[*Tales and Drawings *
of a life by the sea]
Emeritus Professor of Marine Ecology The University of Hong Kong Hong Kong SAR, China
Hardback edition First Published in 2014 by aSys Publishing
eBook edition First Published in 2017 by aSys Publishing
Copyright © 2016 Professor Brian Morton
Professor Brian Morton has asserted his rights under ‘the Copyright Designs and Patents Act 1988’ to be identified as the author of this work.
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Table of Contents
The Tales and Drawings
Zebra mussel, Dreissena polymorpha
Archaster typicus and its parasites: Mucronalia fulvescens and Balcis shaplandi
Eelgrass, Zostera nana
Hemifusus ternatanus feeding on Pinna pectinata
Blue-ringed octopus, Hapalochlaena maculosa
Alpheus rapacida and Vanderhorstia mertensi
Nassarius glans particeps
Gari costulata and Moerella donacina
Phyllosoma larva of Panulirus argus
Surf clam, Donax semigranosus
Widgeongrass, Ruppia maritima
Buoy barnacle, Dosima fascicularis
Red-lined bubble shell, Bullina lineata
Congeria kusceri and[_ Marifugia cavatica_]
Salt-pan plants of Lake Macleod, Western Australia
Dumpling squid,[_ Sepioloidea lineolata_]
Cominella eburnea, Natica gualteriana and Lepsiella hanleyi
Leaf oyster,[_ Dendostrea folium_]
Scintillona cryptozoica and Scintillona daviei
Polinices incei and Donax deltoides
Atlantic mackerel, Scomber scombrus
Plough-snail, Bullia rhodostoma and by-the-wind sailor, Velella velella
Gilthead seabream, Sparus aurata
Sea grape, Coccoloba uvifera
American jacknife clam, Ensis americanus
Paradise threadfin, Polynemus paradiseus
Golden grey mullet,[_ Liza aurata_]
Yellow broom-rape, Cistanche phelypaea
Pelican’s foot, Aporrhais pes-pelecani
Sea rocket, Cakile maritima
Beach pea, Lathyrus japonicus
Silverfish, Trachinotus ovatus
Sprat, Sprattus sprattus
Silver sea stock and dry purslane
Chinese mitten crab, Eriocheir sinensis
Athanas dorsalis and Anthocidaris crassispina
Curled picarel, Centracanthus cirrus
Monrovian surgeonfish, Acanthurus monroviae
Atlantic emperor, Lethrinus atlanticus
Red-bellied piranha, Pygocentrus nattereri
Armoured catfish, Hoplosternum littorale
Tropical two-winged flying fish, Exocoetus volitans
Sea purslane, Halimione portulacoides and Bembidion (Cillenus) laterale
Four-horned spider crab, Pisa tetraodon
Boarfish, Capros aper
Mediterranean sand smelt, Atherina hepsetus
Newly settled juvenile of the cockle, Cerastoderma edule
King angelfish, Holacanthus passer
Scampi, Nephrops norvegicus
Atlantic herring, Clupea harengus
Shame-faced crab, Calappa granulata
Dyschirius impunctipennis and Dyschirius salinus
Bay squid, Photololigo duvaucelii
Burrowing eel goby, Trypauchen vagina
Blue-rayed limpet, Patella pellucida
This book is dedicated to all those friends who persuaded me to publish some of my drawings and the written vignettes that accompany them. But, it is especially dedicated to those friends, former students, colleagues and those precious soul mates who have walked on the shores and sailed the seas of this journey of discovery with me.
‘Science and art, the highest cultural achievements of humankind’
Irenäus Eibl-Eibesfeldt. In: Haeckel, E. 1998. Art forms in Nature. The prints of Ernst Haeckel. Munich and New York: Prestel.
This e-book is provided by the author for private use only. All information and illustrations in the book are the copyright of the author. Full colour, high quality hardback copies of the book may be purchased from the author at [email protected] and at a cost plus postage provided on request.
I have few recollections of, as a boy, neither drawing nor crayoning at home nor of any school art classes. Having failed my (the old) 11 plus examination, and leaving Hertfordshire for West Sussex, I was assigned to Littlehampton County Secondary School for Boys. Here, in the early years of the 1950’s, I was clearly destined for a vocational career and most instruction had a heavy bias towards woodwork, metalwork, gardening and the like and the general pursuit of toughening us all up with regular outdoor sports in all weathers and in the gymnasium. Some of us were also, from the age of about 14, enrolled in the Duke of Edinburgh’s Outward Bound scheme. This seemed, at that time, to involve no more than lots of hiking, climbing, hare and hounds running and corporal punishment for any minor infringement of rules poorly understood. I did, however, gain a love of English, both its language and its literature, through a more enlightened teacher, Mr Don Long. I also developed a mind adept with mental arithmetic through the simple but effective expedient of obtaining a cane across the hand for every one of the questions got wrong during each morning’s quiz of ten. This took up some time with a class of 20 or so boys – the caning not the quiz, I mean. And I cannot remember the name of the teacher responsible, if that is the right definition of him.
At the age of 15, my parents discovered that the school had no (again old) O[rdinary] level stream, in fact there was not a single examination a boy could gain a qualification in and I was duly removed from its premises and sent to Crawley Technical College where I was enrolled in the said stream. O levels obtained –many in science subjects - I remained at Crawley to complete (again old) A[dvanced] level examinations, again in science subjects -zoology, botany and chemistry - but obtained with not overly impressive grades.
At that time, botany and zoology practical classes mostly involved simple experiments but mainly comprised observations and the recording thereof, and an understanding of anatomy through pencil drawings of animal dissections and examination of plants. Except for the zoology teacher, Mr A. Jack, who was studying for an external London Ph.D. degree on mites. He asked for a volunteer to stay on after classes to help with this, involving the drawing of their tiny parts, again with a pencil, and as seen through a microscope with a camera lucida attached. To the amusement of the other boys and girls, I volunteered. He then took my pencil drawings home for inking and, I assume, labelling but I never saw the finished products. Perhaps because he did not finalise them before I left Crawley.
Post school, I hesitated about going to University and took a year off to work as a research assistant at the Glasshouse Crops Research Institute in my home town of Littlehampton. I was assigned to Mr J. Hesling to work with him on heteroderan nematodes. He too was reading for an external London Ph.D. degree and hearing of my work with Mr Jack at Crawley, I was, again, assigned a microscope with a camera lucida. And, after dissecting out the miniscule genital apertures of various pest nematodes with the finest of needles and mounting them on slides, I was also given the task of drawing them with a pencil for him too to finish up inking at home in the evenings.
A year of this was enough and I decided to try for a university place. I knew I had no chance of obtaining a placement either at the Oxbridge or red-brick universities and so plumped for London and, particularly, Chelsea College, which at the time was negotiating for full collegiate status of the University of London. In this, I was encouraged by my uncle, Frank Morton CBE, who was then Professor of Chemical Engineering at Manchester University and founding Vice-Chancellor of the Unversity of Manchester Institute of Science and Technology. He was, as too was my father, a committed socialist, friend of Harold Wilson and very keen on helping students such as I to obtain tertiary level education. I can rember being told (with great pride) as a young boy that my grandfather had taken these two boys to see the Scottish socialist and labour leader Keir Hardie (1856–1915) being installed, with a cloth cap, not a top hat, into the Houses of Parliament of the United Kingdom as the first elected Independent Labour Party Member. But, why Chelsea?
The answer to this lies in its foundation as the South West Polytechnic in 1895, becoming Chelsea Polytechnic in 1922, Chelsea College of Science and Technology in 1966 and a constituent college of the University of London, as simply Chelsea College, in 1971. The college occupied a single site on the corner of Manresa Road and King’s Road in Chelsea (SW3) and, bearing in mind the importance of such a location, with the famous Six Bells Public House opposite, to a late teenage lad in the swinging sixties, where else would anyone in their right mind not wish to be a student? As an aside, Chelsea merged with nearby Queen Elizabeth College in 1985 and soon, thereafter, the merged colleges were themselves amalgamated into King’s College. I now, thereby, find myself as an alumnus of one England’s most famous and prestigious colleges by fortuitous default. An alma mater via the delivery-man’s back gate as it were. Quite appropriate, I do, however, feel as the son of a working class family and inheritor of Keir Hardie’s cap.
But, I still had to gain admission to Chelsea and there I think my Uncle Frank had a hand, because, by virtue of history, the college was steeped in socialism. Its origins are exceptionally old and have been documented by S. J. Teague (1969), the college librarian when I was in attendance. At the time I was seeking admission, the college still stuck to its socialist roots and, hence, I was by all accounts an eminently suitable applicant, which I think Uncle Frank put to his like-minded London academic friends. That is the way it was in those days and maybe still is and always will be, to some extent at least. But I still had to be accepted, after being offered an interview.
My interview, in early 1963, was with the newly appointed Professor of Zoology, R.D. Purchon (1916-1992) (Morton 1993), recently returned to England from his Foundation Raffles Chair of Zoology at the still colonial University of Singapore, via a mutually disagreeable and very brief sojourn at the University of Ghana, Legon Accra. I think it fair to say that Dick Purchon and I were as different as chalk and cheese. From his military, prisoner-of-war and colonial background, he sat, stood and walked erect, had an officer-corp bearing complete with immaculately trimmed moustache and a clipped accent. For some inexplicable reason, however, he either seemed to like me or, more likely, took sympathy on me (I was, however, later to discover it was the former) and I was, there and then, accepted to read for an honours degree in zoology with physiology as a first year ancillary subject. I was very happy to accept.
The physiology component of the degree was fascinatingly experimental, but the zoology was classical anatomy based. A whole year of invertebrates, a second of vertebrates and a third of miscellaneous topics – such as embryology but also with two lectures on behaviour and two on ecology and no field work. Very different from the modern biological curriculum. Practical classes were a stream of microscopic and dissecting experiences, the subjects of which all had to be drawn with a pencil and were assiduously corrected and marked. I graduated in 1966. Before leaving, however, and much to my surprise, I was invited to see Dick Purchon and there and then he offered me a University of London Studentship to read for a Ph.D. under his supervision. I was to be his very first higher degree student. The subject was the invasive freshwater mussel, Dreissena polymorpha Pallas, 1771.
Importantly, however, Dick told me that I could research any aspect of the zebra mussel I wished but that, first of all, I had to draw its anatomy, starting with the shell, then the living body and finally examine it microscopically. No photographs were to be taken. We ordered paper (Bristol Board) and Rotring pens ® and ink and I set to work. Every illustration I drew was scrutinised by Dick on which occasion he pointed out errors in dimensions, scale and configuration. ‘Is it _][*really *][_like that?’ he would ask sniffily. The offending diagram was torn up, sometimes by him, and I, shocked, started again. Over and over and over again. Finally, he would say that the latest version was acceptable and I would begin anew on another drawing. His philosophy was simple. One had to know one’s subject intimately before one could even begin to contemplate researching other aspects of it. By this expedient, however, my first paper on the anatomy of [_Dreissena _]was published before I graduated, which happened in early 1969 (Morton 1969a). My first lesson in biological, and by that I mean scientific, drawing had been learnt – be as accurate as possible. And, it is true: one does not know either a plant or an animal until one has drawn it. Something that has been largely forgotten in modern biological degree courses.
Without going into further career details, towards the end of 1969, I was offered an assistant lectureship in Zoology at the University of Hong Kong. Once again, this was teaching a still classical zoology and botany honours degree except, of course, many of the demonstrated species were wholly new to me and the students were all Chinese. With a very amateurish local natural history society, little local appreciation of the native flora and fauna, an absence of conservation awareness and, save for a small flock of expatriate bird-watchers, little interest in nature photography, I had to put my infant drawing skills to work. I also had to learn a whole new way of looking at the life around me. And, with no like-minded colleagues to work with, I was floundering in a sea of sub-tropical marine life of amazing diversity such as I had never encountered in Great Britain and that was virtually unknown and unstudied. In order to get to grips with this, something had to be done.
My true intellectual mentor was Sir Charles Maurice Yonge FRS (1899-1986) (Morton 1992), Dick Purchon’s Ph.D. supervisor and, then, still occupying his Regius Chair of Zoology at the University of Glasgow, but shortly to move on to the University of Edinburgh. His simpler, yet nevertheless illuminating, illustrations for the research papers he published, provided me with another aspect of scientific drawing. Do not embellish and do not make the subject too complicated. In Hong Kong, I continued to correspond with Maurice and he suggested I contact a namesake (of mine) biologist in New Zealand, Professor John Morton (1923-2011) (Morton 2011). In 1968, John had published (with Michael Miller) [The New Zealand Sea Shore. _]This was a revolutionary book and, after reading it, I knew I needed to work with this man. John and I corresponded and, through Maurice, I was awarded a Leverhulme Fellowship to work with John for three months in the summer of 1972 at the University of Auckland. John Morton was a true eccentric genius: a brilliant lecturer, world-renowned scientist, popular educator and a talented illustrator of equally wonderful research papers and books. I learnt something from him of how to illustrate the complexities of intertidal life and eventually we would together write _The Sea Shore Ecology of Hong Kong (Morton and Morton 1983). It took me that long to just begin to understand the complexity of Hong Kong’s marine life.
In 1977, I was invited to be the keynote speaker at a conference in Fort Worth, Texas on the ontroduced bivalve Corbicula, organised by Professor Joe Britton (1942-2006). From that very first meeting, Joe and I became the closest of friends and colleagues – we were, for the start, almost exactly the same age and, more importantly, our research interests were wholly similar. We wrote numerous research papers and two books together – Seashore Ecology of the Gulf of Mexico (1989) and Coastal Ecology of the Açores, the latter in co-operation with Professor Antonio M. de Frias Martins (b. 1946), Professor of Biology at the University of the Açores on the island of São Miguel and another good friend and colleague. Tragically, Joe died in 2006 (Morton 2007a) and I lost one of the nicest men ever to embrace biology.
A final hero of mine was Thomas Alan Stephenson (1898–1961), a British naturalist and marine biologist, specialising in sea anemones. Alan developed an interest in natural history from an early age and went to University College, Aberystwyth, where he began to study the local sea anemones. Subsequently, Stephenson held a number of academic posts in Great Britain and at the University of Cape Town, South Africa, completing his career at his alma mater in Aberystwyth (Yonge 1962). I never met Alan Stephenson but for me and all coastal ecologists of my generation, his seminal work with Anne, his wife, on intertidal communities (Stephenson and Stephenson 1973) holds a special place in my professional life and, moreover, assisted me greatly in the illustration of my own studies of Hong Kong’s marine life.
Through Dick Purchon, Maurice Yonge, John Morton and Joe Britton, plus Alan Stephenson and Tony Frias Martins, therefore, I obtained the tools that would enable me to commence a career in the two areas of molluscan research and intertidal ecology in particular and to be able to illustrate my work if not artistically, at least satisfactorily for professional and popular readers alike in published manuscripts and popular books, respectively. This volume, therefore, represents a series of mostly un-published illustrations undertaken between 1966 and 2014, that is, they encompass almost fifty years of marine and freshwater study.
The portfolio of illustrations presented herein comprises an ecletic, miscellaneous, assortment of illustrations, not the usual sketches and paintings of lions, giraffes and elephants, more the smaller creatures of the sea and shore. The illustrations also reflect my own special interests of the plants and animals of the coastline and the intertidal, particularly the Mollusca that are my speciality, especially the typically sedentary bivalves, but also gastropods which have found so many ways of obtaining nutrition. From, particularly, and circuitously, their bivalve cousins, the most prolific and nutritious prey available to them at the sea’s margin. On reflection, however, I seem to have drawn, at some time or other, representatives of most invertebrate groups, even the ciliate protozoan Folliculina, but perhaps the most rewarding of creatures are the fishes, to be either caught or discovered in seafood markets the world over and providing the most wonderful examples and stories about the group’s adaptive radiation as well as proving to be a delicious and nutritious meal subsequently.
Most of the illustrations have never been published before and where they have, an acknowledgement is made. And these particular illustrations are included because of some especially tantalising snippet of information about them. The reader may also become aware that the number of the un-published drawings has increased approximately year by year. This is because, early in one’s teaching and research career, there was less time to engage in recreational illustrations, one’s focus, of necessity, being on more explanatory ones, suiting the purpose of adorning scientific facts with appropriately scientific figures. Only later, a family grown up, release from teaching, higher degree student supervision, the tortures of academic meetings and yet more meetings, did time become available for more thoughtful, esoteric and, hopefully, diligent work – or rather recreation. Lastly, the reader may detect an improving trend as the years unfold with each illustration. I do hope so.
Today, in retirement, I still write, but indulge myself more with a pen and ink and the reader may, finally, observe that over the years and as the drawings are perused, I am turning to colour. This is a reality and an ongoing experiment with life’s challenges. For, that is the beauty of drawing, particularly for one’s own pleasure. One can experiment continually, be critical in private and, I have to say, that the concentration involved relaxes the mind, opening it up to ancillary contemplations, suggesting new projects and the continued planning for new and exciting promises of discovery.
The gifts I was given by my mentors, friends, colleagues and heroes thus remain with me and are fulfilled today as an appreciation of them, their kindnesses to me, their talents but especially their services to the marine biological sciences and, thereby, the greater eduction and edification of all whom they have served so diligently.
1 May 2014
The portfolio represents illustrations undertaken over a period of some 48 years from my first attempts to draw Dreissena polymorpha (Drawing No 1) as a Ph.D. student under the critical eye of my supervisor, Professor R.D. Purchon, to the present day as a retired academic. I have a lot to thank Dick Purchon and my spiritual mentor Sir Maurice Yonge and other scientific colleagues for, but I am deeply grateful to all those past students, friends and colleagues in many parts of the world including Hong Kong, China, Australia, Croatia, Denmark, the U.S.A., the Açores and the United Kingdom who have both facilitated and encouraged me to publish this element of my portfolio of biological illustrations. I am, however, especially grateful to my friends Dr K.F. Leung and Dr S.F. Leung, both of Hong Kong, for their unswerving enthusiasm for this book project. Nathalie Yonow, Swansea University, U.K. is thanked for commenting on the drawings and reading the first drafts of the Opisthobranchia vignettes, not one of my specialities, and for her similar enthusiasm for the book. Professor R.S.S. Wu, Head of the School of Biological Sciences at the University of Hong Kong, Professor K.M.Y. Leung and Helen Leung of the same school, not forgetting Sylvia Yiu of the Swire Institute of Marine Science, also the University of Hong Kong, are acknowledged for their support of the project. Notwithstanding, any errors of either fact or illustrative detail are entirely my own and for which I take full responsibility.
The history of natural history illustration has been reviewed and, often, copiusly illustrated by many authors including Dance (1978) and Desmond (2003) – two volumes which the reader is referred to for a greater appreciation of the subject. This then is not a history of natural history illustration but a personal view of how one’s own interests influence what one does in life. This is because one cannot be an illustrator without reading and perusing what has gone before. This personal view of natural history illustration has been mostly gleaned from books in my own library, the plants and animals that I sometimes research and experiences as a career biologist. This is not, therefore, a comprehensive history of biological illustration, but my own impressions gained during an academic career in the illustrative sciences – botany and zoology.
From prehistoric times, coinciding with the earliest known evidence for [_Homo sapiens _]in Europe, that is, some 40,000 years ago, there is evidence of cave painting scattered throughout some 350 sites mostly in Spain, France and Africa. The oldest known cave art comes from El Castillo in northern Spain.The second-oldest known is that of Chauvet Cave in the Ardèche department of France, the finest gallery of cave art, and paintings which may be 35,000 years old, from the Upper Palaeolithic.
Plate 1 shows a group of, presumably, wild horses. Other examples of cave art may date from as late as the Early Bronze Age (~15,000 years ago) but the style seen here and Altamira in Spain died out around 10,000 years ago, coinciding with the advent of the Neolithic.
The most common themes in cave paintings are large wild animals, such as bison, horses and deer. In the Chauvet Cave, there are illustrations of mammoths with spears in them, either anticipating or recording successful hunts. In the cave of Cueva de las Monedas in Spain, drawings of reindeer provide evidence for the last Ice Age.
A wall painting of horses in the Chauvet Cave in the Ardèche, France. The paintings are over 35,000 years old.
The Tito Bustillo site is part of the Altamira Caves in northern pain and is a declared UNESCO World Heritage Site dated between 30,000 and 13,000 years ago. Almost all the site is painted, the prehistoric representations varying according to the period and are superimposed one over the other, depending on the preferences of the cave’s inhabitants. In the Hall of the Horses, the paintings are coherent with the natural architecture of the cave’s form so that they are virtually three dimensional. Of all the pictures of animals, there is one of a cetacean that is particularly interesting, as these animals are very infrequent in prehistoric art and presumably derive from strandings, in turn suggesting early coastal gleaning.
In southern Africa, the cave art of the San people, or bushmen of the Kalahari, has hardly changed for 20,000 years and is still added to today. Their greatest masterpiece is the spectacular cavern in Zimbabwe called Inanke Cave. It is located high in the Matobo Hills, not far from Bulawayo. Inanke is called the Sistine Chapel of cave art and at its centre is a ten metre-long frieze of scampering, surging, stamping animals. Giraffe, eland, kudu and many more species are all painted with exquisite precision on the cave’s roof. A reflection of man’s relationship with the species he shares this planet as well as a spiritual understanding of that bond.
In Australia too, cave paintings have been found on the Arnhem Land Plateau showing megafauna which are thought to have been extinct for more than 40,000 years. This makes the site a candidate for the oldest known cave paintings. Another site, Nawarla Gabarnmang, has charcoal drawings that have been radiocarbon-dated to 28,000 years, making it the oldest dated site in Australia and among the oldest in the world for which reliable dating evidence has been obtained. As in Europe, some caves probably continued to be painted for a period of several thousands of years.
The illustrations, in their homes (albeit caves), of the animals people saw around them and hunted, shows that such activities are a basic feature of humanity occurring in widely-separated Europe, Africa, Australia and Asia. Throughout our subsequently recorded history, such painting and drawing has progressively become our most visually obvious art form. Today, it adorns the walls of our institutions and our homes. But, it also brings our books, magazines and journals to life – a picture speaking a thousand words. Indeed, before writing and reading became a near-universal pastime, illustrative art was the principal means of not just adornment but also of visualising thoughts and effecting communication. Because of this, such art spread rapidly throughout the world.
Mythical winged horse. Wei and Jin Dynasties, Xinjiang Province China (3rd century AD). (Photo: B. Morton).
To illustrate this, Plate 2 is a carved mythical winged horse excavated from 3rd century AD sites dating from the Wei and Jin Dynasties, Xinjiang Province, China while Plate 3 depicts a hunting scene adorning the roof of grotto No 249 from the Mogao Cave complex at Dunhuang, Xinjiang Province, China. This too dates from the Western Wei Dynasty (304-589 AD).
Hunting scene, Cave No 249, Mogao Caves, Dunhuang, Xinjiang Province, China. Western Wei Dynasty (304-589 AD. (Photo: B. Morton).
Prior to and up to the above Chinese dates, in Europe, the Romans had perfected the art of mosaics. A floor mosaic showing a stylised dolphin from the 3rd century Roman villa at Bignor in West Sussex, England, is shown in Plate 4.
A floor mosaic showing a stylised dolphin from the 3rd century Roman villa at Bignor in West Sussex, England. (Photo: B. Morton).
Plate 5 shows a more natural representation of a bird and a snail (with an axe!) and derives from the apse of the Basilica di San Clemente in Rome, Italy. This, however, is a 12th century piece of art but, nevertheless, reflects the sustained influence of the Romans upon the subsequent mosaic adornment of significant buildings. The Romans also decorated the internal walls of the houses and this is best seen today at Pompei in the shadow of the city’s demise – the volcano of Vesuvius.
A bird and a snail with an axe. From the apse of the Basilica di San Clemente, Rome, Italy. (Photo: B. Morton).
Because the spread of writing and reading was slow, perhaps deliberately so, the very earliest hand-written books were sometimes profusely illustrated. The earliest bestiary was an anonymous 2nd century Greek volume called the Physiologus, which summarised knowledge about animals in the writings of classical authors such as Aristotle’s Historia Animalium and the works of Herodotus, Pliny the Elder, Solinus, Aelian and the other early students of the natural world. Later, bestiaries became a compendium of creatures. These beastiaries reached a zenith of popularity during the European Middle Ages (5th to 15th centuries) in illustrated volumes, or Bestiarum Vocabulum, that described various animals and birds. An illustration of each animal was usually accompanied by some aspect of its natural history, providing an additional moral point, that is, for the time – a religious lesson. This was because, it was believed that every living thing had its own particular meaning. Soon, St Isidore of Seville and St Ambrose elaborated upon the religious message with reference to passages from the Bible. The often fanciful accounts of illustrated beasts began to be read more widely and generally were believed to be true.
