dactylogyridean monoge-
neans of the siluriform fishes of the Old World’ and ‘Diversity of monogeneans in
Southeast Asia’. In 2002, she co-edited and contributed to an important book titled
Diseases and Disorders of Finfish in Cage Culture’. She also contributed to teaching
units on animal diversity for the Open University in Malaysia. Susan was an excellent
218 Bulletin of Zoological Nomenclature 71(4) December 2014
artist and quickly converted to making digital drawings, publishing some of the first
such illustrations of her group in colour. However, her interests were not all related
to classical morphology. Even in her early studies during the 1980s she published a
paper on the use of Jaccard’s Index of Similarity for distinguishing congeneric
monogeneans. Later, she developed an interest in functional morphology and
described an entirely new mechanism of attachment in the form of net-like structures
formed by the coagulation of secretions emanating from the posterior attachment
organ of some of her monogeneans. As reflected in the work of her students, in recent
years she also embraced a very wide range of topics and disciplines, including
ultrastructural and molecular studies, 3D imaging, biotechnology, information
technology and biodiversity database management. She was also heavily involved in
the development of a database of the metazoan parasites of Malaysian wild animals.
In addition to her university teaching duties, Susan supervised many postgraduate
students. Arpah Bt Abu, Tan Wooi Boon, Wong Wey Lim, Neeta Devi Sinnapah and
Theerawoot Lerssutthichawal all completed their PhDs under her guidance; she was
still supervising another five PhD students at the time of her death. In addition, 11
MSc students benefitted from her supervision. Susan arranged for some of these
students to undertake part of their studies abroad at Queens University, Belfast, UK,
and the University of Perpignan, France. Regular checks on her students meant that
she was a regular visitor to the Natural History Museum, London, to discuss joint
projects and examine material.
Susan had a great interest on passing on her expertise and in the training of
taxonomists for filling present and future roles in biodiversity and wildlife manage-
ment. She presented several talks on this topic at international meetings, emphasizing
the lack of available training and job-opportunities in taxonomy. In relation to this,
and mainly for younger people, in 2004 she organised a ‘Workshop on Parasitic
Invertebrate Collections & Relational Database Management’ and a ‘Forum on
Biodiversity Inventories & Data-sharing A Framework for Malaysia’ with a signifi-
cant international specialist involvement, and in 2006 she persuaded the editor of an
international parasitological journal to run a course on ‘Publishing in International
Journals’. In view of her interest in systematics, she became responsible for the
type-collection of the Zoological Museum at the University of Malaya and fought for
a national collection of natural history specimens. Since 1979 she had been a member
of the Malaysian Society of Parasitology & Tropical Medicine, was its Honorary
Secretary twice and was awarded a life membership in 2009. These were in addition
to serving on various university and national committees. Her productivity, the
quality of her work and sociable nature lead to increased international recognition.
This resulted in more travel opportunities and co-operative studies, with research
visits to Japan, India, Australia, Canada, New Caledonia, South Africa and various
European countries, the longest being a year spent at the University of Guelph,
Canada, in 1995. In 2006, Susan was elected into the International Commission on
Zoological Nomenclature; an international group of taxonomists tasked to manage
and regulate how zoological names are used. As one of only three women in the
ICZN at the time (and the only Malaysian ever elected to this prestigious body), she
brought her expertise on parasites to the international stage. Ever the advocate for
systematics, she defended the science of taxonomy tooth and nail, and was a perfect
candidate for the job!
219Bulletin of Zoological Nomenclature 71(4) December 2014
Susan was always positive, energetic and, as in the case of many women who have
made it to the top in a male-dominated world, had considerable strength of character,
defending her work (and the study of her animals) aggressively. More than one
reviewer has taken on the job of refereeing a Lim paper with some degree of
trepidation – her rebuttals sometimes had to be ‘moderated’ by co-authors! She
singularly disliked self-righteous and condescending characters; her scowl (and
growl) for such people was well known. She despised what she saw as unfairness and
cronyism, and was a firm believer in meritocracy; and few could ‘out-work’ her. This
advocacy often got her in trouble with senior management, but this never stopped
her. Courage characterised her many fights for fair treatment for staand students;
and she rarely gave in. One of us (PKLN) had on more than one occasion to calm her
down when she worked herself up over what she saw as wrongdoings; and got
growled at in the process for being too naïve or diplomatic! Susan always had this
‘fire in her belly’ – one of her remarks to DIG many years ago was proof that this
started young – in primary school as a prefect, she commented that ‘‘I preferred to
play than to guard and got a ticking ofrom my headmistress who told me that that
was not how a prefect should behave – I never like authority nor understand it’’. That
was Susan and until her last day – a fighter! This same ‘spunk’ made her a friend one
could count on through thick and thin. Her ‘defence’ of her animals and her science,
and her intolerance of prima donnas belied her more usual convivial nature. The fact
is, Susan was a genuinely nice person, with a good heart and jovial (and often cheeky)
disposition – always popular with international colleagues, always with a greeting
smile. She was never a person overly worried about her appearance – her sartorial
elegance usually extended to a t-shirt and pair of jeans; however, on occasions when
she got fully ‘togged out’, and Lim’s limbs made a rare appearance, she could look
rather stunning.