Medieval bestiaries were particularly popular in England and France around the 12th century and comprised mainly compilations of earlier texts. The Aberdeen Bestiary is one of the best known of some fifty such texts surviving today. Two illuminated Psalters, the Queen Mary Psalter and the Isabella Psalter, contain full bestiary cycles. The bestiary in the Queen Mary Psalter is configured with marginal decorations that typically occupy the bottom quarter of each page. This bestiary was expanded beyond the source in the Norman bestiary of Guillaume le Clerc to include ninety animals. Others are placed in the text to cohere with the psalm they are illustrating. Even Leonardo da Vinci made his own bestiary.
In more modern times, artists such as Henri de Toulouse-Lautrec and Saul Steinberg have produced their own bestiaries. The Argentine poet Jorge Luis Borges (1899-1986) wrote a contemporary bestiary of sorts – the Book of Imaginary Beings – which collected surreal creatures from bestiaries and fiction. Most recently, the American, John Henry Fleming’s Fearsome Creatures of Florida published in 2009 borrows from the medieval bestiary tradition to impart moral lessons but not about religion but about the environment. This bestiary was illustrated by David Hazouri. The Englishman, Caspar Henderson’s The Book of Barely Imagined Beings published in 2013 is actually subtitled A 21st Century Bestiary and deals with an eclectic range of creatures, reflecting the human condition. For the academic biologist, however, Professeur Dr Harald Stümpke’s [_Anatomie et Biologie des Rhinogrades, un Nouvel Ordre de Mammifères _](1962) is the most compellingly and amusingly modern bestiary combining anatomy, biology and humour, without the necessity of either a religious or environmental moral message – simple plain fun. The actual author and illustrator of Professor Stümpke’s monograph was Gérolf Steiner who, by inventing a new order of wholly fictitious mammals that moved around using their noses (Plate 6), proved that even, normally austere, taxonomists have a sense of humour.
Modern writers of fantasy fiction, and the cinema films that arise from them, draw heavily from the beasts described in mythology and bestiaries. In fact, the worlds created in such fiction can themselves be regarded as bestiaries because the images portrayed each represent some moral attribute – good, bad, kind, evil and so on. Today too, the authors of fantasy role-playing games sometimes also compile bestiaries, for example, one of the most popular video games – Dungeons & Dragons.
Hopsorrhinus aureus. Reproduced from Stëmpke’s ‘Anatomie et Biologie des Rhinogrades un nouvel ordre de Mammifères’ (1962, Plate VI). The author and artist was Gérolf Steiner.
The invention of printing is credited to Johannes Gutenberg (ca. 1395–1468) a German blacksmith, goldsmith, printer and publisher who introduced printing to Europe in around 1439. By 1450, a fully operational press was in operation. In 1455, Gutenberg completed copies of a beautifully executed folio Bible (Biblia Sacra) – the so-called 42-line Bible which is now also known as the Gutenberg Bible although the text did not have a date nor a printer’s name. Nevertheless, until that time, books had been hand-written for the few. Gutenberg’s invention of mechanical moveable type printing, however, started the Printing Revolution, which, in turn, played a key role in the development of the Renaissance. Similarly, the Reformation was sparked by the publication in 1522 of Martin Luther’s 95 Theses , leading to the Age of Enlightenment and the Scientific Revolution. This, in turn, laid the basis for our modern knowledge- and material-based economy and the spread of learning to ordinary people, Gutenberg’s invention ultimately allowing the mass production of printed books.
Gutenberg’s former partner, Johann Fust (ca.1400–1466), with Peter Schöffer (ca.1425–ca.1503), published the Mainz Psalter in 1457, which was the first printed book complete with a date and publisher’s name. The English merchant and diplomat William Caxton (ca.1415/1422–1492) settled in Bruges, in modern Belgium, in around 1453. His trade also led to more continental travel, including a visit to Cologne, a short distance from Mainz, where he observed the new printing industry and was influenced significantly by it. So much so that on his return to Bruges, Caxton quickly set up a printing press there, in collaboration with Colard Mansion and, in 1473, printed the first book in English – Recuyell of the Historyes of Troye – a translation by the talented Caxton himself. This translation became so popular that he began to reproduce more copies, thereby bringing such books to a wider audience. Subsequently, bringing his talents back to England, he set up a press in 1476 at the Sign of the Red Pale in London at Westminster. The first book known to have been produced there was an edition of Chaucer’s The Canterbury Tales. He is, therefore, credited with being not only the first English person to work as a printer and the first to introduce a printing press into England but also the first British retailer of printed books and is credited with printing 108 editions of 87 titles.
At the dawn of printing, the illustration of books was rare. The Liber Chronicarum or Nuremberg Chronicle, published in 1493, was a ground-breaking book in several ways. It was, firstly, one of the only incunables (books published before 1500) to include illustrations. Secondly, it was one of the first books that paraphrased world history in relation to the Bible. Thirdly, it was the first printed book to successfully integrate hundreds of complex illustrations into its (Latin) text.
Intuitively, it would be forecast that such a book was quickly emulated. Not so, however, and Francis Bacon’s (1561-1626) Sylva Sylvarum or A Naturall Historie in Ten Centuries was not published until 1639 by William Lee of London.[* *]Even so, it featured only illuminated capitals and decorative headings and tailpieces. This did not last long, however and the Historiae Animalium Angliae tres Tractatus by Martin Lister, published in London in 1678, contained beautiful lithographic plates of shells, in particular. Martin Lister (1639–1712) was an English physician and naturalist, his career culminating when he was appointed physician to Queen Anne from 1709 until his death. Other than Historiae Animalium, his[_ ]principal works were _Historiae Conchyliorum (1685-1692) and Conchyliorum Bivalvium (1696).
Gilbert White (1720-1793) was an English naturalist and ornithologist. White is best known for his The Natural History and Antiquities of Selborne. This was a compilation of his letters to Thomas Pennant (1726-1798), the leading British naturalist of the day, and the Hon. Daines Barrington (1727-1800), an English barrister and naturalist. The letters and the book contained White’s discoveries about local birds, animals and plants, which he distinguished by observation rather than by collecting specimens. Even so, the original book, first published by the Ray Society in 1789, was un-illustrated although the Gilbert White Museum edition, first published in 1977, contains ten drawings by the contemporary artist Frederick Marns and five plates of bird drawings reproduced from Thomas Bewick’s History of British Birds published from 1797-1804 in two volumes, Land Birds and Water Birds, with a supplement in 1821. It is worth pointing out that Gilbert White’s work has been in print continuously since its first publication (1789) and is one of the most famous and frequently published books in the English language.
One of the most remarkable illustrated books on my own speciality – the Mollusca – was published between 1791 and 1796, in three volumes, by the Italian scientist[* ]Giuseppe Saverio Poli (1746-1825) in which not only are shells illustrated but also incredibly fine paintings of their anatomies in stunningly intricate (for the time) detail. A total of 57 plates were included in Poli’s Testacea Utriusque Siciliae published between 1791-1796 and they were all produced by a large range of artists and engravers. Eatly artists were Franc°. Morelli who drew plates 1, 2, 7, 14, 15, 20, 22-24, 36-40 and 42-43 and Lo Manto who produced plates 3, 4, 8-13, 16, 18, 19, 26, 28, 32 and 41. Among the engravers Ioh. Brun was responsible for plates 1, 2, 15, 25, 27, 29 and 41; Vin. Scarpati for 7, 9, 10, 12, 13 and 18; Scarpati for 7, 17 and 21; and Ant[onio] Zaballi for 11, 19, 22-24, 30, 31, 34, 38 and 39.[ *]
The Italian scientist Giuseppe Saverio Poli (1746-1825) who commissioned artists to illustrate his book ‘Testacea utriusque Siciliae……’ (1791). (From the book in the library of the Natural History Museum, London). (Photo: B. Morton).
Plate 7 shows, in profile,[* *]Giuseppe Poli who commissioned the artists to illustrate his outstanding book and Plate 8 is the book’s Plate XXII, which was painted by Franco Morelli and engraved by Ant[onio] Zaballi. In 18th century Italy, artists of this calibre must have been all too common and available to undertake commissioned work such as this and of this quality, much as mobile, television and computer repair shops are common today in our British high streets.
From this 18th century peak in illustrative excellence, the Englishman George Montagu (1755-1815) published his [Testacea Britannica, _]or _Natural History of British Shells (1803-1808) and, wherein, the illustrations are crude in comparison to those produced by Poli’s artists. To explain: some of the Poli illustrations are of mollusc dissections that would be considered acceptable by 20th century and, even, many 21st century anatomists. At approximately the same time, the British naturalist George Perry (b. 1771) published a similar book on Conchology, _][_or the natural history of shells in 1811. In reviewing this book, Petit (2003) concluded that most of its illustrations are of good quality whereas others show shells that are poorly drawn and coloured garishly. The name(s) of the artist(s) for the Conchology _]is (are) not stated but a reading of the introduction lead Petit to the conclusion that they were probably mostly undertaken by Perry, who stated that ‘[_the plates are engraved, and coloured after the original drawings, by Mr. John Clarke.’
Plate 22 of Poli’s ‘Testacea utriusque Siciliae……’ (1791). This plate was painted by Franco Morelli and engraved by Antonio Zaballi.. (From the book in the library of the Natural History Museum, London). (Photo: B. Morton).
Mention was made earlier of Thomas Bewick (1753–1828) who was an English ornithologist and wood engraver. Despite having no formal art training, Bewick showed a talent for drawing at an early age and, at 14, was apprenticed to Ralph Beilby (1744–1817), an engraver in Newcastle. They became partners ten years later, although this did not last as the two men disagreed about authorship of their books. The texts in Bewick’s and Beilby’s A General History of Quadrupeds (1790) and History of British Birds:[_ Land Birds_] (1797) and Water Birds (1804) were drafted by Beilby and revised by Bewick. According to Bewick, Beilby wanted to have his name appear in Land Birds as the sole author. Upon Bewick’s disagreement, however, neither was named as the author and the partnership came to an end in 1797. They were somewhat reconciled in 1800, co-operating again for some projects, including the publication of [_Figures of British Land Birds _]published by S. Hodgson in 1800.
Bewick’s and Beilby’s A General History of Quadrupeds deals with mammals from all over the world and some domestic animals. It includes bats and seals but does not include the cetacean mammals. Bewick was helped by his intimate knowledge of the habits of animals acquired during his constant excursions into the country. He also used information passed to him by acquaintances and local gentry and that obtained in natural history works of his time including the earlier mentioned Thomas Pennant’s (1726-1798) Synopsis _]([_and later History)[_ of Quadrupeds_] published in 1771. In 1766-1767, Pennant published his British Zoology.
Pennant, however, was no artist whereas Bewick was a highly skilled engraver of harder woods, notably box, cut against the grain using fine tools normally favoured by metal engravers.[* ]Bewick was also influenced by the French naturalist[ ]Georges-Louis Leclerc, Comte de Buffon (1707–1788), an encyclopaedic author, but also no artist, his books being illustrated by others. During his lifetime, Buffon published 36 volumes of his Histoire naturelle with additional ones based on his notes and further research being published in the two decades following his death. His works influenced not just Bewick, but the next two generations of naturalists, including the equally eminent Jean-Baptiste Lamarck and Georges Cuvier. Ernst Mayr (1904-2005) said of Buffon (in 1981) that he ‘was the father of all thought in natural history in the second half of the 18th century’. Similarly,[ *]Bewick’s art is considered today the pinnacle of its [engraving] medium (Uglow 2006).
Undoubtedly, however, the pinnacle of bird illustration was reached by[* *]John James Audubon (1785–1851), a French-American ornithologist, naturalist, and painter who was actually born in Haiti. His major work, the colour-plate The Birds of America (1827–1839) cost US$115,640 (over US$2 million in today’s money) to publish, paid for from advance subscriptions, exhibitions, oil painting commissions, and the sale of animal skins, which the author had shot. To put the set in the modern context, on 20 January 2012 a complete copy of the first edition was sold, by the heirs of the Fourth Duke of Portland, at Christie’s in Manhattan, for US$7.9 million.
In the latter part of the 18th century it became fashionable for the aristocracy and people of intellect to construct cabinets in their homes, of diverse scales, and to fill them with curios from the natural world. Of particular interest were seashells. In 1815, Samuel Brookes (died 1838) published An Introduction to Conchology and in which he explains (p. vi) that the figures of the shells [in 11 plates] ‘are drawn and engraved chiefly from specimens in my own collection’ with no mention made of an artist. It seems he was drawing them himself although they were obviously hand coloured after production.
Thomas Brown (1765-1862) in his The Elements of Conchology or Natural History of Shells: according to the Linnean System, published in 1816, makes no mention of an artist but each of the nine plates has a name beneath it. Plate 9 of the book was painted by R. Scott an Edinburgh sculptor (Plate 9),[* *]possibly explaining its poor quality. In 1835, Brown published The Conchologists Text-Book embracing the arrangements of Lamarck and Linnaeus with a Glossary of Technical Terms.
Here, the plates are specifically attributed to either himself or, again, to the artist R. Scott and which show improvements over the intervening course of time. In Brown’s subsequent [Illustrations of the Land and Freshwater Conchology of Great Britain and Ireland, _]published in[ _]1845 all of the 27 plates were stated to have been specifically drawn by the author except for two of slugs, which were drawn by the Rev. B.J. Clarke. Brown’s 1833 book, although compact, perhaps for a pocket-carried and, thus, field-collecting reason, was remarkable in a number of ways because it not only described how to collect (with an annotated illustration of a dredge drawn presumably by himself) and clean shells but also how to arrange them in a cabinet. Most importantly, however, Brown was assiduous in identifying his artists, though he was clearly as equally talented as they, and was thus on a number of counts not just a shell collector but also the forerunner of modern museum curators.
Plate 9 of Thomas Brown’s ‘The elements of Conchology’ (1816). The artist was R. Cook. (Library of B. Morton).
Conversely, many of Brown’s contemporaries, including the British naturalist, William Turton (1762–1835) who, in 1831, published, the somewhat long-windedly titled A manual of the land and freshwater shells of the British Islands arranged according to the more modern systems of classification; and described from perfect specimens in the author’s cabinet: with coloured plates of every species makes no mention of an artist. Others were slightly more appreciative of the artistic endeavours of their employees, John Mawe (1766-1829) recording on page xii of his 1823 book The Linnean System of Conchology that ‘[The plates which embellish this work are taken from specimens in our own cabinet, and we are indebted to the kindness and indefatigable exertions of an _][but still un-named][ artist of the greatest talent, for the accurate and elegant delineation of them_]. The lithography was, however, identified as being undertaken by E.A. Crouch. In an earlier, now rare, book, [The Voyager’s Companion or Shell Collector’s Pilot _]first published in[ _]1821, but going into four editions with only one copy each of the first two editions surviving, Mawe reproduced colour plates with no mention of an artist but they were probably by his own hand. Importantly, however, after (reportedly) sailing the globe collecting mostly shells, the author describes how they (and other plants and animals) should be prepared, preserved and packed, even advising on how to get them through customs undamaged. This was, therefore, the world’s first, global, shell-collecting guide. This book was, however, a subsequent edition of his earliest un-illustrated [_A Short History on Natural History _]published in 1804. This perhaps nicely demonstrates how slowly it was that illustrations came to be used in published books, that is, no more than two hundred years ago.
In 1820, John Wodarch published an[_ Introduction to the study of Conchology_]. This was subsequently reproduced, first in 1822, but again and again, until 1932, by John Mawe as Wodarch’s Introduction to the study of Conchology. The last edition of 1832 was published posthumously by Mawe’s wife, Sarah, three years after his death – whether out of love or money (or both) is unknown. The coloured plates in this book were executed by Edward Crouch. William Pinnock (1782-1843) published[_ Pinnock’s Catechism of Conchology_] in London in 1824 (2nd edition in 1829) and it is, at 8.8 × 13.7 centimetres, again perhaps a pocketbook, the smallest text ever written about shells but contains fine plates by the artist T. Bradley.
Lovell Augustus Reeve (1814–1865) was an English conchologist and publisher. The chance purchase of some shells as a boy led to a lifelong interest in conchology and, eventually, publication of his Conchologia Systematica in London (1841–2). His work culminated, however, in his Conchologia iconica, or, Illustrations of the shells of molluscous animals, which spanned 20 volumes and contains about 27,000 figures and was published by his own company Reeve’s Brothers in London between 1843 and 1878. The colour plates in these volumes are exquisite.
One of the most famous 18th and 19th century conchologists, however, was George Brettingham Sowerby I (1788–1854) who was the second son of James Sowerby. Together with his brother James De Carle Sowerby he continued their father’s work on shells, publishing his most important work – Thesaurus Conchyliorum, that was continued by his son and grandson, George Brettingham Sowerby II and III, respectively. One of his first works was cataloguing the shell collection and cabinets of the Earl of Tankerville (1743–1822). He also published[_ A Conchological Manual ]that ran into four editions with ‘[_All the plates…by his hand]’. Plate 10 is Plate 2, figures 51-59 in the 3rd edition of Sowerby’s Conchological Manual of 1846. G.B. Sowerby also assisted in illustrating shell books for others and William Wood (1774-1857) records in his Index Testaceologicus (1825, pp. iv-v:) ‘It may be not improper to remark that all the plates [38 + 2,300 figures] (with exception of the first six by that excellent artist the late Mr. Sowerby) have been executed under the immediate inspection of the Author’. Slightly patronising of Sowerby one feels but, nevertheless, he probably only cared that he got paid for doing them.
Petit (2003) points out that many figures in the old works were copied from those of others and that such a practice was common. In the 18th but more so the 19th centuries, engravers and lithographers were so skilled they could copy figures, almost like photocopying and, sometimes, plates were re-arranged and some figures were not copied, but redrawn from different specimens. Similarly, even with the famous engravers, such as the generations of Sowerby’s, plates were often re-engraved several times. Petit (2009) shows that many Sowerby plates were re-lithographed at various times and that hand-coloured copies of some monographs were, surprisingly, still being produced into the 1950’s. An example, it seems, of the old adage that ‘there is nothing new under the sun’ (Ecclesiastes 1:4-11).
In France, Jean Claude Chenu (1808-1879) published his beautifully illustrated four volume Illustrations Conchyliologique ou description et figures de toutes les coquilles connues vivantes et fossiles, classées suivant le système de Lamarck modifié d’après les progrès de la science et comprenant les genres nouveaux et les espèces récemment découvertes in Paris between 1842–1854.
Plate 2, figures 51-59 of G.B. Sowerby’s ‘Conchological Manual’. 3rd edition. (1846). The artist was the author. (Library of B. Morton).
Jabez Hogg (1817–1899) was Consulting Surgeon to the Royal Westminster Ophthalmic Hospital and who, in 1854, with many other editions printed subsequently, published The Microscope: its History, Construction and Application. Hogg’s book was illustrated profusely with eight colour plates by Edmund Evans and 354 numbered figures. Others were engraved by a Mr George Pearson. I use this book only to demonstrate that by the middle of the 19th century illustrators had, well and truly, come of age. And, far from drawing shells and other cabinet-held creatures for the rich, famous and otherwise elite, were now actively engaged in the development of science in all its varied aspects for a much-wider audience.
Walter Hood Fitch (1817-1892) was, historically, one of the most prolific and remarkable botanical British artists who published over 12,000 drawings of plants and their flowers especially for Curtis’s Botanical Magazine (Lewis 1991). For this magazine alone Fitch drew over 2,700 plates between 1834 and 1877. He also painted some 500 plates for the majestic Icones Plantarum (1836-76) produced by Sir William Jackson Hooker (1785-1865). They are, however, as were the early molluscan tomes, works of artistry produced for the gentry whereas a later illustrator, William Keble Martin (1877-1969) painted for a broader public. The Rev. Keble Martin was a British botanist and botanical illustrator, best known for his Concise British Flora in Colour published in May 1965 when the author was 88. The book was the result of sixty years’ meticulous fieldwork and highly developed painting skills, and became an immediate best-seller. He completed over 1,400 paintings in colour and many black and white drawings before the book was finally published. Today, however, it is the first book I turn to when trying to identify my own British coastal plants.
One of the British societies responsible for publishing and sometimes re-publishing historical and beautifully illustrated natural history texts is the Ray Society founded in 1844 to honour the name of John Ray (1628-1703), one of the most eminent and influential naturalists of his time. Up until the Society’s 150th Anniversary in 1994, it had published 163 such volumes and its work continues today.
Between 1845 and 1855, the Ray Society published an eight-part portfolio of illustrations and descriptions of the British nudibranch molluscs by Joshua Alder (1792-1867) and Albany Hancock (1806-1873). A ninth part by Sir Charles Norton Edgecumbe Eliot (1862-1931) was published in 1910. Eliot was a British diplomat, colonial administrator, academic, historian, linguist (being fluent in sixteen languages and conversant in twenty more), botanist and zoologist with a particular interest in sea slugs, and, probably, a spy. Interestingly, Eliot was appointed the founding Vice-Chancellor of the University of Hong Kong – my near career-long domicile. His appointment to this position must have been due not only to his administrative abilities, modesty and self-effacing charm, but principally his highly-valued linguistic skills because he quickly became fluent in Putonghua (and almost certainly Cantonese as well). He served as Vice-Chancellor until July 1918 when the British Government sent him to Russia to investigate the death of the Romanov Royal Family and report back on what had happened. Eliot concluded that Tsar Nicholas II and his family, their doctor, two servants and the maid had been shot in Yekaterinburg’s Ipatieve House by the Bolsheviks. After Hong Kong, Eliot became British Ambassador to Japan in 1919 but died in 1925 on his way back to Great Britain into retirement and was, fittingly, buried at sea.
Joshua Alder was a British zoologist and a malacologist, but he also specialised in the Tunicata, and the Gastropoda – especially sea slugs. Hancock was born in Newcastle-upon-Tyne and became an eminent naturalist, biologist and supporter of Charles Darwin (1809-1882). His brother, John, was also a naturalist and the Hancock Museum in Newcastle-upon-Tyne is named after them, both playing an instrumental part in getting the museum built initially.
One could be forgiven for assuming that either Alder or Hancock (or both) were the artists responsible for the beautiful prints that make up the nudibranch portfolio. Not so: the Advertisement to part I of the portfolio set reads ‘[The plates are accurate transcripts of their _](Alder and or Hancock)[ original drawings. They have been executed by Mrs Holmes, an accomplished artist in Lithotint, an invention of Mr. Hullmandel, admirably adapted for pourtraying ][sic][ the delicacy and beauty of these fragile inhabitants of the sea._]’
There is no doubt, however, that Alder was himself an illustrator because in the preface to the four volume A history of British Mollusca and their shells published in 1853 by Edward Forbes (1809–1885) and Sylvanus Hanley, Alder was acknowledged as ‘[gifted with the power of delineating, at once accurately and artistically, the animals whose external clothing and internal structure had alike engaged his attention….’ _]and[ ‘the pencil of Mr Alder has, in many instances, embellished our illustrations with figures exceedingly precious’.] Plate 11[* ]is[ ]Plate 33 of[ *]Alder & Hancock’s four volume portfolio (1844-1848) and Part III shows _Eolis picta (Eubranchus pallidus) – the artist, as noted above, identified only as Mrs Holmes.
Plate 33. Eolis picta (Eubranchus pallidus) (Alder and Hancock, 1842) from Alder and Hancock 1844-1848. Part III. The artist was identified only as Mrs Holmes. (Reproduced with the permission of the Council of the Ray Society).
Forbes was a British conchologist and malacologist and was, himself, a gifted illustrator and contributed his own illustrations to the work as did a certain Mr Clark. Forbes was in other ways also a remarkable man and, in 1848, the Ray Society published his monograph on the British naked-eyed Medusae. His History of British Starfishes and other animals of the class Echinodermata (1841) was embellished not only with his own drawings of various species, but also with an assortment of tiny and charming vignettes, which were stuck in wherever there was an open space at the end of a section of chapter. Subsequently, Forbes became curator at the Museum of the Geological Society of London (1842), Professor of Botany at King’s College, London (1842), and paleontologist to the British Geological Survey (1844). Forbes is most famous for proposing, in 1846, that a great land mass had existed in the Miocene encompassing northern Europe and Spain, and extending out from the Mediterranean far westwards into the Atlantic Ocean virtually to the coast of North America. On his return to England, from the voyage of H.M.S.[_ Beagle_], Charles Darwin became sceptical of Forbes’s lost land and set up his own experiments in the glasshouse at Down House where he immersed the seeds of 87 species of common plants in seawater for a month. He then tried to germinate them and found that over half (64) had survived. By his own calculations, ocean currents could thus have taken such seeds well over half way across the Atlantic Ocean. He undertook similar work on dried muds collected from the feet of migrating birds and concluded that no Forbesian landmass was necessary to explain Hooker’s biogeographic similarities in the flora’s of Europe and North America. Not true, however, but then Darwin was unaware of the future science of plate tectonics and continental drift.
Famously, the Ray Society also published Charles Darwin’s two volume monograph on the barnacles (Cirripedia) (1851 and 1854) although it is worth noting that the illustrations for the volumes were by George Brettingham Sowerby. Similarly, the Ray Society published the four volume, in nine parts, monograph by W.C. Macintosh of the British Marine Polychaetes (1873, 1874, 1900, 1908, 1910, 1915a, b, 1922, 1923) (not referenced). To complete this monumental, even monolithic, task, it was said that his wife did the drawings for it in their home’s attic. Other volumes published by the Society were N.J. Berrill’s The Tunicata in 1950 and which describes the 75 sessile and pelagic species that occur in British waters. Over 60 years after its publication, The Tunicata remains one of the most important works on the Ascidiacea. In passing, Berrill’s book, The Living Tide (1951) was singled out by Rachel Carson (1907-1964), author of The Sea Around Us (1951) and Silent Spring (1962), as one of the best books of 1951. Also published by the Ray Society in 1951 was the [_British Mysidacea. _]This was the work, principally, of W.M. Tattersall (1882-1943), Professor of Zoology at the University of Cardiff, although it was put together posthumously by his wife Olive S. Tattersall (1890-1978). Kemp (1980) records in a paper about the Tattersall’s lives, that it was Olive who made the meticulous illustrations for the some twenty described species.
Finally, but by no means the sum total of the works on marine plants and animals produced by it and not all identified herein, the Ray Society published the two-volume monograph of British Sea Anemones (1928 and 1935) by T.A Stephenson. Thomas Alan[* *]Stephenson (1898–1961) was a British marine biologist with a fascination for sea anemones. The books contain over 100 text figures and 33 plates including exquisite colour illustrations of the species, all undertaken by the author plus discrete thumbnail sketches of his naked wife as a mermaid nymph. These wonderful illustrations by T.A. Stephenson make him, as a direct contemporary of R.D. Purchon and C.M. Yonge, my two mentors, the third in the triumvirate of British marine biologists who constitute my 20th century biological heroes.
Stephenson was a remarkable man in many ways. He enrolled at University College, Aberystwyth, Wales, to study biology but had to abandon his studies because of ill health. Despite not completing his degree, he was made a staff member and was later awarded a doctorate for the body of research work that he had produced. Subsequently, Stephenson held a number of academic posts in Britain and at the University of Cape Town, South Africa, eventually returning to his alma mater in Wales as Professor and Head of the Department of Zoology at Aberystwyth. The National Marine Biological Library at the Marine Biological Association of the United Kingdom’s laboratory in Plymouth holds some of his paintings, but I own one of them (Plate 12) undertaken while he was a member of the Great Barrier Reef Expedition of 1928-1929 and of which C.M. Yonge was the leader. Stephenson was also the lead co-author of virtually every coastal ecologist’s bible – his and Anne’s, again pioneering Life Between Tide Marks on Rocky Shores (Stephenson and Stephenson 1972). Not just this but his papers on coral reef structures on the Great Barrier Reef are similarly significant, including the paper describing his ecological surveys of coral reefs, undertaken in co-operation with another contemporary British marine biological luminary, Sidnie Milana Manton (1902–1979), an authority on the Crustacea (Manton and Stephenson 1935).
Staying with the sea, Gunnar (Axel Wright) Thorson (1906–1971) was a Danish marine zoologist and ecologist, who, in 1957, was appointed Professor of Marine Biology at the University of Copenhagen. Thorson studied planktonic larvae of marine benthic invertebrates (Thorson 1946). He conceived the idea that in the tropics, benthic animals tend to produce large numbers of eggs developing into pelagic and widely-dispersing larvae, whereas at higher latitudes they tend to produce fewer and larger eggs and offspring. This idea was later termed Thorson’s Rule (Thorson 1957). Thorson founded the Marine Biological Laboratory of the University of Copenhagen and was a professor there from 1958-1968. Every year, Thorson sent a Christmas card (Plate 13) to his many colleagues and in these signed illustrations of North Atlantic marine life we can identify his artist – Poul H. Winther (1898–unknown).
Painting by T.A. Stephenson of ‘coral and Fungia’ undertaken on the Great Barrier Reef (1928-1929) of Australia. (Original in the collection of B. Morton).
Gunnar Thorson’s Christmas card for 1955. It shows the whelk Neptunea despecta (Linnaeus, 1758) with egg capsules and hatching embryos. The artist was Poul H. Winther.
I conclude this introduction by discussing Ernst (Heinrich Philipp August) Haeckel (1834–1919), an eminent German biologist, naturalist, philosopher, physician, professor, artist and illustrator who, between 1859 to 1887, discovered, described, and named thousands of new species. Haeckel first studied medicine in Berlin and Würzburg and, indeed, attained a doctorate in medicine, and had a license to practice. He did not enjoy treating suffering patients, however, and left medicine to study zoology at the University of Jena. After three years he earned a doctorate and became Professor of Comparative Anatomy at the University of Jena, where he remained for 47 years, from 1862 to 1909. Between 1859 and 1866, Haeckel worked on many invertebrate groups, including radiolarians, poriferans and annelids and during a single trip to the Mediterranean he named nearly 150 new species of radiolarians.
Haeckel advanced a version of the earlier recapitulation theory, previously set out by the French physician and embryologist Étienne Serres (1786-1868) in the 1820s which proposed a link between ontogeny (development of form) and phylogeny (evolutionary descent), summed up by Haeckel in the phrase ‘ontogeny recapitulates phylogeny’. The theory has since been discredited in its original form and Haeckel’s embryological drawings, researches and conclusions derived therein have also become controversial. For example, his illustrations of the embryos of a dog, chick and a turtle are identical and presented to the reader three times in a row and with three different captions: a perfect case of scientific facts being distorted to fit a pre-conceived concept. Nevertheless, his fertile imagination and considerable artistic talents led to the production of some of the most beautiful illustrations of marine organisms, ever made. The artwork of Haeckel includes over 100 detailed, multi-colour illustrations of sea creatures and which were re-published in Kunstformen der Natur (Art Forms in Nature) (Haeckel 1998).
Where illustration becomes art. Ernst Haeckel’s illustration of four species of Discomedusae. (From: Ernst Haeckel, ‘Art forms in Nature’, Tafel 8, 1998).
Haeckel’s life is documented by Richards (2008). His story was tainted in his later years by objections to his science but also accusations of fraud. His early, signed, illustrations of radiolarians are undoubtedly by his own hand although the more magnificent illustrations in his many treatises are unsigned and undoubtedly have to be attributed also to his lithographer Adolf Giltsch who translated them from sketch to print.
To conclude, there is, I discover, a silken thread linking, through time, the author-illustrators commencing in the earliest moments of modern human history, some 40,000 years ago finding sublime scientific expression in the works and illustrations of Martin Lister (1639–1712), the artists hired by Giuseppe Saverio Poli (1746-1825), Thomas (1753–1828) and George Brettingham Sowerby I (1788–1854), to name only a few, but culminating in the works of T.A. Stephenson and, especially, of Ernst Heinrich Philipp August Haeckel (1834–1919) all of whom turned biological illustration into an art form.
This portfolio comprises 80 illustrations of mostly marine but also some estuarine, coastal and one or two freshwater plants and animals, all drawn by myself. Each one is accompanied by a vignette that identifies and describes points of interest about the species and which first inspired me to draw them in the first place. I also provide a few references for each vignette, so that the more interested reader can find his or her own way into the appropriate literature. In compiling and publishing this portfolio and accompanying vignettes, I make no artistic claims, only illustrative and interpretive ones – both of which are necessary pre-requisites for descriptive biologists. I do, however, hope that something of what is illustrated herein will stimulate others to take up pencil and pen because, today, fewer and fewer people, even professional biologists, are attempting to enlighten their readers in such a way and have also lost the satisfaction and fulfilment of doing something using their own hands and eyes.
The freshwater zebra mussel, [_Dreissena polymorpha _](Pallas, 1771), is native to the Black, Caspian, and Azov Seas and was first described from the Caspian Sea and Ural River. It was introduced into the British Isles in the early 19th century and first made its appearance in the London Docks in 1824. It had probably arrived aboard a cargo of timber from Europe. Although its origins were in the Caspian Sea, the species spread throughout Europe and into Great Britain along the growing canal network that was being built to transport the products of the Industrial Revolution. In Great Britain it similarly spread along the canal network this greatly facilitated by the fact that being byssally-attached, individuals simply settled onto the hulls of barges and were thence transported virtually and literally everywhere (Kerney and Morton 1970). Today, the species occurs in the north in Denmark, Sweden and Finland, down through Holland and as far east as Italy. It is a significant pest in Russia too.
The series of papers published on Dreissena polymorpha (Morton 1969a, b, c, d), collected from Reservoir No. 2 in the Lee Valley Reservoir complex at Walthamstow, North London, as a consequence of my Ph.D. research, received a muted response until about 1988 when the bivalve[_ ]was introduced into the U.S.A., notably the Great Lakes of North America. The impacts of this animal were devastating and a brief moment of malacological fame was enjoyed – my papers being the only ones hitherto published on its anatomy, population dynamics and feeding. My first ever drawings of the shell and siphons of _Dreissena polymorpha are illustrated here (A-E).
Because Dreissena polymorpha was introduced into Great Britain during the 19th century, the companies that supply us with potable water have grown up with the pest and it is largely controlled and prevented from causing too many problems. A different scenario was enacted after it was introduced into North America. There, it shut down power stations, including nuclear ones, by blocking the cooling water systems but, most incredibly, by virtue of the massive filtering capacity of the untold numbers of individuals present within them, has turned the Great Lakes from planktonic- to epibenthic-based communities and hence from eutrophic to oligotrophic ecosystems. This one introduced, invasive, species has thus dramatically altered the ecologies of the water bodies that constitute one fifth of our global reservoir of freshwater. Since 1988, there has been an annual conference and exhibition convened in the U.S.A. and Canada to provide the scientists and managers with the latest details of its biology and spread and the newest technological advances needed to control it. Today, in 2013, the species has spread throughout much of the U.S.A., but is especially prevalent in the northeast and the Mississippi drainage, but has spread to as far west as California and is continuing its march wherever it can find a toehold.
In this North American introduction, Dreissena polymorpha was accompanied somewhat later, in 1991, by a second dreissenid, the quagga mussel, Dreissena bugensis Andrusov, 1897, another Ponto-Caspian species, that was also first recorded in North America from the Great Lakes. It too was probably introduced in discharged ballast water from trans-Atlantic shipping sources. Since that time, it has spread to numerous scattered locations throughout the U.S.A. as far west as California. The two species of Dreissenidae are pernicious pests of natural and potable water supply systems throughout much of North America and Europe as far east as Russia. They cost their recipient countries billions of dollars each year to control and the amount of research that is conducted on them is just enormous.
In early 1975, Winston Ponder, Curator of Mollusca at the Australian Museum in Sydney, Australia, organised a research meeting on Lizard Island on the Great Barrier Reef, to celebrate the opening and completion of his institution’s marine laboratory. Only about a dozen people were to attend, but they included Sir Maurice Yonge (1899-1986) who was especially invited not only to celebrate his 75th birthday but also because he had been the leader of the Great Barrier Reef Expedition of 1928-1929 and so was responsible for initiating research on the reef. It would be an opportunity for me to work alongside Great Britain’s most famous marine biologist and, so, in December of 1975 I arrived on Lizard Island from Hong Kong, [_via _]Cooktown where I became the sole passenger on a single-engined Cessna that would fly me out to the island.
It was an amazing experience to fly low over the reef with its myriad islands and half exposed coral reefs but was soon landing on an airstrip on Lizard Island. The pilot pointed me towards a track that would lead to the laboratory and I set off, rucksack on my back. I remember thinking ‘I wonder why this is called Lizard Island?’ when out of the track’s bush verge stepped an enormous, two metre long, lizard – the perentie (Varanus giganteus [Gray, 1845]). I stopped dead in my tracks, but it just gave me a glance and strolled off into the bush on the other side of the path.
Although built in 1973, the marine laboratory had been clearly little used and participants in the meeting were sleeping in bunks in wooden shacks and digging one’s own latrines as and when the need arose. There was a couple of resident scientists who also dived, serviced the compressors and boats and generally helped facilitate research. The lizards, quite a lot of them, became attracted to the camp and were a nuisance at mealtimes when they foraged for scraps and became so bold as try to eat our food off the tables and even open refrigerator doors. One became quite blasé about them eventually and shooed them away despite their warning hisses. Then, we did not realise that their bite was venomous.
Gradually, the participants settled down to work and I found two fascinating bivalves to research – the coral-sand dwelling Fimbria fimbriata (Linnaeus, 1758) and one of the smallest of the giant clams – Tridacna crocea (Lamarck, 1819) – the latter boring into old coral rocks. These researches were eventually published (Morton 1978, 1979a). One of the pleasures of the meeting was, however, that in addition to undertaking one’s own research, we were able to examine and discuss the other animals the divers brought back alive to the laboratory and one of these was a cuttlefish[. _]Clyde Roper from the Smithsonian Institution in Washington D.C., was studying cephalopods – the octopuses, squids and cuttlefishes – and one species that came back and was kept alive in an aquarium was the flamboyant little cuttlefish[ Metasepia pfefferi ](Hoyle, 1885).[* *]This little cuttlefish, no more than between 2-3 centimetres long, lives in the shallow waters of the tropical Indo West Pacific and ambles (there is no better word to describe it) along the sea bed of sand and coral rubble hunting for its prey – typically small shrimps. But the most remarkable thing about it is its coloration. Normally camouflaged, when disturbed _Metasepia pfefferi can display the most remarkable colours. Typically, the body takes on a dark purple background broken up with bars of white and flashes of yellow marginally and on its arms and head. There may be patches of bright red too on the tips of the arms. Such a striking colour pattern is clearly a warning signal and recent research[_ ]has shown that the species[ _]has a venomous bite like the notorious blue-ringed octopus.
Though a friend, Clyde Roper, who was working on the animal, was rather sniffy about the drawing I made of the little [_Metasepia pfefferi _]but many years later asked if he could use it for an illustration of the species in a paper he was writing about colour patterns and behaviour in cephalopods from the Great Barrier Reef (Roper and Hochberg 1988).
One of the finest beaches in Hong Kong for intertidal life is Starfish Bay or, in Cantonese, Hoi Sing (Starfish) Wan (Bay). In the early 1970’s it was near-pristine and it derived its name from the fact that the beach was literally covered by huge numbers of a seastar -[_ Archaster typicus_] Müller et Troschel, 1840. Here, the seastar formed an interlocking mesh of arms, lightly dusted by a layer of sand, that covered virtually all the lower intertidal. Archaster typicus is usually five-limbed with long, slightly tapering, arms with pointed tips. Adults grow to between 12-15 centimetres in diameter, with males often being smaller than females. Each individual lies buried in the sediment during high tides and moves over the sand surface to feed during low tides. The upper, aboral, side is typically grey-brown and variously marked with darker and lighter patches. The oral side is pale and demarcated from the aboral side by a marginal fringe of short, flat and blunt spines.[_ _]
This seastar has a very wide Indo-West Pacific distribution from the Bay of Bengal in the west out into the Pacific as far as Hawaii in the east and from Japan in the north to Australia in the south. Archaster typicus is a detritivore, using its everted yellow membranous stomach to seive the sand. The engulfed detritus is brought inside the seastar when its stomach is inverted to its normal position.
Eventually (Morton 1979), a year was spent working out the life cycle of the animal. Individuals reach sexual maturity at a radius of three centimetres. About two months ahead of spawning, in Hong Kong’s spring, the seastars begin to congregate – males and females, probably recognising each other by chemotactic contact. On recognising a female, a male climbs on top of her and may remain there for two months during the summer season, each of their five arms interdigitated. During this time their gonadal cycles are synchronised, so that spawning, which occurs in late summer, is too. This is called pseudocopulation since there is no penis or vagina. Fertilisation is by release of sperm and eggs into the sea for external union. Nevertheless, the mass of pseudocopulating seastars is a strange sight to see. Larval settlement is said to occur among mangroves, individuals gradually moving seaward to sandy habitats as they age.
It was also quickly noticed that virtually all the seastars were parasitised by two species of small, apically slightly curved, gastropods – Mucronalia fulvescens (A. Adams, 1860)[_ ]and _Balcis shaplandi Melvill, 1910. Both are species of the Eulimidae, all the representatives of which are parasites, mostly of echinoderms The lighter, creamy yellow,[_ Mulcronalia fulvescens_] lived in the lower (oral) interstices of the seastar’s arms and fed by sticking its proboscis, armed with penetrating radula teeth, into the host’s tube feet and sucking out the fluids of the water vascular system. Conversely, the darker, grey-green, [Balcis shaplandi _]lived on the[ _]upper (aboral) surface of the seastar and eroded away a circular patch of epidermis to facilitate insertion of its proboscis into the host’s coelom and suck out those fluids (Morton 1976). Effectively, therefore the two parasitic snails were not only sympatric on their seastar host but had also partitioned and were exploiting two different food resources.
Over the 1980’s, however, it became apparent that the numbers of seastars on Hoi Sing Wan were declining as were their parasite numbers. And in the winter of 1989-1990 they disappeared from the beach. This was attributed to pollution and I wrote an article in Hong Kong’s main English language newspaper, the South China Morning Post, about this disappearance, ending with a caustic comment that Starfish Bay should now be re-named. Today, the species is rarely seen on Hong Kong beaches, except for a few survivors on the sandy beach in the marine park of Hoi Ha Wan.
Two small sea grass species, Halophila ovata Gaudich and Halophila beccarii (Aschers.), characterise Hong Kong’s muddy intertidal flats. They occur low down on sandy flats well below the fringe of pioneer mangrove plants that also characterise such shores. Similarly, such beaches are, landward, the abode typically of the pioneer mangroves, [Kandelia candel _](Linnaeus) Druce,[ Avicennia marina_] (Forsk.) Vierh. and [Aegiceras corniculatum _](Linnaeus) Blanco, the backshore mangroves, [_Bruguiera gymnorrhiza _](Linnaeus) Savigny,[ Excoecaria agallocha_] Linnaeus and [Lumnitzera racemosa _]Willd., which[ ]grade into scrub and, typically, then, paddi-fields and vegetable plots. Such plants would once, however, have effected the transition to forest, prior to human habitation, but this has now mostly been felled. Since Nature hates emptiness, some plant species should occupy the intertidal space between mangrove-fringed land and seaward _Halophila.
It is not well known outside Hong Kong, but some 40% of this Special Administrative Region (SAR) of China comprises country parks. These mainly encompass the higher hills above the rainwater catchments but, nevertheless, the parks constitute a vital source of recreation for Hong Kong’s largely urban and overcrowded population. One weekend in the summer of 1977, I climbed up to and walked along the Dragon’s Back in the north-eastern quadrant of Hong Kong’s New Territories from Fanling, along the ridge of the line of nine peaks and then down to the beautiful coastline of Crooked Harbour, tracking the shoreline by means of the paths that connect all such coastal villages in Hong Kong’s New Territories. These were built by the Kadoorie Agricultural Aid Association (KAAA) to allow villagers to interact and, subsequently, are now mainly used by country park hikers.
At the village of Lai Chi Wo, virtually abandoned apart from a few elderly residents, I noticed that the mangroves were fringed landward by a grove of huge trees. This was most unusual and they proved to be Heritiera littoralis _]Dryand., Aiton, a backshore mangrove species that would naturally effect the transition, as it was virtually doing here, to forest. Winding through the coastal fringe of mangrove plants, I emerged from the dwarf pioneers to find the mud covered by a thick mat of the seagrass [_Zostera nana[* *]Roth.[_ ](Hodgkiss and Morton 1978)[. ]This eelgrass has long, bright green, ribbon-like, leaves some 10 millimetres wide (A). The whole seagrass bed is inter-connected by an extensive network of white (no chlorophyll) branching rhizomes and from which the stems arise at periodic intervals. The plant’s flowers are enclosed in the sheaths of the leaf bases and contain both stamens and anthers ©. Eventually, tiny sigma-shaped seeds are produced that can float and thereby achieve dispersal by the plant (B). Beds of _Zostera, of which there about twelve species world-wide, are important in that they stabilise bay sediments as here at Li Chi Wo and the epiphytes on the leaves are grazed by numerous snails. Eelgrass beds are grazed by the Eurasian widgeon, Anas penelope Linnaeus, 1758, which also occurs in Hong Kong, so that the plant is sometimes called, but erroneously, widgeongrass (see the illustration of Ruppia maritima).
Looking landward from the eelgrass bed, I realised the full significance of what I had found. Not just a new record for this species in southern China, but an intact coastal mangrove system – something I had instinctively felt should be here but was now seeing for the first time. Because of its remoteness and because it has effectively been abandoned, the village of Li Chi Wo has survived and left around it, free from intensive agriculture and shore gathering, there is a coastal environment that is near natural. A complete ecosystem of former natural times grading from forest to Heritiera and another backshore mangrove, the beach bean [Canavalia maritima _](Aublet) Thouars, to back mangroves, _Lumnitzera, Excoecaria and Bruguiera, to the pioneer plants, Avicennia, Kandelia and Aegiceras, and finally to the maritime eelgrass, Zostera, with [_Halophila _]completing the picture of plant succession at the low tide mark.
Today, the Agriculture, Fisheries and Conservation Department of the Hong Kong SAR Government has designated the area around Li Chi Wo as a marine park, which has achieved, hopefully permanent, protection for the unique mangrove ecosystem here because it is united with the Plover Cove (East) Country Park (Morton 2003).
[S _][_pondervilia bisculpta _](Gould, 1861)[ ]is a tiny bivalve, no more than a few millimetres in length, and could only be found by sieving for it in Starfish Bay in the New Territories of Hong Kong, at this time in the 1970’s. Lightly coloured it is patterned anteriorly and posteriorly and antero-ventrally by little streaks and flashes of brown and yellow. It lies vertically in the sediment extending its siphons vertically to the sand-water interface above. The thinner exhalant siphon is ringed and it seems likely that it can autotomise these segments if it is nibbled by little gobioid fishes, such as the local _Bathygobius hongkongensis Lam, 1986 for example, that shares this miniature world. The collected specimens of Spondervilia bisculpta were sent to the eminent Japanese malacologist Tadashige Habe (1906-2001) and he confirmed its identity. But I noticed that the generic name Spondervilia, erected by Iredale 1930, used to have a different one, that is, [_Ervilia _]erected by Turton in 1822. The equally eminent Australian malacologist, Tom Iredale (1880–1972), was an English-born malacologist (and ornithologist) who lived for most of his life in Australia becoming Curator of Mollusca in the Australian Museum, Sydney. He was an autodidact who never went to university and lacked any formal training. This was reflected in his later work because he never revised his manuscripts and never used a typewriter. Iredale was a notorious generic ‘splitter’ believing, essentially, that everything that occurred in Australia was different and so warranted a new, different, generic name. Japanese scientists too were/are similarly serious ‘splitters’, like Iredale believing that anything that lives in Japan must be different and hence deserving of a new, unique, name. Anyway, because of this problem, the specimens of [_Spondervilia bisculpta _]were never investigated further, but I kept the drawings I made of it.