PKLN last saw her when the ICZN convened in Singapore in November 2013 to
discuss the fate and future of this organisation, as it faced a series of huge financial
and scientific challenges. Susan attended the proceedings and contributed in her usual
way – energetically and positively. As is typical of Susan – she dragged George down
to Singapore with her and, while she was engaged in ICZN matters during the day,
he was out in the field collecting parasites from marine fishes with an assistant! And
in the evening, she would look at parasites with them. Talk about work ethos and a
love for monogeneans! Encouraged by her active participation, and that she had
apparently overcome her fight with cancer, the ICZN was hoping she would be a
force for change. Sadly, the illness returned and, this time, she lost the fight. Her old
friend and mentor, Dr. A. Sasekumar, told PKLN that she was true to form right to
the very end, talking about science with her usual passion the day before she passed
on. Vintage Susan!
Susan’s passing, at the young age of 62, is a great loss – not only in terms of
expertise to her country but to science as a whole. Susan was a good friend to many
and will long be remembered by monogenean specialists throughout the world for her
contributions. Immortal in the form of the many new taxa she described, her
influence will also live on in the form of her students, her published work and in the
memories of her family, friends and colleagues.
220 Bulletin of Zoological Nomenclature 71(4) December 2014
Full-text available
Background. Anchors are one of the important attachment appendages for monogenean parasites. Common descent and evolutionary processes have left their mark on anchor morphometry, in the form of patterns of shape and size variation useful for systematic and evolutionary studies. When combined with morphological and molecular data, analysis of anchor morphometry can potentially answer a wide range of biological questions. Materials and Methods. We used data from anchor morphometry, body size and morphology of 13 Ligophorus (Monogenea: Ancyrocephalidae) species infecting two marine mugilid (Teleostei: Mugilidae) fish hosts: Moolgarda buchanani (Bleeker) and Liza subviridis (Valenciennes) from Malaysia. Anchor shape and size data (n = 530) were generated using methods of geometric morphometrics. We used 28S rRNA, 18S rRNA, and ITS1 sequence data to infer a maximum likelihood phylogeny. We discriminated species using principal component and cluster analysis of shape data. Adams’s Kmult was used to detect phylogenetic signal in anchor shape. Phylogeny-correlated size and shape changes were investigated using continuous character mapping and directional statistics, respectively. We assessed morphological constraints in anchor morphometry using phylogenetic regression of anchor shape against body size and anchor size. Anchor morphological integration was studied using partial least squares method. The association between copulatory organ morphology and anchor shape and size in phylomorphospace was used to test the Rohde-Hobbs hypothesis. We created monogeneaGM, a new R package that integrates analyses of monogenean anchor geometric morphometric data with morphological and phylogenetic data. Results. We discriminated 12 of the 13 Ligophorus species using anchor shape data. Significant phylogenetic signal was detected in anchor shape. Thus, we discovered new morphological characters based on anchor shaft shape, the length between the inner root point and the outer root point, and the length between the inner root point and the dent point. The species on M. buchanani evolved larger, more robust anchors; those on L. subviridis evolved smaller, more delicate anchors. Anchor shape and size were significantly correlated, suggesting constraints in anchor evolution. Tight integration between the root and the point compartments within anchors confirms the anchor as a single, fully integrated module. The correlation between male copulatory organ morphology and size with anchor shape was consistent with predictions from the Rohde-Hobbs hypothesis. Conclusions. Monogenean anchors are tightly integrated structures, and their shape variation correlates strongly with phylogeny, thus underscoring their value for systematic and evolutionary biology studies. Our MonogeneaGM R package provides tools for researchers to mine biological insights from geometric morphometric data of speciose monogenean genera.
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