Now we jump forward a decade and in 1988, on a research visit to the Açores, another species of Ervilia, that is, Turton’s type species for the genus [Ervilia castanea _](Turton, 1822) was collected. In 1990, a research paper on this published (Morton 1990a) showing it to be a similarly tiny bivalve that in the locality of the oceanic Açorean Archipelago, occurs in such huge numbers that its washed up empty shells can stain the intertidal sand red. Quite unlike the rare[ Spondervilia bisculpta ]in Hong Kong. This study, however, showed that the genus _Ervilia was placed in its own sub-family the Ervilinae Dall, 1895 in the Mesodesmatidae Gray, 1840. It was quickly realised this was incorrect and, in that same year of 1990, another paper was published with Paul Scott, a colleague from the Santa Barbara Museum, in California, showing that [_Ervilia castanea _]should, on the basis of anatomical features and differences, be re-located in the Semelidae Stoliczka, 1876 – a family with a quite different evolutionary history from the Mesodesmatidae (Morton and Scott 1990).
Although dealing here with two tiny, possibly insignificant, little bivalves, one from the Pacific, the other the Atlantic, they do expose the problems biologists face continually. That is, in remote locations such as the Açores, and especially where marine faunas are little understood, as in Hong Kong, the simple tasks of giving animals (and plants) names can be an horrendously difficult task. And that has to be undertaken before one can even begin to study them as living, sentient, beings. Sometimes, in Hong Kong, other tasks of Sisyphean proportions, similar to the Spondervilia/Ervilia question, were encountered, and despair at not finding other ways of pushing the Greek rock up the hill became a common ailment.
In December 1981, a short research trip was made to Hawaii. The visit to the University of Hawaii was co-ordinated by a colleague, E. Alison Kay, then a Chair Professor at the University. Through Alison, I was introduced to an amateur shell collector Tom Burch who had a small vessel, [Janthina VII _](but wondered what had happened to[ Janthina I-VI_]) equipped with a box dredge. We set off one day, and I had worried needlessly, Tom was an exceedingly competent skipper, and sure enough, we collected the species that was of particular interest for research purposes.
Having obtained and partially examined the animal I had wanted, the remaining days were spent pottering on the rocks and on one good low tide found, in rock pools with sandy bottoms, two individuals of [Julia exquisita _](Gould, 1862)[ ]crawling around on a mat of shallow subtidal algae made up notably, among many others, of species of[ Laurencia ]and _Gracilaria. Never having seen such a strange bivalved-gastropod before, I decided to draw it, especially as early malacologists, mistakenly believed that it and its cousins were a link between the Bivalvia and the Gastropoda, showing how the former had evolved from the latter.
Although tiny, no more than six millimetres long,[_ Julia exquisita _]is a very interesting animal. It is a sea slug, belonging to the gastropod order Sacoglossa, all representatives of which have a radula that resembles scissors in their mouths with a single row of dagger-like teeth and which they use to pierce the cells of algae and suck out the cell sap. As a consequence, [_Julia exquisita _]is coloured a brilliant green, even the shell, with patches of brown ringed in white. The head bears a pair of long rhinophores at the base of which are tiny eyes. At first, I thought that a pair of white patches, with central black dots, further back were the eyes but they were not – probably representing false eyes for the purpose of alarming potential, but also tiny, predators. But the most remarkable thing about [_Julia exquisita _]is that it bears on its back not a typical gastropod shell but one comprising two separate hinged valves. That is, the shell divided from front to back along the dorsal mid-line so that to all intents and purposes it looks like a bivalve mollusc and not a gastropod – hence the strange colloquial name.
The species was first described by A.A. Gould (1862) from shells collected in Hawaii during the United States Exploring Expedition of 1838-1842. Naturally, for that time, Gould believed the shells to be those of a bivalve. It was not for another 100 years that Myra Keen (1960) suggested that when living individuals of Julia exquisita _]were eventually found they would turn out to be gastropods not bivalves. And so it transpired, because Alison Kay did find the living animals eventually and, yes, they were gastropods not bivalves (Kay, 1962). Alison found the animals ‘[_in sand patches on small rocks about one metre under water’ near Koloa, Kauai, in Hawaii. The rocks were also covered with algae, which they were presumably feeding upon. In addition to her work on micro-molluscs, Alison Kay (1928-2008), served as editor of the journal Pacific Science from 1972 to 2000 and was a dedicated conservationist, helping for example, to found the Save Diamond Head Association and to protect Hawaiian limpets (locally called opihi) from over-exploitation by shaping state regulations to limit their collection. Another colleague, Kathe R. Jensen from the University of Copenhagen, Denmark, has made a life-long study of the Sacoglossa (Jensen 2007).
Hemifusus ternatanus (Gmelin, 1791) is a member of the Melongenidae, the crown conches and their allies. It lives at depths of between 20-160 metres, on the sandy-mud sea-bed, that is, the continental shelf of the Indo-West Pacific region, from Japan in the north to the Philippines in the south. It can reach an impressive shell length of between 70-300 millimetres. It shares this habitat with its congener, Hemifusus tuba (Gmelin, 1791), which reaches a shell length of half that of Hemifusus ternatanus. The Hemifusus ternatanus shell is ponderously slender, coronated, unequally fusiform, with a spire that is shorter than the long aperture and siphonal canal, but shows geographical variations in sculpture and colour. The shell colour is usually light brown, while the aperture is creamy-white. [_Hemifusus tuba _]is similarly coloured leading sometimes to confusion between the two when small.
The two species of Hemifusus are regularly seen for sale in Hong Kong’s sea food markets, Hemifusus ternatanus more rarely than Hemifusus tuba. It was soon discovered that virtually nothing is known about any aspect of the biology of either melongenid and in the 1980’s a long-term study was made of them in the marine aquarium of the (then) Department of Zoology of the University of Hong Kong. Because Hemifus tuba was more readily available in Hong Kong, this species became the principal focus of the main study of melongenid biology. The species could be obtained in large numbers, readily acclimated to aquarium life and fed happily. Moreover, the species reproduced readily in captivity, males and females copulating with the latter subsequently laying large numbers of egg capsules. From these hatched, eventually, crawl-away juveniles and which, initially, fed on organic detritus, grew, allowing for estimates of consumption and growth rate to be obtained. Over the course of three years, these various aspects of the biology of Hemifusus tuba were detailed and the results published (Morton 1985, 1986a, 1987). The principal aim of this large study was to provide the information necessary for the eventual cultivation of the species because it is a highly-valued fishery resource in Asian waters.
It was discovered that in captivity, [Hemifusus tuba _]was principally a predator of bivalve molluscs, each prey item being grasped delicately by the muscular foot and the similarly stout proboscis thrust between the shell valves of selected prey individual when it relaxed and the tissues rasped out by the blade-like stenoglossate radula in the mouth. [_Hemifusus tuba _]would, however, also feed on carrion, which made estimates of consumption easier to calculate. Such a lifestyle is similar to that of the Atlantic representatives of the related Buccinidae, for example[ Buccinum undatum _](Linnaeus, 1758), which also feeds on bivalves (Nielsen 1975) and carrion making it an equally important fishery resource.
[Hemifusus ternatanus _]was, however, a more difficult species to keep for study in numbers necessary for statistical analysis but two individuals were held in captivity for 13 weeks and offered a wide range of living potential prey items (Morton 1986b). They too, however, like _Hemifusus tuba, only fed on bivalves, the preferred species being fan shells, such as Pinna pectinata Linnaeus, 1767. The illustration of [Hemifusus ternatanus _]feeding on such a prey item was made in[ ]February 1985. As with _Hemifusus tuba, the prey fan shell is approached stealthily, gently eased from its sandy habitat and the proboscis thrust between the posteriorly-gaping shell valves and then Hemifusus ternatanus sets to work eating it – alive.
Working on the beaches of Princess Royal Harbour at Albany, Western Australia, from a jury-rigged laboratory, the team of accompanying divers brought in a Southern blue ringed octopus [Hapalochlaena maculosa _](Hoyle, 1883). I was allowed to draw it after its morphometric data had been recorded. There are three or four octopus species belonging to the genus _Hapalochlaena and which live in tide pools and coral reefs in the Indo-West Pacific from Japan to Australia mainly around southern New South Wales and South Australia but clearly here also in Western Australia. They are some of the world’s most dangerous marine animals and I for one had of course had heard about them and how venemous they were but I had never seen one hitherto.
It was, to me, surprisingly small for an animal with such a fearsome reputation – the body only being about 50 millimetres long and the eight arms ten millimetres. When resting, the octopus is a uniform grey to beige-yellow colour with large light brown patches or maculae – hence the specific name. The body is also wrinkled and possesses numerous raised tubercles. When alarmed, however, the colour change is amazing. The brown patches darken dramatically and vivid, irridescent, electric-blue rings appear and pulsate within the maculae. There are some fifty of these rings and the body itself becomes patterned with blue stripes. It was exceedingly dramatic bearing in mind that it uses neurally-stimulated chomatophores in its skin to effect such remarkable changes in appearance. The blue-ringed octopus spends much of its life hiding in crevices, without any form of skeleton, changing its shape easily, to squeeze into orifices much smaller than itself. Like other octopuses too, Hapalochlaena maculosa swims by expelling water from its hyponome, or respiratory funnel, in a form of jet propulsion. Also, if an arm is lost, it can be regenerated within six weeks.
The diet typically consists of small crabs, hermit crabs and shrimps, but the octopus may bite attackers, including humans, if provoked. No blue-ringed octopus antivenom is available. The blue-ringed octopus pounces on its prey from its home crevice, seizing it with its arms and pulling it towards its mouth. It uses its beak to pierce a crab’s exoskeleton, and introduces its venom. The venom paralyes the prey allowing the octopus to rip away the shell with its arms and suck out the flesh.
During mating, a male approaches a female and begins to caress her with his modified arm, the hectocotylus. If the female is receptive, the male mates by grabbing her and transferring packets of sperm, spermatophores, by inserting his hectocotylus into her mantle cavity. Fertilised blue-ringed octopus females lay only one clutch of about 50 eggs in their lifetimes. Eggs are laid and then incubated underneath the female’s arms for about six months (Stranks and Lu 1991). During this process she does not eat. After the eggs hatch, the female dies.
The octopus produces venom contains a number of chemicals, the major component of which is tetrodotoxin, a neurotoxin also found in pufferfish, that causes paralysis and respiratory arrest within minutes of a bite, leading to cardiac arrest due to a lack of oxygen. The toxin is produced by bacteria in the salivary glands of the octopus. In humans, if untreated, death usually occurs within minutes from suffocation due to lack of oxygen to the brain. However, respiratory support, together with reassurance, even if the victim appears dead, often ensures the victims will recover. It is said that, despite its small size, Hapalochlaena maculosa carries enough venom to kill 26 adult humans within minutes. Their bites are, moreover, tiny and often painless, so that most victims do not realise they have been envenomated until respiratory depression and paralysis start to set in. For this reason, my blue-ringed octopus was treated with a great deal of respect and returned to the sea subsequently.
Nellie B. Eales (1889-1989), who was editor of the [_Proceedings of the Malacological Society of London _]when this young scientist was publishing his first papers, and was kindness and tolerance personified, was very interested in the opisthobranch gastropod Aplysiidae – the sea hares. In fact, she wrote the authoritative, worldwide, account of these animals (Eales 1960).
In July 1988, a species of Aplysia was collected from a pool in the low rocky intertidal of the Açorean island of São Miguel. Using Nellie’s key, it was possible to identify it as Aplysia punctata[* *]Cuvier, 1803[_. _]This species is recorded from the North-East Atlantic Ocean, from Greenland and Norway to the Mediterranean and out into the Açores. [_Aplysia punctata _]may achieve a length of 120 millimetres and is olive green, spotted with white splashes and small, dark, spots and veining but is also a much deeper red when young. Purplish black individuals have been reported upon as well. The dark, almost bronze, chitinous, shell is located centrally, with a single gill beneath it on the right side. The body is long and narrow and the parapodia join rather high posteriorly. The head is characterised by a pair of oral tentacles in the front and another pair of rhinophores atop it (Thompson 1976).
All sea hares are gentle creatures occurring in the low intertidal and even rock pools. They typically aggregate in spring to copulate and lay long, orange-yellow, gelatinous, strings of eggs that are coiled around seaweeds and on rocks. Sea hare larvae are thought to settle out of the plankton onto red algae at the bottom of the shore, and on which they feed. As they grow, they gradually move upshore and, as a consequence, their diet changes so that they feed more on brown and green algae. It is reported that [Aplysia punctata _]feeds on species of the red algae _Polysiphonia and Ceramium when young, and then moves on to the brown and green algae Fucus and Ulva as it moves up the shore. In France they have been observed feeding on [_Codium tomentosum _]Stackhouse, 1797.
When handled, Aplysia punctata ejects clouds of purple ink from pallial glands and an acrid, foul-tasting, substance from sub-pallial glands. It has been suggested the latter contains sulphuric acid, which is distasteful to any fish that may try to eat it. These secretions are its initial defences but, in extreme circumstances, the lateral margins of the foot can be expanded into parapodia and, by sending undulatory waves along them, the sea hare can lift itself from the seabed and swim gently.
Species of Aplysia have been studied as a model organism by neuroscientists, because they only have about 20,000 neurons, making them a favourite subject for investigation. In particular, its siphon-withdrawal response is mediated by electrical synapses, which allow several neurons to fire synchronously, known as the Aplysia gill and siphon withdrawal reflex. The quick neural response is necessary for a speedy reaction to danger resulting in the release of the noxious defensive secretions (Kandel et al. 2000).
Eric R. Kandel was one of the recipients of the Nobel Prize in Physiology or Medicine in 2000 for his work on these neural pathways. The final gem of information for this story must, however, remain with Nellie Eales. She had obtained her Ph.D. in the 1920’s with a thesis on the anatomy of a foetal elephant. She wrote five papers on this subject for her Ph.D., the specimen on which they were based coming, dead, from the London Zoo. I provide one reference (Eales 1926) of the five she wrote because the title is itself amazing – ‘The anatomy of the head of a foetal elephant, Elephas africanus (Loxodonta africana)’. As a naïve young student, I asked Nellie how she had achieved this anatomical feat? She answered that the transverse tissue sections she had cut of the head were mounted on sheets of window-pane glass!
Tonna zonatum (Green, 1830) is a big gastropod with a near globular, but thin and brittle shell that can reach up to 15 centimetres in height and lives on the continental shelf of the South China Sea, off Hong Kong. It was believed to feed on holothurians (sea cucumbers), but this had never been confirmed. The opportunity to undertake such an investigation did not arise until April 1989 when a single individual was obtained and kept in the marine aquarium of the University of Hong Kong. It was actually kept for 21 weeks and, during this time, it was not only possible to draw it feeding, in this case, on an individual of the sea-cucumber Holothuria cinerascens (Brandt, 1835) but also to experiment on it by offering a choice of different holothurian prey.
Over the course of the 21 weeks, it was, it transpired, capable of eating five different sea-cucumber prey species and did so extremely rapidly. Tonna [zonatum _]would approach a holothurian and seemingly ‘measure it up’, like an undertaker, tapping it with its paired cephalic tentacles. It would then find one end of the prey and evert its enormous proboscis around it and engulf it with its monstrous mouth, all within the space of between two to five minutes. The feeling of being measured up by _Tonna zonatum caused what can only be described as panic (but on a holothurian scale) by the chosen prey. A tapped holothurian would take water into the coelom and, thereby, not only increase its girth making it more difficult for the predator’s proboscis to engulf it but also make themselves buoyant and, by writhing about, ‘swim’ free of the sediment and thereby attempt to escape the attentions of the Tonna. On another occasion, a large, 300 gram, black [Holothuria leucospilota _]Clark, 1920[ _]was attacked causing it to discharge sticky Cuvierian tubules from its anal end in the direction of the gastropod attacking it. This is the usual holothurian defense mechanism whereby the body-wall musculature contracts, seawater is expelled from the anus and this is then followed by the sudden appearance of a mass of white threads, which, on contact with seawater, resolve themselves into a number of white tubules. These, in turn, swell, writhe and elongate attaining almost a metre in length before breaking away from the holothurian. This is typically followed by evisceration. The ejected tubules are intensely sticky and are generally considered to be organs of defence since they are capable of entangling predatory animals such as crabs and fish. Early research on holothurian Cuvierian tubules was undertaken by Robert Endean (1926-1997), one of Australia’s most eminent marine biologists (Endean 1957).
But for my sea-cucumber, however, there was no escape, the Tonna ate the Cuvierian tubules and the eviscerated guts and, two days later, re-attacked the now slimmer holothurian and devoured it too.
This research was published subsequently (Morton 1991), but the most interesting thing about the Tonna was its effect on people. If Mr Lam Man Hung, the aquarium technician, suspected an attack was about to occur (because not only did the animal feed at least once each week but also because he too became so fascinated by it he was constantly watching for signs of an attack), he would alert not only me but all the postgraduate students plus any undergraduates in the surrounding laboratories. As a consequence, a large crowd would gather to see the spectacle. For that is what it was. And, a rarely seen, remarkable, demonstration of a predator-prey relationship.
The bay of Hoi Ha Wan in the notheastern New Territories of Hong Kong was and still is home to a rich variety of scleractinian corals and it was long argued that it should be protected as a marine park. This eventually come about when on 31 May 1995, Hong Kong’s Legislative Council passed the Marine Parks Bill, but it would not be until 14 July 1996 that Hoi Ha Wan was designated as Hong Kong’s first marine park. The shallow sea-bed of Hoi Ha Wan comprises a long expanse of sand. And, snokelling over this, actually on a number of occasions, it became apparent that there were numerous large burrows in the sand and at some of their entrances sat a small fish – a goby. The local diving fraternity members were asked if they could try and capture some of these little fishes. This turned out to be more difficult than expected but was achieved eventually by putting a mesh net over the holes and then digging into the burrow beneath. In due course, the fish was capture but, surprisingly, along with it was a shrimp. Actually, a so-called snapping shrimp belonging to the Alpheidae.
It took a long time to identify these animals, as this symbiotic association, well known from elsewhere, had never been described from Hong Kong before. Eventually, however, the shrimp was identified as[_ Alpheus rapacida_][* *]De Man, 1908 and the fish as Vanderhorstia mertensi Klausewitz, 1974[_. _]They live together – sharing the same burrow – and do so for mutual benefit. The uniformly brown-coloured shrimp is some 6-7 centimetres long and characteristically has one claw, in this instance the right, bigger than the other. This heavier, larger claw, which is almost the size of the carapace, can be snapped shut, producing a loud crack – hence the common name for these members of the Alpheidae – snapping shrimps. The chela is closed suddenly under strong muscular pressure, a peg on the moveable finger of the claw fitting into a socket on the fixed one. The heavily calcified finger tips come together sharply, and a jet of water is emitted, displaced by the peg. The jet is produced at such speed that it literally vaporises, raising the temperature so high that if it finds contact with, for example, a crab ambling too close to the burrow entrance it is momentarily stunned and, upon recovery, hastens away. The snapping sound produced by the chela, the equivalent being a human’s finger click, may also be used in communication between individuals – signalling their presence.
It is the task of the shrimp in this association with the fish to deter accidental intruders into the burrow. The shrimp, however, performs another function which is to keep the burrow free of silt and, occasionally, one can see it emerge from the entrance pushing a pile of sand and small stones ahead of it with its larger chelae. It toils away incessantly and persistently as each movement of water in the sea above wafts more particles back down the burrow. But, the shrimp also has pleopods beneath its abdomen and these serve the additional function of keeping the burrow aerated by drawing a strong current through it.
The role of the goby Vanderhorstia mertensi in this mutually beneficial relationship is to be the burrow watchman. The goby is some 11 centimetres long with big eyes, five vertical stripes on its body and pretty smudges of white and yellow on its head and on the first element of the dorsal fin. It sits, raised high on its pectoral fins, at the entrance to the burrow and keeps a lookout for danger and, almost certainly, any passing pieces of potential food. At the first sign of danger, the goby vanishes at lightening speed into the burrow, which is actually a highly complex and sophisticated gallery and not just a simple hole. This alerts the shrimp which also disappears downwards. The relationship between the snapping shrimp and watchman goby is thus one that can be classified as mutualism – both benefiting from living one with the other.
Both species are naturally distributed in the Central Western Pacific and Indian Ocean but they have both been introduced quite recently into the eastern end of the Mediterranean probably via the Suez Canal (Özcan et al. 2007, Bilecenoglu et al. 2008).
While participating in a research workshop held on Rottnest Island, off from Perth, Western Australia, I drew Nassarius glans particeps (Hedley, 1915)[* *]but did not research it. In January 1996, another workshop was convened on Rottnest Island and Joe Britton, Professor of Biology at Texas Christian University, Fort Worth, Texas, U.S.A. (1942-2006) did and was joined by his son, David Britton, who was then also studying to be a marine biologist and is now a researcher with the United States Fish and Wildlife Service. Together Joe and David worked on a pair of scavenging snails, the nassariid Nassarius glans particeps and the buccinid Cominella tasmanica (Tenison-Woods, 1878). As with other sympatric scavengers, the pair of snails showed behavoural interactions which David and Joe described (Britton and Britton 1999).
Nassarius glans particeps has a beautiful shell. It has a creamy brown shell, that is streaked with light brown rays. The animal itself is almost pure white but is prettily patterned with dark brown to black dots. There are a pair of trailing tentacles to the rear of its foot and the animal glides elegantly over the clean sands of Rottnest Island’s intertidal pools like majestic royalty. Joe and David showed that, in Thompson Pool, [Nassarius glans particeps _]would arrive at experimental bait very quickly and feed on it. [_Cominella tasmanica _]was altogether slower arriving only after the nassariid had departed. Moreover, if two _Nassarius glans particeps arrived simultaneously, the dominant individual would demonstrate a shaking behaviour. If this did not deter the subordinate, the more dominant individual would extend its proboscis and scrape at the tissue of the subordinate with its radula. In this way, the dominant individual had first choice of the meal. Such research was pioneering in demonstrating complex intra-specific behaviour hitherto unappreciated.
Cominella tasmanica is interesting too, but a member of the Buccinidae not Nassariidae, and is a scavenger, as Joe and David also demonstrated. Its congenerics are also predators feeding on bivalves, as later demonstrated when undertaking research on Cominella eburnea (Reeve, 1846) in Princess Royal Harbour in Albany, also Western Australia, but this time alone (Morton 2006). It is generally believed that representatives of the predatory Buccinidae gain access to their prey by means other than drilling. It had previously been reported, however, that in southwest Western Australia, Cominella eburnea, drills its bivalve prey. Research in Princess Royal Harbour showed that the species feeds principally upon the venerid bivalve Katelysia scalarina _](Lamarck, 1818) but also upon a range of sympatric gastropods and always by either marginal ([_Katelysia scalarina) or apertural (little gastropods) access using the proboscis. Numerous observations of [_Cominella eburnea _]attacking [_Katelysia scalarina _]in the laboratory, similarly by marginal proboscis insertion, substantiated these records: drilling was never observed.
[Clinocardium nuttallii _](Conrad, 1837) known commonly as the Pacific cockle is found from San Diego in California north to the Bering Sea and across the Pacific to Japan (Quayle 1960). The species generally occurs in bays in sediments ranging from coarse sand to silt/clay, and from the shallow intertidal to depths of 200 metres. It lies barely buried at the surface or just below the surface of the sediment, often in eelgrass, [_Zostera marina _]L., beds. _Clinocardium nuttallii was examined at the Moss Landing Marine Laboratory, Monterey, California, U.S.A. in July 1991.
Cockles have broad, high, shells with radial ribs and Clinocardium nuttallii can grow to a height of 14 centimetres. The cockle’s profile from the side is heart-shaped, providing it with another common name. It also has a thick shell that is, typically, characterised by more than thirty distinct radial ribs covering the entire surfaces of both shell valves. The ridges are so strong that they become undulations, which interlock with one another at the ventral margins of the valves. Growth rings encircling the shell are also prominent. The shell is usually light tan, mottled with various bands or blotches of brown, especially in younger individuals. Its siphons are short and when a heart cockle is buried in the sand, their edges and whole of their outer surfaces are covered with a mass of white papillae.
The mantle margin also has tiny tentacles with tiny eyes. The thick shell is needed because its habitat, especially the seagrass areas are full of predators. Such predators include the sunflower seastar,[_ Pycnopodia helianthoides_] Brandt, 1835. This is the largest sea star in the world, with a maximum armspan of over one metre. Sunflower seastars usually have 16 to 24 limbs which they use to feed mostly on sea urchins, gastropods and [Clinocardium nuttallii. _]Another seastar predator is the giant pink sea star, [_Pisaster brevispinus _]Stimpson, 1857[. Pisaster brevispinus ]can reach a diameter of 60 centimetres and weigh over one kilogram, It has a soft, flabby texture. _Metacarcinus magister (Dana, 1852) is a species of crab that also inhabits eelgrass beds and typically grows to twenty centimetres across the carapace. To escape such, relatively huge, predators, Clinocardium nuttallii has a strong escape response rapidly extending its large foot and leaping away in a series of explosive jumps. It responds to the touch of the seastars and crabs but the mantle eyes also provide it with a shadow reflex, detecting subtle changes in light intensity, and responds by shutting its interlocking shell valves tightly.
Unusually, Clinocardium nuttallii is a simultaneous hermaphrodite spawning in July and August at two years of age (Gallucci and Gallucci 1982) but can live to between 15-19 years in Alaska. The small pea crab, Pinnixa faba (Dana, 1851) can be found living inside the mantle cavity of this cockle and other co-occurring large bivalves.
Despite the general availability of the Pacific cockle, it is only harvested occasionally by recreational clam diggers but has received relatively little scientific attention in contrast to the European cockles, [Cerastoderma edule _](Linnaeus, 1758)[ _]and [_Cerastoderma glaucum _](Bruguière, 1789), which are not simultaneous hermaphrodites, and whose growth and reproductive cycles are well known (Rygg 1970, Boyden 1971, Kingston 1974).
Although many research vessels, including the famous H.M.S. Challenger, had dredged off the Açores, no such work had been undertaken in modern times. In the summer of 1991, it was considered time to rectify this situation and a research meeting was held at the village of Vila Franco do Campo on the island of São Miguel. The specific aim of the research workshop was to undertake dredging of the near-shore Açorean sea bed. Also a participant in the workshop was the Danish scientist Jørgen Knudsen (1918-2010) (Morton 2009), famous for his monumental studies of deep-water molluscs (Knudsen 1967). Accordingly, Jørgen was prevailed to give us plans for a dredge and these were taken to the local blacksmith where the structure was made. Lowered into the water for the first trial, the dredge quickly filled up with stones and could not be lifted off the seabed. A passing fishing vessel with a powerful winch came to our assistance and it was eventually dragged to shore. Undaunted and with plans scaled down by 95%, a new dredge was tested but even so took a winch and four strong men to bring aboard. We were, however, left with a basin full of material that contained a few animals, including the two bivalves here illustrated – Gari costulata (Turton, 1822) and [_Moerella donacina _](Linnaeus, 1758). These were the first two deeper-water endobenthic bivalves ever drawn, alive, from the Açores.
Gari costulata ranges from the west coast of Great Britain south to the Mediterranean, the Canary Islands, Madeira and, now the Açores. It typically lives at depths greater than sixty metres. [Gari costulata _]is a member of the Tellinoidea and is, as a consequence, a deposit feeder with long separate siphons[. _]The shell is pinkish white with patches, streaks and radiating rays of purple or red.
[Moerella donacina _]is also a member of the Tellinoidea and, hence, also a deposit-feeder with long siphons. This shell also has a whitish-yellow background colour but is ornamented with pink rays radiating from the shell’s apex. The two very fragile shells are both given the general term of sunset shells although later there is an illustration of _Tellina tenuis da Costa, 1778 more fully deserving of the name.
The outcome of this first modern attempt to dredge the deeper sea bed of the Açores has been, after two more subsequent expeditions, the production by their co-ordinator, Professor A.M de Frias-Martins, of a list of the marine Mollusca of the Açores (Frias-Martins et al. 2009).
The shell of [_Finella pupoides _]A. Adams, 1860 (Scaliolidae) is unmistakable, owing to the peculiar profile of the spire and the shape of the aperture, which is broadly rounded and lacks a siphonal canal. The shell varies in length between 2-4 millimetres and in colour from white to pale yellow, sometimes even purple-brown. The shell typically has between 7-8 chubby whorls and indistinct brown bands in the middle of each one. The shell is elongate, fusiform, with a blunt-ending apex. The protoconch is smooth and comprises two whorls. The sculpture of the teleoconch comprises characteristic flat-topped spiral cords with rather weak axial ribs. These form a fine reticulate pattern on the upper whorls, but this is reduced on the body whorl.
The head has a pair of long, slender, translucent, cephalic tentacles and possess moderately large black eyes set in weak swellings at the outer tentacle bases. The most distinctive feature of the animal is the large, long, mobile and translucent snout. The snout, dorsally adorned with white spots, is capable of great distension when feeding and at this time, the yellow-pink-orange odontophore occupies the distal end of the snout. When feeding, the cowl-like snout is lifted at one side and then the other in a see-saw-like manner and surface detrital particles are captured and can be seen moving down the oesophagus. The foot is expanded laterally anteriorly and pointed posteriorly, the latter also possessing a small brown operculum.
The most characteristic features of [Finella pupoides _]are the pallial tentacles[. _]Two small ones arise from the left and right sides and a third long one also arises from the right side, to trail behind the animal as it moves. These are translucent, colourless and covered in cilia. They are probably sensory.
[_Finella pupoides _]has an Indo-West Pacific distribution from Réunion in the Indian Ocean eastwards into the Pacific Ocean and including Southeast Asia and Japan (Hasegawa 1998). It lives in sand or mud in the shallow waters of bays at depths of up to approximately ten metres. Live specimens are, however, rare and the illustrated individual was collected from Tai Tam Bay on the southern coasline of Hong Kong Island by seiving sediments low on the shore (Ponder 1994) and drawn at the Swire Institute of Marine Science of the University of Hong Kong at Cape d’Aguilar in April 1992.
Though uncommon in the Asian focus of its distribution, Finella pupoides has been introduced into the Mediterranean Sea through the Suez Canal and thereby into European waters. This marine alien has been recorded in low abundances in the Eastern Mediterranean for over half a century. Recently, however, populations inhabiting the southeastern Levantine coastline have grown so that the species is here extremely abundant (Öztürk and Can 2006). Samples collected off the coast of Israel during 2010–2011 by the Israel Oceanographic and Limnological Research Institute contained up to 3,300 individuals per square metre (Bogi and Galil 2013). The larvae of Finella pupoides are planktotrophic, thereby ensuring long-distance dispersal but it seems probable that this non-indigenous invasion was effected by transport in the ballast water of ships passing through the canal from the Indian Ocean via the Red Sea.
I visited Bermuda during August 1993 to sign an agreement between the Swire Marine Laboratory of the University of Hong Kong and the Bermuda Biological Station for Research, now both re-named in 1994 and 2006, respectively, the Swire Institute of Marine Science and the Bermuda Institute of Ocean Sciences. In addition to signing the agreement for research co-operation, I wanted to examine the flora and fauna of Bermudan shores for comparison with those on the also Atlantic islands of the Açores and those of both North America and mainland Europe at Portugal. This was in the context of a book I was co-writing on the coastal ecology of the Açores (Morton et al. 1998)
It transpired that the visit coincided with a period of a full moon and so was able to see one of the great marine spectacles in the world – the mass spawning of the palolo worm, [Odontosyllis enopla _]Verrill, 1900. The spawning takes place every lunar month, three days after the full moon and 57 minutes after sunset. At this time, females swim rapidly to the surface of the sea and emit a bright green fluorescent chemical that attracts the males to them. The bioluminescing worms create a mini-firestorm in the sea and in fact _Odontosyllis enopla is also called the fire-worm.
Finally, however, examining the marine laboratory’s collection of marine life and seeing how it was organised, I came across a specimen of the phyllosoma larva of a spiny lobster. At the time, the identity of which species of panulirid lobster the specimen on display was the larva of was unknown. But, admiring its amazing form, I decided to draw it. Later, it was identified as [_Panulirus argus _](Latreille, 1804).
Only relatively recently has Goldstein et al. (2008) successfully reared [Panulirus argus _]larvae in the laboratory through all its planktonic (phyllosomal) larval stages from egg to early juvenile – the puerulus – which resembles a miniature adult and will settle onto the sea bed and become one. These authors showed that the fertilised eggs of _Panulirus argus go through ten phyllosomata stages identified as phyllosoma I-X and examination of their paper suggests that my drawing is of the VI phyllosomal developmental stage and almost, if the limbs are extended, some 35 millimetres across – a huge size for any marine larva.
[_Panulirus argus _]is the most widespread, commercially important, and extensively studied spiny lobster in the western hemisphere, the adults of which live on reefs and mangrove swamps in the western Atlantic Ocean. What amazes me is that to obtain an adult lobster, the species has to go through such a long (a mean of 174 days, half a year) period in the plankton. There, it must be not only at the mercy of ocean currents which may or may not take it to a suitable site for adult occupation, but that each stage must be completed successfully in order to move onto the next and eventually attain adulthood. Not just this, but each larva must not just find sufficient food to survive and moult into its next stage, but not be eaten by predatory fish of which there must be many. Perhaps the larva’s large size helps protect it from such predation.
A favourite beach in Hong Kong was Starfish Bay, or Hoi Sing Wan, on the southern shores of Tolo Harbour. In the early 1970’s, it was near pristine but, slowly, as the new city of Shatin was built, on the tiny shoulders of the original fishing village of the same name, the water quality of Tolo Harbour deteriorated dramatically. Slowly, but surely, the intertidal biodiversity of Hoi Sing Wan was lost and only one animal species seemed to benefit from it. This was the tiny, approximately ten millimetre long, nassariid snail [_Nassarius festivus _](Powys, 1835). And it now occurs on the sand surface of Starfish Bay in countless numbers. In the late 1980’s, research on the behaviour of this species began and it was discovered that it was attracted to carrion, notably any broken bivalve it was presented with.
The most obvious feature of this little snail is that as it approaches any piece of carrion, such as the broken and dying venerid bivalve Tapes philippinarum (A. Adams & Reeve, 1850), illustrated opposite, it extends its proboscis in anticipation, the so-called proboscis extension reflex, which allows the observing researcher to identify almost precisely the moment it begins to feed. The other important behavioual feature of Nassarius festivus is that as soon as it has fed to satiation, it departs the food. Hence, one is able to calculate exactly for how long each animal fed and by using an extremely fine balance, how much food was eaten can be calculated. By sacrificing some animals and obtaining a relationship between shell height and tissue weight, one can then calculate very precisely how much each individual snail ate in one meal or, for example, in one day as a proportion of its body weight and/or shell height.
It turned out that _Nassarius festivus _ could eat something like 25% of its body weight in a single meal – the equivalent of us eating 200 Big Mac’s at a single sitting. Another interesting behaviour discovered was that if one crushed up a couple of snails and sprinkled them over a group of carrion-feeding brothers and sisters – they would all rush away from the food – leaving it uneaten. That is, the crushed conspecifics made the snails believe that a bigger predator was eating them. But, if one starved the snails, so that when they were really, really, hungry, when crushed conspecifics were put on top of the group of feeding snails, they did not flee in terror. They stayed put. That is, their hunger overcame their fear of being eaten themselves.
We also showed that not only did the snails compete among themselves for food (intraspecific competition) but also with the local sandy shore hermit crab, Diogenes edwardsii (De Haan, 1849) (interspecific competition), so that in reality each piece of carrion, for example a dead fish, that ends up on a sandy beach becomes a little struggle for survival as snails, hermit crabs and liitle intertidal gobies, fight for access to it. Darwinian natural selection in action. I and my co-workers wrote many papers on [Nassarius festivus _](Morton 1990b, Britton and Morton 1992, Morton _et al. 1994, Morton and Chan 1999, 2003, 2004, Morton and Yuen 2000, Chan and Morton 2001, 2003, 2005)[_, _]and it became a favoured research animal, recognising that here was a little hardy creature that could so easily become an educational tool, an environmental monitor of pollution and an ecological and behavioural test species for elucidating and testing the theory of evolution itself.
The Arminidae are poorly known nudibranch (naked gilled) gastropod molluscs. They burrow into sediment, live mainly in deep water, and virtually nothing is known of their biology. Even knowledge of the anatomy of arminid nudibranchs, other than Armina,[_ _]is fragmentary. Characters shared by members of the family are an elongate, flattened body narrowing towards the tail, longitudinal ridges or pustules on the notum, a distinct oral veil and paired retractile rhinophores sometimes with a caruncle between them. In the mouth, each radular row comprises a broad and denticulated central rachidian tooth and partly denticulated laterals (Kolb 1998). Radular morphology is apparently especially important in that it shows remarkable inter-specific variation in form, probably related to the feeding diversity of the group on various cnidarians (Gosliner and Fahey 2011).
Within the Arminidae, [Armina _]is the largest taxon with more than 70 species described. Kolb and Wagele (1998) concluded that the Arminidae probably originated in the western Pacific, near Japan and have radiated in a westerly direction. Accordingly, the most primitive species of _Dermatobranchus are restricted mainly to the western Pacific while the more derived species of Armina have a world-wide distribution. New species of the Arminidae are now being collected constantly and identified: Gosliner and Fahey (2011), for example, recently described twenty new taxa.
In April 1994, a research trip was made to the Third Institute of Oceanography at Xiamen, Fujian Province, China. The research trip was undertaken to perform benthic dredging and trawling in the waters around the port city of Xiamen, which is situated at the junction between the East China and South China Seas. It is also located in the Taiwan Strait opposite the island of Taiwan. During the dredging/trawling programme, a single individual of Armina longicauda Lin, 1981 was collected. This species lives on sandy-mud sea bed at depths of between 170-260 metres (Lin 1981) and is restricted in distribution to the East and South China Seas; at least, no other records have been published.
The elongate body of Armina longicauda is some 12 centimetres in length, with the hind end forming a black tail. The head-veil is semi-circular and covered with three series of approximately forty short conical papillae, sometimes an iridescent blue with a yellow margin. The paired rhinophores are small, white apically, and are longitudinally lamellate. There does not appear to be any eyes. The mantle covering the main body is somewhat broad and rounded anteriorly but posteriorly tapers into a long tail. The back is black-brown divided into between 18-20 yellowish-white longitudinal ridges. The foot is narrow, rounded in front but with left and right corners produced into horn-like processes. Little else is known about the biology of [Armina longicauda. _]Information on other arminids, however, has shown them to be predators: species of _Armina are said to feed on pennatulacean cnidarians, species of sea pens. In the waters off California, sea pansies, Renilla kollikeri Pfeffer, 1886, are preyed upon by Armina californica (J.G. Cooper, 1863). When attacked, the sea pansy raises its flattened rachis, so that water currents lift its stalk out of the sediment and it tumbles away from the predatory arminid (Kastendiek 1976). In British waters, the similarly deep sea Armina loveni (Bergh, 1860) feeds on the slender sea-pen Virgularia mirabilis (O.F. Müller, 1776), which may grow to 60 centimetres upwards out of the sediment and comprising a slender central stem with leaves arranged in two opposing lateral rows.
Four species of surf clams occur on Hong Kong’s exposed sandy beaches such as Big Wave Bay on the island itself. These are Donax incarnatus Gmelin, 1791, [Donax faba _]Gmelin, 1791,[ Donax cuneatus ]Linnaeus, 1758 and[ Donax semigranosus_] Dunker, 1877 (Ansell 1985). Alan D. Ansell (1934-1999) was a friend and eminent British marine biologist, who died too young, and after whom the Alan Ansell Research Aquarium at the Scottish Marine Institute at Oban, Scotland, is named. Another of Alan’s friends, Lloyd Peck of the British Antarctic Survey, summarised Alan’s life as follows: ”Despite all of the fine accomplishments in his career, being an excellent scientist with a humble demeanour, possibly his greatest attribute was enthusiasm.”
There are over fifty species of surf clams, colloquially termed pipi’s scattered throughout the world’s tropical oceans and the four Hong Kong species are distributed widely in the Indo-West Pacific. [Donax cuneatus _]and[ Donax semigranosus_] are the commonest species locally and the former can reach a shell length of almost 40 millimetres, whereas the latter has a maximum length of approximately 15 millimetres. Of the two, however, Donax semigranosus is the commonest species.
Species of Donax are remarkable for their ability to migrate up and down sandy beaches, maintaining their position relative to the wash zone of the waves by co-ordinated movements involving emergence from the sand, surfing in the wash or backwash of the waves, followed by rapid reburial as the energy of the wash is reduced. Donax semigranosus was the first donacid for which this behaviour was described from beaches in Japan (Mori 1938, 1950). Surf clams do this because they have exploited a niche in the coastal marine environment where they can collect, by filter feeding, fine particles of detritus stirred up by the crashing waves. To do this most efficiently, however, they have to keep in the position of the breaking waves and which changes over the tidal cycle. In Hong Kong, the maximum tidal range is four metres and because the tides are generally semi-diurnal, this means they are migrating vertical distances, up and down, of two metres, approximately twice a day.
To do this, they jump out of the sand and go tumbling down the beach, or in the opposite case, up it. They can re-bury in a matter of seconds using a proportionately large, axe-shaped, foot. Similarly, the shape of the shell is anteriorly pointed and posteriorly foreshortened, so that it too is beautifully adapted for rapid re-burial. Hence, the vast majority of people who visit the same sandy beaches to party, sunbathe, swim and paddle, never see the clams.
Another advantage of occupying such beaches, in contrast to flat, protected shores of sand and mud, is that few predators occur here – being unable to catch the speedily migrating and fast re-reburying clams. As we will see later in the book, however, one predator, the moon shell [Polinices incei _](Philippi, 1873) has exploited the juveniles of the pipi _Donax deltoides Lamarck, 1818, by the simple expedient of itself surfing up and down wave-exposed shores on the coast of Queensland, Australia.
In 1977, I drew the small, only 25 millimetres across, and distinctively brightly patterned scallop Minnivola pyxidatus[* ]Born, 1778.[_ ]The under valve is a uniform dull cream. Conversely, the upper shell of [_Minnivola pyxidatus][ *]is red-brown but variously blotched with patches of white framed in dark brown to black. The exquisitely patterned little animal is, however, highly polymorphic in terms of coloration so that some individuals are just a dull mauve whereas others develop the full pattern. This is an example of cryptic polymorphism since separate morphotypes are not identifiable, and allows the animal to be camouflaged against a wide range of background sediment colours and textures. Later, more time was spent with the animal and this research published information on its behaviour and anatomy (Morton 1996a).
[Minnivola pyxidatus _]has a wide geographical range in the Western Pacific and occurs in Hong Kong only in the cleanest southeastern oceanic waters at depths of between 13-44 metres. Typically, each individual attaches to the sediment, clearly relying on camouflage for protection, by just two three fine byssal threads. If attacked, however, as for example was demonstrated when some individuals of the co-occurring muricid gastropod predator _Rapana bezoar (Linnaeus, 1767) was placed in an aquarium with it, it can swim (Morton 1994).
When touched by the muricid, the scallop breaks its byssal attachment to the sediment and undertakes an escape reaction that involves a dozen or so phasic adductions of its centrally located and single adductor muscle that lifts the individual clear from the sediment by a few centimetres. Minnivola pyxidatus can then swim a short distance by clapping its shell valves together, thereby achieving lift, and with it a form of jet propulsion. That is, the clapping together of the valves, pumps jets of water out from the mantle at gaps on each side of the shell’s auricles dorsally. This makes the animal swim forward in bouts of jerky lunges and, most importantly, removes it from the immediate vicuinity of the predator.
But, just how does it detect the predator? It was shown that the array of yellow tentacles which arise from both mantle margins are interspersed with gleaming black dots which are actually quite sophisticated eyes that are also seen in other shallow water scallops, such as the ecological equivalent of Minnivola in British waters, that is, albeit much larger, Pecten maximus (Linnaeus, 1758). The pectinid eye is extraordinarily complex and first described in any detail by W.J. Dakin (1883-1950) for Pecten maximus. Born and educated in England, William Dakin here undertook his research on pectinid eye structure (Dakin 1909) but eventually settled in Australia, first in Perth at the fledgling University of Western Australia and then at the University of New South Wales in Sydney. Here, he wrote another classic book on the intertidal ecology of Australian shores published in 1952 (Dakin 1952) and assisted by two other Australian marine biological luminaries, Isobel Bennett (1909–2008) and Elizabeth Pope (1912-1993).
The eyes of both Pecten maximus and [Minnivola pyxidatus _]comprise a double retina and focussing lens and, thus, resemble closely the structure of the mammalian eye. It is probable, however, that on the sea bed they are only able to distinguish changes in light intensity and react to those, that is, they share a shadow reflex that initiates swimming. _Minnivola pyxidatus, however, also possesses small pallial papillae adjacent to they eyes and these are thought to be mechanoreceptors that are sensitive to touch, as for example the slowly creeping and not shadow-forming, gastropod Rapana bezoar giving the scallop a double means of detecting a wide variety of predators.
The northern shore of the island of São Jorge in the Açores is steep and precipitous, the road down to the base from the top of the island performing numerous, terrifying, hairpin bends. On this side of the island are two fajãs, _]or lagoons. Fajã de Santa Cristo and Fajã dos Cubres were created by massive rock falls from the mountain behind, during a massive earthquake that shook São Jorge on the 9 July 1757 – “[_the most violent seismic phenomenon felt in these islands after their settlement” (Agostinho 1935). In both cases, over time, the enormous piles of resulting rocks have been moulded by the sea into concave boulder ramparts behind which have formed the lagoons. In the case of Fajã de Santa Cristo, the lagoon has access to the sea via a channel, but in the case of Fajã dos Cubres, there is no channel and the water in it appears altogether more hypertrophic with mats of algal gut weed Enteromorpha intestinalis Linnaeus, 1753 and a species of Chaetomorpha floating on the surface. Both fajãs have inputs of freshwater into them but the lagoon at Fajã dos Cubres is fringed for most of its circumference by the sharp or spiny rush Juncus acutus Linnaeus. This lagoon also is fringed, internally to the Juncus, by a sea grass – variously called ditch-grass, tassel pondweed, beaked tasselweed and widgeongrass, [Ruppia maritima _]Linnaeus. Widgeongrass because it is reputedly a favourite food of the duck which its name bears. When initially discovered, little did I know that I had accidentally stumbled upon the rarest plant in the Açores and known only from Fajã dos Cubres (Morton _et al. 1995) in the archipelago.
The stems of [_Ruppia maritima _]bear dark green alternate or opposite dark green leaves which are exceedingly, one millimetre, thin (A). The flowers are borne on a short peduncle which arises at a growing point and elongates after anthesis. The two stamens lack a filament and each possesses a pair of two-celled anthers (B). Following release of the anthers, the peduncle elongates to a length of about 20 millimetres and two female flowers develop each with four styles ©. [_Ruppia maritima _]is basically self pollinating, the pollen grains being illustrated in D, and the peduncle elongates yet further, coils and bears black drupelets (E). All these stages were present in the lagoon at Fajã dos Cubres and, hence, clearly, the species is here a success making it, arguably, one of the most important wetland sites in the Açores.
To protect such delicate wetlands, the Convention on Wetlands (Ramsar, Iran 1971), thereafter called the Ramsar Convention, is an intergovernmental treaty that embodies the commitments of its member countries to maintain the ecological character of their Wetlands of International Importance and to plan for their wise and sustainable use. Fajã dos Cubres and a sister, more marine, lagoon a few kilometres along from it – Fajã de Santo Cristo were designated as Ramsar Sites in 2005. I have concluded, however, that this level of protection is neither strong enough nor sufficiently appropriate to protect the sister fajãs of Cubres and Santo Cristo on São Jorge and I have, therefore, recommended to the island’s governor in 2014 that these two locations be designated as World Heritage Sites (Morton 2014).
In June 1996 a teacher from The National University of Singapore, Dr Tan Koh Siang, was invited to work in the Cape d’Aguilar Marine Reserve at the Swire Institute of Marine Science (SWIMS) of the University of Hong Kong. Koh Siang had an interest in predatory gastropods and we resolved to tackle one of the smallest species in the reserve, the 8 millimetre long buccinid whelk [Engina armillata _](Reeve, 1846). Not just the smallest, but the most enigmatic – virtually nothing being known of not just it but its congeners either. The species occurs under stable boulders in the reserve living amongst the encrusting mat of sponges, bryozoans, barnacles and tubeworms plus a range of other more mobile little animals. A few trial experiments discovered that, of all the animals inhabiting such boulders, _Engina armillata preferred to prey on encrusting tubeworms.
But, the question was then: how to examine its diet in more detail and discover just how important it was in controlling the abundances of such common encrusting animals? A search duly began for empty bivalve shells on the seabed of Hok Tsui Wan, that is the focus of the reserve, and which had tubeworms attached to them. Many such shells were found and by firstly removing empty tubes and secondly by removing some of the living tubes, we ended up with little ‘platters’ of tubeworms of not just different numbers and sizes but also of different species compositions. These shells were placed in their own little pots through which ambient seawater flowed and to them were added Engina armillata individuals, also of different sizes. They were then left to make their choices and feed.
It transpired, that of the four commonest species of tubeworms in the reserve, that is, Hydroides elegans (Haswell, 1883), [Pomatoleios kraussi _](Baird, 1865) _Simplicana pseudomilitans and Spirorbis foraminosus Grube, 1872, the first of these was the favoured prey item (Tan and Morton 1998).
An individual of Engina armillata is illustrated feeding on an Hydroides elegans tubeworm. The predator aligns itself along the length of the worm and everts its proboscis into the aperture of the worm’s tube and attacks its prey using the radular teeth that are located at its tip. The illustrated specimen appears to have a look of surprise in its eyes as though caught in the flash of a camera’s glare. This is an illusion, however, although this is the first and, as far as I am aware, only illustration of an Engina species attacking its prey.
There are many species of barnacles most often noticed encrusting rocky beaches as a distinctive and crowded band in the mid-intertidal. Species of the goose barnacles, Lepas, which are characterised by a stalk or peduncle, are also commonly washed ashore attached to natural floating objects (flotsam) such as sea bird feathers, cuttlefish bones and wrackweed, and pieces of wood or discarded fishing nets and the inevitable floating waste plastics (jetsam). In the days before it was realised that birds migrate, it was thought that the barnacle goose, [Branta leucopsis _](Bechstein, 1803), developed from such a barnacle, since they never nested in temperate Europe and to ignorant eyes just seemed to disappear. Hence the colloquial English names goose barnacle and barnacle goose and the scientific name of the commonest former species _Lepas anserifera Linnaeus, 1767 from the Latin anser for goose. The Welsh monk Giraldus Cambrensis, or Gerald of Wales (c.1146–c.1223), made this claim in his Topographia Hibernia. Since barnacle geese were thought to be neither flesh nor born of flesh, they were conveniently allowed to be eaten on days when religion forbade the eating of (other) meat.
The buoy barnacle, Dosima fascicularis, differs from other floating stalked barnacles of the Lepadomorpha, however, in its ability to secrete a float of its own. The barnacle’s pelagic larval cyprid settles onto a floating particle in the sea, perhaps a tiny piece of feather or a fragment of polystyrene, and produces a sponge-like, foamy, secretion from modified cement glands in its head. The continued production of the secretion as the larva metamorphoses into an adult barnacle results in the production of a spherical, 20 millimetre wide, float that keeps its passenger close to the sea surface. It appears that, over time, other barnacle larvae attach to the float so that eventually a small colony of individuals results, each contributing to the growth of their floating home. In contrast to the huge popularity and significance of his book The Origin of Species, Charles Darwin was fascinated by barnacles and wrote a Ray Society monograph about them too (Darwin 1851).
The buoy barnacle has a transparent, basally swollen, stalk and a head-like capitulum. Like other barnacles, the capitulum has shelly plates, in this case five large ones. One of the plates, the carina, is bent and has a prominent umbo and terminates in a large flat basal disc. Unlike other stalked barnacles, however, the five plates are thin and flexible. Like most other barnacles too, and save for parasitic species, [_Dosima fascicularis _]is a suspension feeder and ‘combs’ the water with its cirri, modified limb appendages, to collect either particles of floating seston or tiny plankters.
The [_Dosima fascicularis _]colony illustrated was found washed up on the Southern Ocean sandy shores of temperate Albany in Western Australia, although the species occurs throughout the warmer waters of the Pacific and Atlantic as well as the Indian Ocean. In 2011, huge numbers of buoy barnacles were washed up on the shores of South Africa and Ryan and Branch (2012) took an opportunistic study of them there. These authors confirmed that the initial float was secreted by a significant first builder, which grew into a mature adult with a capitulum length of 50 millimetres. Other larvae swimming in the sea find the float and settle on it to create the multi-sized colony, which becomes a cross-fertilising unit. [_Dosima fascicularis _]is also occasionally stranded on the shores of north Cornwall and the west coast of Ireland, especially after storms and was, in fact, first described from such material by the British biologists James Ellis and Daniel Solander in 1786.
Psammotaea elongata (Lamarck, 1818) occurs in muddy-sand in the low intertidal and sublittoral to a depth of around 30 metres. In the marine park of Hoi Ha Wan, in the northeastern New Territories of Hong Kong, where the illustrated individual was collected, the species is uncommon (possibly because of intensive collecting), but elsewhere it occurs in extensive beds often in sandy bays near mangroves. The species has a widespread Indo-West Pacific distribution, from East Africa, including Madagascar and the Red Sea, to the Philippines; north to Japan and south to northern Queensland.
The shell of Psammotaea elongata is elongate and sub-trapezoidal-ovate in outline. The anterior margin is narrowly rounded, the posterior obliquely truncate. The ventral margin is nearly straight. A low, rounded, ridge radiates from the umbones to the postero-ventral end of each shell valve. The outer surface is smoothly polished in juveniles, but duller in larger shells. The young shell, which is much shorter and ornamented with marked radial bands of colour, as illustrated, has been considered a different species and given the name Gari minor (Deshayes, 1855). It is now known, however, that shell form and colour change with age the adult shell being externally purple but duller and often with fewer paler rays passing from the umbones to the posterior ventral margin. Psammotaea elongata can reach a maximum shell length of seven centimetres almost double that of the illustrated individual.
Psammotaea elongata _]has a large, muscular, digging foot enabling it to burrow deeply and rapidly if disturbed. Because it lives deeply in the sediments, it also has long separate, segmented, siphons. Although few studies have been made of this species, it seems likely that the siphons can autotomise if the animal is attacked, for example, if they are nibbled at by fish. In such situations, the siphons can autotomise their distal tips, losing them to the fish, but allowing survival of the remainder. Afterwards, the siphons simply re-grow. A similar predator–prey relationship is seen between juvenile European plaice ([_Pleuronectes platessa Linnaeus, 1758), which feed extensively on the extended siphons of the burrowing bivalve Tellina [= Angulus] tenuis (da Costa, 1778). Such predation activities result in a significant proportion of the prey’s energy being diverted away from growth and reproduction into siphonal regeneration leading to reduced bivalve recruitment, a reduced food supply for the fish and, eventually, prey switching (Trevallion et al. 1970). It is unknown if a similar, co-evolutionary, relationship exists between [_Psammotaea elongata _]and its predator(s).
In the focus of its range in the west central Philippines, Psammotaea elongata prefers bay areas with salinities of less than 15‰ and can survive near-freshwater conditions. Growth estimates indicate that Psammotaea elongata is a fast-growing, short-lived species, which is typical of many tropical marine organisms. Longevity was estimated to be around three years (Del Norte-Campos 2004). Although reproduction was shown to be continuous throughout the year, again typical of many (most) tropical marine species, recruitment suggested two peaks with major and minor spawnings occurring between December-January and May-June, respectively. These two spawning peaks were adjudged to be probably influenced by the monsoons, which, in turn, affect the precipitation and, hence, the salinity levels in the species’ habitat. The species has great potential for aquaculture, especially in the Philippines.
The shallow, offshore, marine environment of Shark Bay, Western Australia, is dominated by the cockle _Fragum erugatum _ (Tate, 1889). This small, less than 11 millimetre long, bivalve occurs in large numbers (4,000 per square metre) and probably completes its life cycle in one year. Cyclones, with a periodicity of ~50 years have, over the last 5,000 years, deposited huge numbers of this bivalve’s shells onto the shore, as windrows, creating the unique Shell Beach, or Hamelin Coquina. Lharidon Bight and Hamelin Pool are adjacent arms of inner Shark Bay and are separated from the outer bay by the Faure Sill. The two arms of Shark Bay are extremely hypersaline (>60‰) and radiocarbon datings suggest that the sill developed about 4,200 years ago, that is, some 1,800 years after the sea rose to its present level at the end of the Flandrian Transgression. The inner areas of Shark Bay have thus been hypersaline for a relatively short period of time. The development of the Hamelin Coquina must also, therefore, stem from this time.
The waters of Shark Bay, as well as being hypersaline, are oligotrophic and the question arises as to what is the food source that supports such large numbers of the filter-feeding bivalve Fragum erugatum? Representatives of the Tridacnidae, which are closely related to the cockles, are epibenthic heliophiles (sun-loving) using the light to sustain the huge colonies of photosynthesising dinoflagellates ([Symbiodinium _]sp.), or zooxanthellae, contained in their greatly enlarged mantle tissues. Conversely, species of _Fragum are endobenthic sciaphiles (shade-loving) but also possess zooxanthellae. Fragum erugatum positions itself at the sediment/water interface and is covered, in inner Shark Bay, by transparent water of ~50 centimetres depth. It is clearly maximising its ability to obtain the benefit of sunlight for its entrained zooxanthellae.
In virtually all anatomical respects, Fragum erugatum is a typical cockle except that in possessing symbiotic zooxanthellae it probably obtains the same nutritional benefits from the association as do the giant clams and which facilitate its survival in the hyperosmotic environment of Shark Bay. Giant clams obtain nutrition in the form of a soluble sugar, either glycerol or glucose, from their zooxanthellae and it seems probable that Fragum erugatum does too (Morton 2000).
C.M. Yonge (1936, 1975) argued that the symbiotic (mutualistic) relationship between giant clams and their zooxanthellae has led to the gigantism for which representives of the Tridacnidae are so well known. Perhaps the question of gigantism is not, however, the point about the association. Rather, it is the enhanced productivity and, hence, growth that the zooxanthellae confer upon the clams. It has been established that Tridacna gigas is the fastest-growing bivalve, accumulating shell material at a rate of about sixteen times that of other well-known bivalve fast growers – the oysters (Ostreidae) and pen shells (Pinnidae). This amounts to a shell growth rate of about 60 millimetres each year. Assuming that Fragum erugatum obtains the same nutritional benefits from the metabolites of its zooxanthellae and bearing in mind that it lives in highly unproductive, oligotrophic, waters, then the productivity advantage this species obtains from the relationship, pertains not to the achievement of a huge size but allows it, in a short life-span of probably little more than one year, to grow, mature and reproduce.
The symbiotic relationship between Fragum erugatum and its zooxanthellae has resulted in high levels of intra-body primary productivity and high levels of secondary productivity, expressed at this level by the numbers of cockle individuals present in this unique, hypersaline, bivalve-dominated, environment. Over the centuries, empty shells of the cockle washed ashore by storms, have formed the one kilometre wide and 15 kilometre long Shell Beach, or Hamelin Coquina of Shark Bay. Remarkably too, over time, the windrows of Fragum erugatum shells become mineralised to form, landward, a concrete-strong matrix. This was once cut up with huge saws into blocks by early Shark Bay inhabitants and used to make buildings – some of which survive.
[_Bullina lineata _]is a cephalaspid gastropod colloquially known as the red-lined bubble shell. It grows to a length of around 25 millimetres and has a delicate, globular shell. The individual illustrated was collected by a fishing boat hired during the spring of 1998 to be a research vessel dredging and trawling over the seabed in the southern waters of Hong Kong. This has to be one of the most exquisite of all gastropod shells. It has a white background with two bright red-purple bands encircling it. Running the length of the shell is a variable number of bright red to purple rays. These are crossed by three or four, horizontally spiraling, bands of the same colour. Much of the shell, however, is covered in life by a milky-white mantle with iridescent blue edges. The mantle also has a marbled texture and a pinkish hue, and may be yellow apically. The animal itself is nearly transparent and has an iridescent blue hue. The margin of the broad foot is an even brighter blue. The front of the foot is broadly bi-lobed and the head itself also bears broad, leaf-like, extensions. There are two small black eyes between the head shield processes. Despite being so fleshy, the animal can retract itself completely into its shell and there is a small, thin, transparent and barely visible operculum situated posteriorly on the foot.
[Bullina lineata _]is widely distributed in the Indo-West Pacific from Japan in the north to Australia in the south and out into the Pacific islands. It occurs in the low intertidal and in the shallow subtidal over the central part of its range, but is commoner in deeper depths in the more temperate extremes of its range, as in Japan, New Zealand and, now, from the Cape d’Aguilar Marine Reserve in Hong Kong. It roves over shallow algal reefs, most often at night, seeking its prey, which are believed to be cirratulid polychaetes. J.D. Taylor discovered what [_Bullina lineata _]fed on during a research trip he made to Piti Bay in Guam (Taylor 1986). An examination of the gut contents of two individuals from Guam contained remains of a species of _Cirriformia (Cirratulidae). Cirratulid polychaetes are deposit feeders, consuming mainly sediment with the aid of grooved and ciliated palps. The worm lies just beneath the shallow seabed sands in a J-shaped tube. When feeding, it extends up to four ciliated, tentacle-like, palps over the sediment surface, collecting surface deposits, which pass down the groove of each one directly to the mouth. The cirratulid body resembles the rather plain earthworms but has numerous branchial filaments arising from its body segments and which also project up into the water column to effect respiration. Shin (1982) has recorded the cirratulid Tharyx filibranchia Day, 1961 from Hong Kong waters, but it remains uncertain what[_ Bullina lineata ]feeds on here[._]
I n Western Australia’s far northwest is situated the town of Broome, first visited in July 1998. Broome was named after Sir Frederick Napier Broome KCMG (1842–1896), who in 1873 was Governor of Western Australia. Broome is a truly amazing town that has its origins in the pearling industry that commenced following the discovery in 1861 of rich beds of the giant (up to 28 centimetres across) gold- or silver-lipped oyster Pinctada maxima (Jameson, 1901) in nearby Nickol Bay. Men from the world over would come to Broome to seek their fortunes, but the divers who actually collected the shells, principally came from Malaya, China and, even, Japan. As a consequence, not only does Broome have a rich history, it has a richer cultural diversity as, one would imagine, a pioneering town like this should have. Since its inception, the cultured pearl industry based around Broome has grown apace and now its pearls are considered to be among the finest in the world.
The town of Broome is located alongside Eighty-Mile Beach, which is, as its name suggests, a long expanse of glistening sands. Behind it sits the holiday resort of Cable Beach. Some may say that the town of Broome should be re-named McAlpine after the British industrialist, Baron Robert Alistair McAlpine of West Green (b. 1942-2014) who so loved the area that he not only converted the erstwhile pearling port into the present-day attractive town of Broome, but also built the Cable Beach Resort.
The tides along this stretch of coast are huge – greater than nine metres at the time of springs and as such a mass of water retreats, approximately twice each day, it exposes a huge expanse of sand. But the coastline is also heavily influenced by waves so that Eighty-Mile Beach is also a surfers’ paradise. As illustrated elsewhere in this book, such beaches in South Africa and Eastern Australia are home to two snails – the nassariid [Bullia rhodostoma _](Reeve, 1847)[ ]and the naticid _Polinices incei (Philippi, 1873) – that surf up their home beaches to eat stranded hydrozoans and jellyfish and catch surf clams or pipi’s, respectively.
On Eighty-Mile Beach occurs a third gastropod, the nassariid, [Nassarius bicallosus _](E.A. Smith, 1876) (A). This nassariid has a shell that grows to be about 20 millimetres tall and is white when juvenile and nearly transparent. As it ages, the rim of the shell thickens (B) and it becomes coloured a pale yellow in parts with the darker internal organs also adding to a generally darker appearance. Unlike the two other gastropods, however, which can be found even as the tide is high when they search for their prey, _Nassarius bicallosus is only seen as the tide recedes and when they emerge from buried repose in the sand. They do not surf either upwards or downwards with the swash and backwash of the waves but, instead, crawl about using their expanded foot and search for the finest of particulate detritus. To achieve this, the proboscis is rarely extended, instead the mouth is applied directly to the sand and they suck up the silt that is focused as a micro-layer on its surface. As the tide recedes, they emerge in increasing numbers to feed and then all retreat into it as the tide returns (Morton 2011b).
It is of interest that such a lifestyle may be a clue as to how the carrion-feeding lifestyle of other nassariids has evolved. For, if one gets down to its level on hands and knees, and places tiny pieces of fish in the path of its advance, it will extend its proboscis quickly to try and seize it. Because its habitat is so dynamic, however, with waves arriving regularly to wash away such tiny amounts of protein and because the animal cannot surf up the beach like [Bullia rhodostoma _]to feed on stranded carrion, _Nassarius bicallosus is compelled to survive as a specialized deposit feeder on Eighty-Mile Beach. Being the only scavenging gastropod species that can survive on such shores in Northern Australia, however it does have the evolutionary advantage of exclusivity.
In early 1996, a letter was received from Professor Boris Sket of the University of Lubljana, Slovenia, asking me to look at some shells of a bivalve from caves in his country. When they arrived, I was very surprised. I had previously researched [Dreissena polymorpha _](Pallas, 1771), a byssally attached bivalve that had invaded Britain in the 1800’s and is a member of the Dreissenidae. The other extant genus of this family is _Mytilopsis which has a natural range encompassing North America but species of which, notably Mytilopsis sallei (Récluz, 1849) and Mytilopsis leucophaeta (Conrad, 1831) have colonised the brackish waters of Asia and Europe, respectively. A third genus of the family, Congeria, was believed to be extinct. But, what Professor Sket had sent was a living Congeria, namely Congeria kusceri Bole, 1962 from caves in the Dinaric karst of the former Jugoslavia.
Plans were made for a research trip to Slovenia and try and find living animals in July 1996. This was unsuccesful but a week was spent working on the preserved individuals Boris had in his laboratory. We published a paper on this work (Morton et al. 1998) and, as a consequence, a series of research visits have been made subsequently to Split in Croatia near where there is a pit called Jama u Predolcu near Metrović. From here, the Croatian Ministry of the Environment has given permission for specimens of [Congeria kusceri _]to be collected for further research. This has resulted in further publications on this amazing species (Stepien _et al. 2001, Morton and Puljas, 2013, Bilanzija et al. 2013) in which it has been demonstrated, respectively, that Congeria is a distinct dreissenid genus, that Congeria kusceri has an amazing reproductive strategy involving both ctenidal and pallial brooding and, finally that Congeria is represented not by one but three species distributed separately throughout the Dinaric Alps.
Congeria kusceri _](B) is a pretty plain creature, the shell being an uniform dull brown and the tissues lacking any pigment and not possessing any kind of light receptors – all adaptations to a troglodytic existence. Living with [_Congeria kusceri _]is another surprising animal, the serpulid tubeworm [_Marifugia cavatica[* *]Absolon & Hrabe, 1937. Marifugia cavatica (A) has a white tube and is similarly virtually colourless but possesses an array of nine palmate tentacles with which it collects food plus another tentacle that is modified into an operculum that functions rather like a sink plug to block up the aperture of the tube after the animal has retracted into it. It also functions, moreover, as a brood chamber for fertilised eggs to be held within for eventual release as late larvae
But, the real interest in these animals and the subterranean habitats they live in is the incredible diversity of other specie that occur in these Balkan caves, including the incredibly rare European cave salamander, or olm, Proteus anguinus Laurenti, 1768 This amphibian too is blind, colourless and even has external gills – a phenomenon called neoteny wherein larval features are retained into adulthood.
The greatest interest in many of these cave animals is where they arose. During the Middle Miocene, the area of what is now the Mediterranean Sea evaporated almost to dryness in the Late Messinian. – this event being called the Messinian Salinity Crisis. During this time, when sea levels fell by as much as 40-70 metres, most marine animals would have been killed off. Simply put, some of these marine animals such as the ancestors of Congeria and Marifugia survived in a gigantic Pliocene marsh system called Lake Pannon in the Western Paratethys and eventually went underground into the caves and rivers of what is now seen as the Dinaric karsts of the eastern coastline of the Adriatic Sea. Congeria kusceri (and now its sister species) and Marifugia cavatica are thus living marine fossils, suviving underground today and, as such, are now highly endangered species as the pristine waters they live in become more and more polluted from human activities above them.
Lake Macleod is the most western lake in Western Australia and is located north of the small coastal, fishing, city of Carnarvon and over 1,000 kilometres north of the state capital Perth. Queensland in northeastern Australia is located in the wet tropics, whereas northwestern Western Australia is in the dry tropics. This part of Western Australia too is greatly influenced by the north-flowing Western Australian Current that takes cool water from Antarctica northwards so that, unlike the wet, humid and hot eastern side of the Australian continent, the flat coastal plain of Western Australia is near desert. In this arid area is huge Lake Macleod. This lake was connected at its southern end some 6,500 years ago to the Indian Ocean but became separated from it by the build up, beginning about 1,500 years ago, of a coastal barrier of sand dunes.
The lake has an area of 225,000 hectares, is 110 kilometres long, 40 kilometres wide, and lies between 3-4 metres below sea level so that Indian Ocean seawater percolates into it around its margin forming vents and seepage mounds. Very high air temperatures, combined with dryness, cause the water to evaporate creating vast, shallow, brine pans and gypsum deposits. This is mined and Dampier Salt currently produces over 1.5 million tonnes of gypsum annually. This whole area of northwestern Western Australia also produced 8.5 million tonnes of salt in 1998/1999, this being used to produce sodium hydroxide and chlorine, which are used to manufacture polyvinyl chloride resins.
Importantly, however, Lake Macleod has Australia’s largest inland community of mangroves, mainly the grey mangrove – Avicennia marina (Forssk.) Vierh. – a truly weird sight for a marine biologist used to seeing these plants fringing estuaries in the wet tropics. The lake is also an important site for migrating shorebirds and since 2000, over 70 species have been recorded from this seemingly inhospitable landscape.
The fringes of the lake comprise an open scrub that is dominated by low halophytic plants. Three of these are illustrated. Halosarcia indica (Willd.) P.G. Wilson (A) and Halosarcia pruinosa (Paulsen) P.G. Wilson © are called the brown-head and bluish samphires, respectively. The third is Muellerolimon salicornisceum (F. Muell.) Linez. (B). The plants belong to the Salicornioideae (Chenopodiaceae), which are to be found virtually world-wide high on sandy shores.
[Halosarcia indica _]occurs as a dense shrub up to two metres tall, the segments being thick, succulent, cylindrical, turning brown apically. As in all samphires, the tiny flowers occur in groups of three concealed by fleshy bracts with only a single stamen and two lobes of the stigmata projecting above each one.[ Halosarcia pruinosa _]is a more delicate plant, the segments, though succulent, are spherical arranged like rows of bluish-green beads. [_Muellerolimon salicornisceum _]is darker green and has long, thin, segments the rows of which end in a sharp point. These three halophytes occur in coastal flats throughout northwestern Western Australia and where salinities of standing water flooding them on spring tides may rise to between 40%o and, in the blistering heat of summer, an incredible 90%o – three times that of normal seawater.
In August 2000, a group of Australian and overseas scientists congregated within a field laboratory at Dampier on the Burrup Peninsula itself on the northwest coast of Western Australia. One day, the divers brought into the laboratory, in a bucket, a dumpling squid, also known as the striped pyjama squid, [_Sepioloidea lineolata _]Quoy et Gaimard, 1832. This somewhat surprising little cuttlefish is native to the southern Indo-West Pacific occurring off eastern, southern and Western Australia. Although called a squid, it is actually a cuttlefish and hence quite different from the truely pelagic, swimming, squids. This little squid is little bigger than 50 millimetres in overall size and, in an aquarium, sat contentedly on the sand provided eyeing me out of its slitty horizontal eyes, the upper lid of which is bright orange. Normally, the species is found on sand and amongst seagrasses in waters up to 20 metres in depth. It also tends to bury itself in sand so that only the top of its head is visible and from where it emerges to hunt shrimps, crabs and the like over the seabed catching them with the longest two of its ten arms.
The squid is round, hence one of its colloquial names but it is also striped cream and brown, hence its other common name. The body is longitudinally striped, the head vertically so, because of the way the animal sits. Above each optic lobe arises a row of finger-like projections, also giving it the appearance of possessing bushy, Dennis Healey (born 1917 and former British Member of Parliament) – like, eyebrows. The arms are short and webbed. Its lateral fins are translucent and sitting there, apparently asleep, I could not resist drawing it. This little tropical cuttlefish must be deserving of the description ‘cute’ but its bite, unknown to me then, is venomous just like the earlier-described blue ringed octopus Hapalochlaena maculosa (Hoyle, 1883), also discovered in Western Australia.
Franklin et al. (2012) investigated mating in the southern dumpling squid, [_Euprymna tasmanica _]Pfeffer, 1884. In this species, copulation between a male who has enchanted, ensnared and restrained a female in his ten arms, may last for three hours. During this process, the male, who has grabbed his partner from underneath, transfers sperm-filled spermatophores using one of his modified longer arms – the hectocotylous – from his own vas deferens to the oviduct of the female. Such a prolonged bout of copulation halves the swimming endurance of both sexes and they take up to 30 minutes to recover. But, this is not all. Sexually mature for just a few months, males spend much of their short lives iniating and engaging in multiple mating sessions with as many female partners as possible. Such a strategy ensures that his genes are transferred to multiple offspring so that the survival of this short-lived cephalopod, as a species, is guaranteed. Tiring work though. And I am glad our cute but clearly promiscuous little animal was eventually returned to the sea to continue with his amorous adventures.
The rocky beaches surrounding Watering Cove on the Burrup Peninsula at Dampier on the northwest coast of Western Australia occur two chitons or coat-of-mail shells. Low on the shore at Watering Cove was the[_ ]aspinose[ ]chiton[ Acanthopleura. gemmata ](Blainville, 1825). Higher up the shore and, thus, occupying the mid eulittoral, was the highly spinose chiton [_Acanthopleura spinosa _](Bruguière, 1792). Both of these chitons are large reaching a body length of up to 90 millimetres and both species are widely distributed on appropriate habitats throughout the tropical Indo-West Pacific. It seemed self-evident that the long spines of the more exposed _Acanthopleura [_spinosa _]had a defensive role in protecting it, say, from predation by crabs and gulls.
Frequently, however, anatomical adaptations serve more than one function and the decision was made to test the hypothesis that the long spines of [_Acanthopleura spinosa _]also facilitate convective cooling. This is because the black, heat absorbing, surfaces such as occur along the Burrup Peninsula often reach surface temperatures greater than 40oC. In a makeshift laboratory, an experiment was therefore set up to compare the temperature tolerances of the two chiton species. This involved allowing individual chitons of both species to attach to petri-dishes with temperature probe terminals situated beneath them. Once settled, the chitons were subjected to radiant energy for a standard period of time from a powerful overhead lamp set at a standard distance from them, while beneath-foot and ambient air temperatures were recorded.
The study provided evidence that the long, black spines of [Acanthopleura spinosa _]did indeed facilitate convective cooling. That is, _Acanthopleura spinosa warmed more slowly than the spineless Acanthopleura gemmata. With the spines of [_Acanthopleura spinosa _]clipped off with nail scissors, moreover, the two species warmed at approximately the same rate. When intact and spineless [_Acanthopleura spinosa _]were compared, however, the foot of the latter attained a mean temperature of 2.4°C above that of spinose individuals. Clearly the spines were functioning as radiant heat emitters (Britton and Morton 2003).
The simplest way to minimise thermal stress in a difficult, tropical, environment is to seek shelter during daytime periods of emersion and exposure to the sun and employ evaporative cooling by raising the mantle margin from the hot rock surfaces, in the same way that the Saharan desert’s long-footed lizard, Acanthodactylus senegalensis[* *]Chabanaud, 1918, lifts its feet alternately from the hot sand to cool them. The long black spines of Acanthopleura spinosa, however,[_ _]provide it with an extra adaptation, which is to reduce the effects of thermal stress yet further using its spines as thermal radiators. Black body structures, such as feathers and spines are more efficient at reducing thermal loading than if such structures were white. It is true that a substantial fraction of incident short wave radiation is reflected from a light-coloured structure. Black body structures, on the other hand, absorb large fractions of incident solar radiation, but it remains near the surface and is easily removed by either re-radiation or convection. Hence, the black spines of [_Acanthopleura spinosa _]seem to function similarly, minimising the thermal loading to its body. This allows it to occupy a higher position on the shore than [_Acanthopleura gemmata _]and is thus able to exploit an exclusive source of food, that is, the high-zoned micro-algae and lichens.
The spines of Acanthopleura spinosa, however, probably[_ _]also function, as protection from predation, for example, from more terrestrially-adapted crabs and gulls but this was not tested. Convective cooling, the removal of heat from a body by a flowing fluid, is a thermoregulatory mechanism exhibited by a wide variety of organisms from a broad suite of habitats especially in the tropics.
Many phylogenies of the Gastropoda have evolved predatory life styles and this has become an abiding interest. Representatives of three such families are illustrated. On the large expanse of intertidal sand and mud exposed within Princess Royal Harbour at Albany, Western Australia, the buccinid Cominella eburnea (Reeve, 1846) edge attacks the shallow burrowing venerid bivalve [Katelysia scalarina _](Lamarck, 1818). This is illustrated in A and was drawn in October 2003. The gastropod positions itself parallel to the ventral margin of its potential prey and waits patiently until the bivalve is forced to part its shell valves slightly in order to take new amounts of refreshing water into its mantle cavity. When this occurs, _Cominella eburnea inserts its proboscis slowly and deliberately into the slightly parted shell valves and slowly extends it into the body of the prey. At the tip of the proboscis is a radula, which is then used to scrape and ingest the bivalve’s tissues (Morton 2006).
The moon shell[_ Natica gualteriana_] (Recluz, 1844) was observed drilling the burrowing venerid bivalve Marcia ceylonensis (Lamarck, 1818) on a sand flat at Langkawi, Malaysia in February 2004. The pair of molluscs is illustrated in B. Representatives of the Naticidae embrace their prey, typically bivalves, but sometimes other taxa, with the enlarged lobes of the mantle and position their mouth typically above that part of the prey’s shell where the most nutritious tissues lie beneath. They then use the radula at the tip of the mouth plus secretions from glands in the gut complex to drill a characteristically countersunk hole in the bivalve’s shell. The position of the hole is characteristic for each species of prey attacked. That is, each prey species is manipulated, held and drilled in a particular way, as has been demonstrated for other tropical, Hong Kong, representatives of the Naticidae (Ansell and Morton 1985, 1987) but never for Natica gualteriana.
The representatives of the Muricidae, unlike the Buccinidae and Naticidae, which largely capture their prey on sand flats, are primarily predators of rocky intertidal and subtidal habitats. Illustrated is Lepsiella (Bedeva) hanleyi (Crosse, 1864) drilling the mussel [Mytilus galloprovincialis _]along the banks of the Swan River in Perth, Western Australia in February 2004. The pair of molluscs is illustrated in C. Of all the predatory gastropods, the Muricidae are the most well-studied and are widely regarded as keystone predators on rocky shores worldwide. That is, by virtue of their predatory habits, they regulate the population structures of their prey species and thus the ecology of the habitat in total. In the illustrated example of the predator and its prey, an individual of _Lepsiella (Bedeva) [_hanleyi _]has drilled a large individual of [_Mytilus galloprovincialis. _]In this case, however, the drill hole is not countersunk, as is typical of naticid predators, but is smaller and straight-sided (Morton 2012a). Muricids, like naticids, however, do use the radula and secretions from glands in the proboscis to not just etch the hole, but possibly also inject a paralysing chemical into the prey.
Ever since my days, as a post-doctoral fellow between Ph.D. and leaving for Hong Kong, at the Hayling Island Marine Laboratory of the University of Portsmouth, held an interest in oysters, both scientifically and culinary. They are a difficult group to work on because to the untrained eye many oysters look very similar. They generally attach by their left shell valve to the habitats they either occupy naturally or to the artificial cultch that oystermen put out in the sea to cultivate them. The European flat oyster Ostrea edulis Linnaeus, 1758 is round with a shallowly-cupped left valve. Since its near demise in the 19th and 20th centuries in Europe, it has been largely replaced in oyster farms by the Pacific oyster Crassostrea gigas (Thünberg, 1793). Wherever it has been introduced, it has become accepted dogma that this is the name of all populations, everywhere. A post-doctoral student, Dr Katherine Lam at the Swire Institute of Marine Science of the University of Hong Kong, however, examined the Hong Kong oysters of that name and cultivated in Deep Bay for centuries, possibly thousands of years, using mitochondrial DNA analyses. Surprisingly, the so-called Pacific oyster, Crassostrea gigas, cultivated in Hong Kong, was not that all, but a new species which we named Crassostrea hongkongensis (Lam and Morton 2003). Using the same mitochondrial DNA analyses too on oysters from farms in Albany, Western Australia, we showed, rather surprisingly, that the European flat oyster, Ostrea edulis, had been introduced there too and never detected before (Morton et al. 2003). We also produced papers identifying and illustrating the Hong Kong oysters (Lam and Morton 2004) and those from Malaysia and Singapore (Lam and Morton 2009).
Despite them being an important research area, none of these oysters was ever drawn, because of their superficial similarity to each other and the wide variation that each one demonstrates in the various components of their ranges. Except one – the leaf oyster [Dendostrea folium _](Linnaeus, 1758). This species occurs in Hong Kong but specimens were obtained from Langkawi during a research trip to Malaysia in 2004. The species occurs subtidally at shallow depths and attaches to other shells and objects and specimens were obtained from inshore fishermen by the simple expedient of asking if I could examine their fishery discards. From here, thereby I obtained the illustrated oyster attached to the empty shells of the mussel _Modiolus micropterus Deshayes, 1836
The remarkable characteristic of Dendostrea folium is that, firstly, its shell adopts the form of the object it settles onto – a biogical phenomenon called xenomorphism. Secondly, however, unlike other oysters which simply cement themselves to an object, Dendostrea folium produces little finger-like extensions from both sides of its lower shell valve so that it appears to be clutching, hand-like, its mussel substratum. This is so weird that one is left with the indelible impression of a hand, somewhat like that of an inanimate alien creature hanging on to life.
Consider the case of the oyster,
Which passes its time in the moisture:
Of sex alternate,
It chases no mate,
But lives in a self-contained cloister.
(Jerome Tichenor [a pseudonym of Joel Hedgpeth, 1911-2006])
In September-October 2004, a research visit was made to the Institute of Oceanography and Fisheries in Split, Croatia. The aim was to research the Mediterranean muricid gastropod [Hexaplex trunculus _](Linnaeus, 1758)[. _]Though widespread and common in the Mediterranean, actually little was known of this gastropod’s behaviour beyond the presumption, that, being a member of the Muricidae, which are all predators, it was too. [_Hexaplex trunculus _]is a large animal, the shell being up to 10 centimetres tall and sharply angulate, often with spines of variable size. The animal is a commercial fishery resource too, especially in the Adriatic Sea, so that there was another purpose for the research. If one wishes to cultivate it, then it is important to understand its dietary preferences. The snail is also interesting in that its hypobranchial gland is the source of the purple-blue dye, Royal blue, much prized by the ancient Phoenicians.
My host, Professor Melita Peharda, and I made the trip south from Split to Dubrovnik where the famous Croatian marine laboratory located in the walls of this ancient port is located. Just north of Dubrovnik is Mali Ston Bay where the Mediterranean mussel, [Mytilus galloprovincialis _]Lamarck, 1819[, ]was being cultivated on ropes hanging down in the water from rafts. Divers also collected us two other large bivalves, namely _Arca noae Linnaeus, 1758 and [_Modiolus barbatus _](Linnaeus, 1758) nearby plus [_Hexaplex trunculus _]individuals.
On return to Split, the [Hexaplex trunculus _]individuals were weighed and measured[ _]as were the three bivalves and the predators then established in aquaria with a choice of the three bivalves to attack plus individual prey items of different sizes. It transpired that only [_Mytilus galloprovincialis _]was attacked (at least in significant numbers) and this research was published a year or so later (Peharda and Morton 2006).
In the summer of 2005, Professor Peharda made the return trip to England and, at the University of Cambridge, the shells of [Mytilus galloprovincialis _]predated by the [_Hexaplex trunculus _]individuals were re-examined using Scanning Electron Microscopy (SEM), with Dr E.M. Harper. This research too was published (Morton _et al. 2007) and showed that [Hexaplex trunculus _]uses both the tooth-like projections on the outer lip of its shell to chip the posterior valve margins of its [_M. galloprovincialis _]prey. It can, however, like many other muricids also drill the shells of its bivalve prey. Once any kind of access is achieved, the predator pushes its proboscis between the broken edges of the prey’s shell and ingests its tissues. The illustration on the left shows _Hexaplex trunculus with a detail of the spinose shell margin (A). The experimental prey species Arca noae, Modiolus barbatus and Mytilus galloprovincialis are also drawn in B, C and D, respectively, with a detail of the chipped posterior shell margin of the latter species also shown (arrow).
Moreton Bay, in Queensland, Australia, is protected from the mighty waves of the Pacific Ocean by North Stradbroke Island. The wide expanse of protected seagrass-covered sand flats of Myora Springs, within the bay, was investigated in 2005.[_ ]Here, there occur byssally-attached clumps of the mussel [_Trichomya hirsuta _](Lamarck, 1819),[ ]each of which had an empty oyster shell, [_Saccostrea glomerata _](Gould, 1850), a valve of [_Pinctada fucata _](Gould, 1850), or [_Isognomon ephippium _](Linnaeus, 1758), or that of a conspecific, as their basis. The mussel clump so attached comprised more than thirty individuals creating a, byssally-bound, compact mass. The bowl of mud in which each mussel clump sat was not anaerobically black, but clearly aerated and when turned over revealed a little maze of galleries, the most obvious residents of which were the galeommatid bivalves[ Scintillona cryptozoica _](Hedley, 1917) and [_Scintillona daviei _](Morton, 2008). There were other inhabitants of the galleries but the most ubiquitous were a pair of little hairy crabs, [_Pilumnopeus serratifrons _](Kinahan, 1856). It is this crab that creates the galleries in the mussel clumps, and it is to the roofs of these that the two bivalves were attached.
The small bivalve Scintillona cryptozoica occurred in family groups of up to 25 individuals and each one has a distinctive array of apically deep red tentacles and papillae adorning the mantle folds that are reflected to mostly cover the shell, except apically. The mantle edge is a light red around the exhalant siphon and yellow around the inhalant aperture. There are antero- and postero-dorsal tentacles above the inhalant aperture and exhalant siphon, respectively. In addition, there are two pairs of other tentacles anteriorly and four pairs posteriorly. Smaller papillae adorn the reflected mantle fold. An even smaller bivalve, less than 7.5 mm in shell length, often occurred with Scintillona cryptozoica, also in small family groups of up to seven individuals. This turned out to be a species new to science and which was named in honour of Peter Davie of the Queensland Museum. Scintillona daviei _ also has a distinctive array of apically deep purple-red tentacles and papillae adorning the mantle folds that are reflected over most of the shell. The mantle margin is dark red around the exhalant siphon and yellow/orange around the inhalant aperture. There are antero- and postero-dorsal tentacles above the inhalant aperture and exhalant siphon, respectively. There are two pairs of other tentacles anteriorly and one pair posteriorly. Smaller papillae adorn the outer surface of the reflected middle mantle fold. Hence, it was very similar to _Scintillona cryptozoica, except that, in addition to being much smaller, had a purple dorsal shell region and had a distinctive red pigment spot on the foot.
All the bigger individuals of both species were females; some brooding fertilised eggs and larvae inside their ctenidia. Smaller individuals of, again, both species were males and in two cases, two tiny such juveniles were identified inside the females. The males of the two [_Scintillona _]species are thus not strictly dwarf males but probably join a clump of adults and become male if an established, maturing, female is also resident. In the absence of a female, the newly settled individual may become a putative female. This results in a surfeit of either putative or intersex females, with each cluster being optimally dominated by one or two mature, possibly gravid or brooding, females. With completion of their life spans and, as the oldest individuals die, younger females can then assume their places in the cluster hierarchy. Such a pattern of reproduction is called protandric consecutive hermaphroditism.
It was also interesting to speculate upon the nature of the relationship between the two bivalves and their host [_Pilumnopeus serratifrons. _]Not every clump of bivalves comprised both species. Hence, their relationship is not absolute and thus not obligate. The associations between the two bivalves with [_Pilumnopeus serratifrons _]did, however, appear obligate, that is, they were never recorded from clumps of [_Trichomya hirsuta _]without the crab. Clearly, therefore, the bivalves benefit from the protection, that is, aegism, afforded by living in a gallery underneath a [_Trichomya _]clump. Moreover, the crabs, by virtue of their activities, must aerate their gallery and, in so doing, bring in suspended particulates that the bivalves can exploit and collect for their own nourishment. Such an associations is called commensalism, the bivalves benefiting nutritionally from living with the crab, but not doing it any harm by depriving it of food (Morton 2008a).
On the highly exposed shore of Blue Lake Beach on North Stradbroke Island, Queensland, Australia, the naticid moon-shell Polinices incei (Philippi, 1873) surfs up the beach to attack juvenile, tidally migrating, surf clams, Donax deltoides Lamarck, 1818. The moon-shell moves shallowly within the drying sand at the top of the beach, its presence being detected by a raised trail. The surf clams apparently recognise the presence Polinices incei in their vicinity as they, unusually, emerge from the sand and attempt to escape from the predator by leaping. If captured, however, up to three individuals of the surf clam juveniles are held by Polinices incei beneath the posterior margin of its foot. Polinices incei then retreats down-shore where it drills a hole in its bivalve prey[_ _]and consumes the contained flesh
To move up shore and return down shore, Polinices incei surfs the swash and backwash, respectively. Surfing involves rapid inflation of the foot and propodium such that the moon shell floats upside down and is carried by the waves either up or down. Surfing down shore, Polinices incei was often observed holding bivalve prey. Downward migrating moon-shells[_ _]also roll in the backwash, but brake by digging in the anterior end of the foot and resume burrowing rapidly. Other identified behaviours include hunting and galloping. The former is undertaken, since the moon shell is blind, by detecting the movements made by its bivalve prey in the sand. The latter’s galloping behaviour is used to chase its leaping, desperately trying to escape, prey (Morton 2008b).
As far as is known, Polinices incei is the only known naticid predator that has evolved a surfing behaviour, which enables it to attack its prey, the juveniles of Donax deltoides, on highly exposed beaches in eastern Australia. The evolution of naticid drilling predation was part of the Mesozoic Marine Revolution (Vermeij 1977). As predicted by escalation theory, the dangers to infaunal bivalve residents from naticid predation seem to have increased over time but since shallow water benthic assemblages are thought to be particularly susceptible to mass extinction events it seems probable that the Polinices incei _](predator)/[_Donax deltoides (prey) relationship is relatively modern.
The relationship also gives us an insight, however, into the evolutionary arms race that evolves between predators and their prey. In this case, surf clams have evolved to keep well out of the reach of possible predators by migrating up and down with the tides. When the tide is high, they are located high on the shore well away from the reaches of predatory fishes for example. Polinices incei seems to have evolved a behavioural way of attacking them at these times, however, by surfing up the beach with the bivalves and attacking them in what was hitherto believed to be a refuge from predation.
Until this research was undertaken, surf clams have always been regarded as occupying a ‘safe refuge’ niche, at least from marine predators. The relationship is, therefore, a remarkable, and hitherto unrealised, example of adaptive radiation by this family of moon-shell predators.
Cuba has been described by[_ ]González Guillén (2008) as a land snail paradise. Of especial interest throughout the Caribbean is the land snail genus _Cerion. Cerion is from the Greek word kerion, signifying honeycomb, and is given to these shells because their form resembles that of a beehive. They were, hence, at one time known as beehive shells. Species of the Cerionidae are air-breathing pulmonates. That is, they possess a lung instead of the aquatic gastropod’s gills. They are endemic to the islands of the tropical Caribbean. Their distribution ranges from the barrier islands and keys of southern Florida, throughout the Bahamas, Greater Antilles, Cayman Islands, western Virgin Islands, and the Dutch Antilles, but are absent from Jamaica, the Lesser Antilles, and coastal Central and South America. Species of Cerion live beneath coastal vegetation, usually within a few hundred metres of the shore where salt spray reaches them. The illustrated Cerion salvatori Pilsbry, 1927 was collected from precisely this habitat, amongst the coastal sand dune plants at Varadero on the southern coastline of Cuba in March 2006. Picked up, they appeared to be empty shells. Once put in water for an hour or so, however they came to life and began crawling. Presumably, therefore, they spend much time in a state of dormancy, only being resuscitated with the arrival of rain or sea spray.
Cerion salvatori has a shell length of up to 32 millimetres and an above aperture diameter of 12 millimetres. Within the aperture is a single tooth. The species was originally described by the American malacologist Henry Augustus Pilsbry (1862-1957) who, over his long life, published more than 3,000 research papers, mostly from his office in the Academy of Natural Sciences of Philadelphia. Much of the early work on the Cerionidae was thus primarily taxonomic, and resulted in the description of some 600 nominal species and over twenty genera and subgenera. In this work, Pilsbry was a pioneer and described Cerion salvatori (Pilsbry 1927, plate 1, figures 11 & 12).
Species of Cerion live in dense but patchy populations, often containing over ten thousand individuals, typically living in coastal leaf litter. Individual populations tend to be fairly uniform in the size and morphology of their shells. Clench (1938) noted that the greater the isolation of the population, the greater the uniformity among its members. Variation in shell morphology among populations can be enormous, not only throughout the geographic range of Cerion, but even among neighbouring populations separated by less than 100 metres. This extraordinary yet geographically circumscribed diversity has led to an extensive body of literature dealing with the Cerionidae.
As recorded by[_ ]González Guillén (2008), the greatest diversity of _Cerion occurs on Cuba and in the islands of the Bahamas. Modern researchers have, however, postulated that perhaps only 10-20% of the species named within the Cerionidae will eventually be found to constitute valid species. The remainder may, also eventually, be considered to be subspecies, ecophenotypes or merely distinctive combinations of alleles. Gould and Paull (1978), for example, reduced the eleven described species of Cerion from Hispaniola to the Virgin Islands to a single one using multivariate analyses of shell characters.
In the Açores, Portugal, the steeply descending continental shelf is characterised at different depths by two, endobenthic, suspension feeding species: the shallower-living (0-~100 metres) bivalve [Ervilia castanea _](Montagu, 1803)[ ]and the deeper-residing (~100–250 metres)[ ]serpulid polychaete [_Ditrupa arietina _](O.F. Müller, 1776)[.] As a dominant member of the continental shelf fauna, _Ditrupa arietina provides a habitat for a number of epibiont species that attach to its tubes anteriorly. These include the cemented, introduced, serpulid [Hydroides elegans _](Haswell, 1883), three species of Foraminifera and a number of species of, mostly unidentifiable, bryozoans. Tubes of _Ditrupa arietina are also drilled and, hence, probably predated by the naticid prosobranch gastropod [_Natica adansoni _]Blainville, 1825.
Ditrupa arietina lives for ~2 years with most growth occurring during the first year and with sexual maturity also occurring in the first year of life [_ ]so that, post reproduction and death, its tube becomes available for secondary colonisation by the sipunculan worm [_Aspidosiphon muelleri _]Diesing, 1851[ ]At this time, anterior epibionts die, because the sipunculan orientates the tube anterior end down, but further epibionts can now colonise the posterior end of the tube. With time and wear, the tube slowly degenerates. Throughout its life history, therefore, the tube of _Ditrupa arietina functions as an inhabitable substratum and is, thus, a locally important Açorean habitat for a suite of other epibenthic species.
The suggested life history of the tube of Ditrupa arietina was described and illustrated by Morton and Salvador (2008)[. _]A living individual of _Ditrupa arietina (A) has a cylindrical, slightly curved tube that resembles an elephant’s tusk or, more appropriately, a molluscan scaphopod, and is anteriorly swollen although the orifice typically narrows again at its anterior extremity. There are numerous constrictions, or flanges, to the shell that is also variably patterned with circlets of orange-brown pigmentation. Because the contained worm is bright red, the tubes of living individuals are also redder than their empty counterparts. The nineteen branchial filaments are also bright red (sometimes red banded) and the operculum is a membranous cup or funnel closed distally by a flat, brownish, chitinous plate thickened in the centre to form a boss.
Living individuals of Ditrupa arietina (A1) live in the sediment, posterior end down and with the swollen anterior end situated above the surface. The anterior end of the tube is often encrusted with epibionts (B). As described above, drill holes in the tube are probably made by a naticid gastropod (Morton and Harper 2008) and is illustrated in C. Following death, either from drilling predation or other causes, the tube of [Ditrupa arietina _]is occupied[ ]by _Aspidosiphon muelleri (D) at which time the anterior epibionts also die because the sipunculan lives head down in the sediment (D1). With the posterior end of the tube now projecting above the sediment surface, it too becomes encrusted by epibionts (E), such that the tube becomes more eroded with age (F) until only fragments remain (G).
The tube of [_Ditrupa arietina _]thus has a life history that is important for the survival of not only the original polychaete inhabitant, but also the subsequent occupant of the empty tube, the sipunculan [_Aspidosiphon muelleri _]and a suite of encrusting animals that occupy in turn the anterior and then the posterior ends of the tube until it fragments.
There are ten species of mackerel that occur in British waters but by far the most common is the Atlantic mackerel Scomber scombrus. Linnaeus, 1758. My first memory of this fish was as a young boy in the 1950’s when I used to go fishing on the piers flanking the entrance to the River Arun in my home town of Littlehampton. One early summer day, I and a few other lads were fishing when a huge shoal of Atlantic mackerel came into the river either being chased or, more likely, themselves chasing a school of sprat. There was so many, the river’s waters were literally boiling with them and dashing down Nelson’s Steps to reach their level, we were literally able to scoop them out of the water using our dip nets.
The mackerel is beautiful. It is fusiform, or torpedo-shaped, and a brilliant green-blue above and silver below. The head is a steely blue-black with a small yellow patch behind the eye. There are between 23-33 dark wavy bands across the back of the fish down to the midline. Because there is no swim bladder, the flesh is oily, but the animal has to swim continuously in order to ramjet aerate the gills
[_Scomber scombrus _]over-winters in deeper waters where it does not feed but comes inshore during the early summer months when sea temperatures are rising to between 11ºC to 14ºC, and where it feeds voraciously on other shoaling fish, such as the European sprat, [_Sprattus sprattus _](Linnaeus, 1758). Each female may produce up to 300,000 eggs and the newly hatched fry, between 2-3 mm long feed on planktonic copepods. By the end of the first year of life, the young mackerel are between 15-20 cm long. The fish is also cannibalistic and so each shoal usually comprises a single cohort of individuals all approximately the same size. The species matures at an age of about three years but can live for twenty years and reach a maximum length of almost half a metre. Like many other species, the Atlantic mackerel was over-fished in the 1960’s especially in northeastern Atlantic waters, but recovered substantially up to 2008 because the stock was well managed by a group of EU coastal states and Norway who set a quota of 600,000 tonnes and divided it up between them. Recently, however, the Icelandic and Faeroese governments have abandoned quota agreements designed to protect stocks and their fleets are targeting the migrating mackerel specifically. These same two ‘countries’ have already, virtually unilaterally, driven the blue whiting ([_Micromesistius poutassou _][Risso, 1827]) to extinction such that stocks of this species have collapsed and, now, they are intent on doing the same with mackerel.
In 2011, I wrote an editorial for Marine Pollution Bulletin about this situation (Morton 2011c). In 2011, the main Faeroese company, Thor Offshore and Fisheries, which already has six trawlers in the mackerel grounds, is bringing in another factory ship to add to its fleet. Another Faeroese company, Vardin, will add three more industrial-scale trawlers to the growing fleet that will target the mackerel in the North Atlantic. The catches were expected to be record-breaking in 2012, with Icelandic vessels initially setting a quota for themselves of 147,000 tonnes but expecting to take 155,000 tonnes and Faeroese trawlers another 150,000 tonnes. The actual total catch in 2012 was 930,000 tonnes with the two tiny island ‘countries’ accounting for a third of this. As a consequence, the European Union has banned all Icelandic and Faeroese mackerel fishing vessels from its waters, but there is little else that can be done to protect the species in non-EU waters. But there is another aspect to this story. Because the Icelandic and Faroese governments have unilaterally abandoned quotas, other fleets from Russia, the Far East and China have felt free to move into North Atlantic waters in pursuit of the mackerel. It is estimated that there are currently twenty ‘super-trawlers’ working these waters including the Hong Kong-controlled Lafayette, which is currently processing 1,500 tonnes of mackerel daily, mainly for the Chinese market. Even so, the mackerel fishery is still important to British, notably Cornish and Scottish, fishermen and is estimated to be worth £135 million (US$ 220 million) annually.
‘They lie in parallel rows,
on ice, head to tail,
each a foot of luminosity
barred with black bands’
Many predatory gastropods detect their prey by chemoreception. Nassariid gastropods do not generally occupy high energy, wave-exposed, beaches, chemical cues emanating from stranded carrion being better detected on long, gently sloping, beaches under conditions of low exposure to wave action. A remarkable exception to this generalisation occurs on the wave exposed shores of South America, south and west Africa and the Indian Ocean (Brown 1982). On the highly wave-exposed sandy beaches of South Africa, facing the swell of the Indian Ocean, the plough-snail [Bullia rhodostoma _](Reeve, 1847)[ ]‘surfs’ or swash-rides up such shores in search of carrion, for example, stranded jellyfish and pelagic hydrozoans (Odendaal _et al. 1992).
If one walks on such beaches, such as that of Wilderness Bay in the Western Cape of South Africa, they appear empty, few animals and certainly no plants being able to tolerate the continuous onslaught of the pounding waves. If, in particular, however, one of the pelagic colonial hydrozoans such as the Portuguese-man-of-war, Physalia physalis (Linnaeus, 1758), or the by-the-wind-sailor, Velella velella (Linnaeus, 1758), both bottle blue, are washed ashore then they are quickly attacked by plough-snails, most commonly Bullia rhodostma. Illustrated (in A) is an individual of Velella velella that has been stranded and been detected by the first snail to arrive at its doomed body. Already the snail has its radula-tipped proboscis everted in preparation to feed. Other individuals appear in large numbers, as if miraculously, from their repose in the sand. They do this by ‘galloping’ up-shore to the food such that it is quickly engulfed in a mass of feasting individuals.
[Bullia rhodostoma _]lacks eyes, instead it has an acute sense of smell, responding rapidly to amino acids in the water (Hodgson and Brown 1985) although exactly how food can be located even by this means is unknown because any chemical gradient that nassariids are thought to need to successfully arrive at must be destroyed by the turbulence created by pounding wave action. Notwithstanding, arrive they do and in hordes. Another remarkable feature of the plough-snails is that _Physalia physalis possesses extremely powerful stinging cells, or nematocysts, in its brilliant blue tentacles. These are not, however, deactivated upon the stranding of the animal, and, hence, [Bullia rhodostoma _]must have structures or produce secretions in its mouth that protect it from the stinging cells. Within minutes of being stranded, all that remains of a Portuguese-man-of-war[ _]is its float, which unless it is deflated and eaten in the attack, drifts away, and the snails depart the scene too.
They do this, to escape the desiccating heat of the drying upper shore by surfing down it. The foot is inflated like a parachute (illustrated in B) and this catches the backwash of the receding wave train and they are thereby transported back to the cool, regularly moistened, sand. Here they disperse, there being a positive advantage to not remaining together but spreading numbers out far and wide so as to maximise the chances of obtaining a meal of randomly occurring carrion both spatially and temporally.
The gilthead seabream Sparus aurata (Linnaeus, 1758)[* *]occurs along the eastern coast of the North Atlantic and the Mediterranean and grows to a length of up to 60 centimetres. It is either solitary or occurs in small groups and is, like most breams (Sparidae), a predator feeding on small crustaceans, mussels and even oysters that it crushes in powerful jaws equipped with between four to six teeth in the front of the mouth followed by two rows of teeth to the sides. The fish is a dull grey overall but there are long shadowy stripes dorsally and a dark black blotch on the upper edge of the operculum and marking the origin of the lateral line system. There is a characteristic golden band between the eyes and which gives the species its English name
Sparus aurata is either caught wild or raised in fish farms, especially in Spain where it is called dorada. It has a similar name dourada in Portugal and is commonly sold in restaurants and fish markets in the Algarve, which is where it was first encountered and drawn.
In the wild, the species is euryhaline and eurythermal, that is, it can tolerate wide variations in salinity and temperature. This is because juveniles migrate into coastal waters in early spring to feed and grow. In autumn, they return to the open sea where they mature. The species is also a protandric hermaphrodite, that is, juveniles mature first into a male at two years of age and a body length of between 20-30 centimetres. Once they have spawned, a change in sex occurs at between two to three years and a length of between 33-40 centimetres and they develop into females. These females are prolific, producing, if conditions are good, between 20,000 to 80,000 eggs each day for a period of up to four months.
This helps account for its success in the wild, but also the success that has been achieved in rearing this species artificially. Catches of wild Sparus aurata have always been relatively modest, that is, around 9,600 tonnes in 2009, primarily from the Mediterranean. Traditionally, Sparus aurata was cultured in natural coastal lagoons and saltwater ponds. To improve on this traditional industry, intensive rearing systems were developed during the 1980s, when it began to be cultured in either land-based ponds or in cages moored in the sea. Subsequently, Sparus aurata has become an important mariculture species.. Today, it is cultured in most of the countries bordering the Mediterranean and its production reached 87,000 tonnes in 2000, 130,000 tonnes in 2007 and 140,000 tonnes in 2010, thus dwarfing the capture fisheries figures.
Because it is possible to hatchery rear the species from fertilised egg all the way through to adulthood, this species is a significant success story and, theoretically, therefore, the industry could be expanded dramatically. So successful, in fact, is cultivation that the industry based around it is today a high volume, low profit margin, business with Greece being by far the largest producer (50%), followed by Turkey and Spain (~15% each), and Italy (5%). Other countries outside the Mediterranean region are also producing Sparus aurata and it is fast becoming a fish staple in the diets of many. It can today be purchased daily from British fishmongers.
The first botanical name assigned to the sea grape (Polygonaceae) was by Hans Sloane in 1696, who called it Prunus maritima racemosa reflecting the notion of a coastal prune. Linnaeus called it [Polygonum uvifera _]in the 1753 edition of his [_Species Plantarum. _]Subsequently, in 1756, Patrick Browne changed the generic name to _Coccoloba (red-leaf) and in the second (1762) edition of his [Species Plantarum. _]Linnaeus changed the classification to _Coccolobus uvifera.
The sea grape was first encountered on the beaches of southern Texas while researching for the book the Coastal Ecology of the Gulf of Mexico (Britton and Morton 1989). The plant is a sprawling evergreen bush or small tree, with a smooth yellowish bark, that reaches a maximum height of around eight metres, but most plants are little more than two metres tall. It is naturally found close to coastline beaches throughout tropical America and the Caribbean.
Because the sea grape is highly tolerant of salt spray and resistant to wind it is often used to stabilise beach edges and is planted as an ornamental shrub. As a consequence, it has been introduced far outside its natural range. On the waterfront of the port of San Sebastian on La Gomera in the Canaries it is planted in an ornamental row but such is the strength of the wind here that the plants are stunted, wind-wasted and finding it hard to survive. Conversely, along the more sheltered promenade of Santa Cruz on La Palmas, also in the Canaries, the plants grow taller and have been topiaried. But in its natural range, such as on Jamaica, where this illustration was drawn, the plants reach their full magnificent height and form. Here too, one can discover its method of reproduction, uninhibited by stunting and topiarization.
In Jamaica, the sea grape forms large shade trees along the coastal edge and each plant is characterised by large, round, leathery leaves up to 25 centimetres in diameter and close to where the flower stalks (and eventually fruits) are born. The leaves occur in pairs, so that the impression is given of elephant’s ears up to half a metre in width. Each leaf has a red primary vein extending from its base and these radiate outwards along its length, sub-dividing towards the outer edge. The leaf turns red as it ages. In Jamaica, I could not understand why some trees bore masses of fruits and others did not. It was eventually figured out that some plants are males with thin straggling flower stalks whereas others are female that eventually produce the grape-like bunches of fruits (A). Each tiny, whitish, flower (B) has three anthers and eight stamens. Pollen is transferred from the male plants to the females by insects, typically bees. Eventually, the flower stalks of the males die but the female trees grow and bear long, grape-like, bunches of green fruits that gradually turn to orange, then red and finally purple as they mature at about two centimetres in diameter (C-E). When cut open (F), each fruit contains a large stone that makes up most of its fabric.
In Jamaica, the sparse flesh around each ripe fruit is edible and can either be used for jam or eaten right off the tree. This was tried, but it was very bitter. Putting the seeds in the sea, it was discovered that they can float and it seems likely that this is the method of dispersal throughout tropical central America and, now, possibly elsewhere. They are unlikely, however, to have floated acrooss the Atlantic to colonise the Macaronesian beaches of the Canaries – the plants here being clearly planted.
The island of Lanzarote and neighbouring Fuerteventura are the oldest members of the volcanic Canarian archipelago and arose from the sea some 20 million years ago. In the northeastern-most end of the island sits the volcano La Corona that was created between 2,000-3,000 years ago. When it erupted it created one of Lanzarote’s most impressive geological oddities – the lava tunnel of the Malpais de la Corona volcano. With a length of 8,400 metres, this is one of the largest larva tunnels on Earth and extends from a hidden fissure close to the volcano’s apex to 1,400 metres offshore and a depth of 50 metres. The tunnel was formed when liquid larva from the erupting volcano flowed underneath a cooling surface larva field. As the flow of larva decreased and drained away, so a tunnel was formed. In the Lanzarote situation, a second flow has created two tunnels, or galleries, one above the other. In some places, parts of the Lanzarote tunnel have collapsed, the resulting holes being called jameos and these provide access to the tunnel system.
There are a number of jameos lining the route of the Lanzarote lava tunnel, the most famous and accessible being Cueva de Los Verdes. Further towards the sea, however, is Jameos del Agua. Beyond the jameos, the lava tunnel extends out to sea as the Túnel de la Atlántida, which terminates as an array of tiny holes. These, however, allow seawater into the tunnel and this fills an impressive, barrel-vaulted, chamber containing a subterranean lake that is accessed via the more seaward and deeper Jameo Chico. The larger, more landward and more surficial Jameo Grande contains a pool of fresh groundwater. The water in the Jameo del Agua rises and falls in co-ordination with the tides albeit with a slight time lag.
Over time, since its formation, approximately ten species of marine animals, such as amphipods, isopods, ostracods and polychaetes have colonized the Túnel de la Atlántida and, hence, the subterranean lake accessed from Jameo Chico of the Jameos del Agua complex. The most famous of these, however, is the galatheid squat lobster [Munidopsis polymorpha _]Koelbel, 1892[ – Canjego ciego. _]Between 10-20 millimetres long, this little squat lobster appears almost perfectly white in the crystal clear waters of Jameo Chico’s lake and has much reduced eyes. It is thought to be blind, although individuals emerge in greatest numbers from their hiding places amongst the rocks that cover the bottom of the lake in the evening as light levels diminish. And, it appears to avoid the area of Jameo Chico’s lake that is illuminated by a shaft of light from the cavern’s ceiling above. Probably, therefore, it has only limited powers of vision, but sufficient to detect changes in ambient light intensity.
The closest relative of [Munidopsis polymorpha _]is[ Munidopsis subsquamosa ]Henderson, 1885 that[ ]occurs at great depths (deeper than 3,000 metres) on the Mid-Atlantic Ridge and which is associated with hydrothermal vents. In Jameos del Agua, however, the small amounts of light that enter the cavern allow the growth of diatoms (Diatomacea) which form a brownish coating on the subterranean rocks and the crabs feed on this by picking at it with their chelae. Females are smaller than males and, as in all crustaceans, mating occurs following each moult. Each fertilised female carries an average of three large, lecithotrophic, eggs under her abdomen and they hatch after a few weeks to form non-swimming larvae that, after several adolescent moults, develop into adults. In the absence of predators in the subterranean lake, _Munidopsis polymorpha occurs in large numbers and the whole tiny ecosystem created by the Malpais de la Corona volcano has become a significant tourist attraction. The Jameos del Agua complex was the first Centre of Art, Culture and Tourism to be created by Lanzarote’s most famous son, the artist, architect and writer César Manrique (1919-1992) and was opened in 1977.
The American jacknife clam, [_Ensis americanus _](Gould, 1870), is a large species of the Solenidae and has a natural distribution on the Atlantic coast of North America from Canada to South Carolina. Razor calms tend to live at or just below the mean low tidal mark in the sand and mud of bays and estuaries, which means that they are usually covered with water for a great deal of the time, except during the lowest tides. Because of its streamlined shell and strong foot, razor shells can dig themselves deeper into the sediment at an extraordinary speed and, if accidentally dislodged from the sediment, can swim. This is achieved by rapid adductions of the shell valves that pump water out of the anterior, burrowing, end of the animal and the same method is used to achieve burrowing. That is, by the liquidisation of the sediment in the depth of the burrow so that it effectively and literally sinks underground.
Razor clams get their names from the rim of the shell being extremely sharp and its shape bearing a strong resemblance to a straight razor. The best way to collect them, without undertaking all the strenuous task of digging, as it can easily outstrip a human digger, is to identify each one’s keyhole-shaped opening in the sand. This identifies the siphonal apertures at the sand surface. If ordinary kitchen salt is poured into the opening, after just a few minutes, the razor clam will emerge from its burrow and can be easily pulled out of the sand.
Because of this burrowing capacity, Ensis directus has few predators other than humans. Such predators include birds, for example the ring-billed gull ([Larus delawarensis _]Ord, 1815) in North America and the Eurasian oystercatcher ([_Haematopus ostralegus _]Linnaeus, 1758) in Europe. Other species of razor clams such as _Solen corneus Linnaeus, 1758, can autotomise elements of their siphons and thereby sacrifice these to such predators whilst saving itself although in this particular case predation may be by fishes (Morton !984b).
[Ensis americanus _]was supposedly introduced into Europe from North America as larvae in tanker ballast water. Its subsequent spread within European waters has, however, been by pelagic larvae. _Ensis americanus has spread rapidly in the southern North Sea countries from its point of introduction in the Elbe Estuary in the German Bight in 1978, to Denmark and The Netherlands by 1982 (Essink 1985, 1986), to Belgium by 1984 and France by 1986 (Bohr 1984, Luczak et al. 1993). It reached the English Channel towards the end of the 1980’s (Knudsen 1989).
In the United Kingdom,[_ Ensis americanus_] was first found in 1989 on Holme Beach, Norfolk. Currently it is to be found at sites along the British east coast south from the Humber and along the English Channel west as far as Rye Harbour, East Sussex. (Howlett 1990). It is reported to be common in The Wash and in some places, for example, Southend on Sea, Essex, it was reported to be one of the commonest living bivalves on the shore in 1995.
Threadfins are silvery-grey perciform fish belonging to the Polynemidae which contains about forty species in eight genera. As will be described, threadfins have numerous, long, thread-like rays, which give them the appearance of catfish. See for example the illustration of [Clupisoma sinensis _]Huang, 1981. This is illusory, however, threadfins being principally marine and the rays have a different origin. Threadfins are found in tropical to subtropical marine waters throughout the world. _Polynemus paradiseus (Linnaeus, 1758), is known from Sri Lanka, India Pakistan and Thailand and the illustrated paradise threadfin was obtained from the Rangoon River, Irrawaddy Delta, Burma, in November 2008.
Threadfins range widely in length. The black-finned threadfin, Polydactylus nigripinnis Munro, 1964 is only 20 centimetres long whereas the Indian four-finger threadfin, Eleutheronema tetradactylum (Shaw, 1804) and the giant African threadfin ([_Polydactylus quadrifilis _](Cuvier, 1829), can both reach lengths in excess of two metres. Both of the latter are important commercial fishes. [_Polynemus paradiseus _]reaches a length of about 25 centimetres.
The body of Polynemus paradiseus is elongate and bulkily fusiform, with the two dorsal fins widely separate, as too are the ventral fins. The tail fin is large and deeply forked; indicating speed and manouverability. The head has a blunt snout located beneath which and quite far back is the mouth. The jaws and palate possess bands of fibrous, villiform, teeth. The most distinguishing and surprising feature of the paradise threadfin and its relatives is its pectoral fins. These are large and when expanded outwards give the appearance of a trimmed-down flying fish. See the illustration of Exocoetus volitans Linnaeus, 1758 (page 143). The lower component of the fins, however, arising from beneath the operculums of both sides are long, thread-like, flexible, pectoral rays. They comprise seven pairs of independent rays, the anteriormost ones of which are relatively short whereas the posterior ones are long and can extend beyond the edge of the caudal fin. The posteriormost pair of rays extend far beyond the tail fin.
Threadfins frequent open, shallow waters in areas with muddy, sandy or silty bottoms. The pectoral rays are thought to serve as tactile, sensory, structures, helping them to find prey within the bottom sediments. They are thought to feed primarily on bottom-dwelling shoals on crustaceans and smaller fish.
Polynemus paradiseus is able to tolerate a wide range of salinities, that is, it is euryhaline. The adults normally live in the coastal areas of rivers, often forming large schools, but enter estuaries and even rivers to breed. Although virtually nothing is known of the species, it is presumed to be a pelagic spawner, probably releasing many small eggs into the water column which then become part of the plankton.
Polynemus paradiseus is said to be extremely rare, although it is caught as a fishery resource. Because of its euryhalinity, it is also difficult to keep in aquaria. Nevertheless, in 2008, seven paradise threadfin individuals were displayed in the Guangzhou Ocean Aquarium in China. These silver-grey fish were described as “lighting up golden fish tanks and wowed many visitors”, the local newspapers reported. With their long, thread-like, fins, the Guangzhou fish were named “the thousand-handed Kwan-yin”, the Goddess of Mercy and Compassion and, even, the saviour of seamen and fishermen. Each fish was valued at more than one thousand yuan (over £120).
On Gran Canaria in the Canary Islands is a quite unexpected wetland reserve. At Maspalomas on the island’s southern end is an extensive dune field, which, at its seaward edge, plays host to the millions of local Canarian, Spanish mainland and other, mostly European, holidaymakers.
But the question is: how has the little marsh and reserve within the dune field been established? The key lies in the sandy beach seaward of the dunes. The sand has ultimately been derived from the Sahara, westerly winds blowing it out of Africa onto the Canaries, so that on the closest islands to the continental mainland, such as Lanzarotte, Fuerteventura and Gran Canaria, it accumulates on their exposed coastal fringes. On Gran Canaria at Maspalomas, however, seasonally blowing easterly and westerly winds keep the sand in a tight cell, creating the dune field. From the mountains of the hinterland, a small stream drains down onto the sandy beach. In the maze of properties that separates inland mountains and countryside from coastal sand, the stream is largely lost and covered, or both. Towards the coast, however, it emerges and, held back by the seaward bank of sand, the out-flowing freshwater creates a small lake called La Chanca de Maspalomas. If one watches the lake at different times of the day, its level rises and falls, out of synchrony with the tides. On these oceanic islands, the tides are small with a vertical maximum height of little more than one metre. Nevertheless, the rising tide allows seawater to percolate through the sand barrier and refresh it. Similarly, as the tide falls, the seawater now diluted by freshwater, percolates back out through the sand barrier. Hence, the little lake comprises brackish water with salinities that will fall following bouts of heavy rain but rise during the dry season as the sea refreshes it.
The result is that the Chanca is actually a brackish water lagoon that has allowed the development around it of an array of coastal plants such as the soft rush, Juncus effusus L., the galingale, Cyperus laevigatus L., the bulrush, Typha latifolia L., and the introduced South African hottentot fig, Carpobrotus edulis (.) N.E. Br. The edge of the Chanca is also colonized by the native tamarix, [Tamarix canariensis _]Wiild.&,& while the dunes surrounding the lake are the home of the tall Canarian palm, _Phoenix canariensis Chabaud, the date palm, Phoenix dactylifera L., two sedges, [Cyperus capitatus _](Vand.) and _Cyperus laevigatus L., the endemic salado, [Schizogyne glaberrima _]DC, with brilliant yellow flowers, and the desert-loving goosefoot herb _Traganum moquinii Webb. The drier fringes of the lake are home to scattered plants of sea lavender Limonium tuberculatum (Boiss.) Kuntze while the floor of the lake is covered by a mat of the tasselweed, [_Ruppia maritima _]L.
One of the fish in the Chanca is the flank-striped white seabream, [Diplodus sargus _](Linnaeus, 1758). But the commonest is[ ]the golden grey mullet,[ Liza aurata ](Risso, 1810)[,_] or, in Spanish, [_lisas o lebranchos. _]Normally, this is an inshore, or[* *]neritic, fish that swims close to the water surface and has a depth range of only about 10 metres. In the Chanca de Maspalomas, however, it is so abundant that it is the main attraction for the tourists who visit the lagoon and feed it. Attracted to it too is the osprey, [_Pandion haliaetus _](Linnaeus, 1758), which occasionally swoops down to snag a mullet in its claws.
[Liza aurata _]is typically mullet-shaped and is characterised by a dorsally flattened head, large eyes and a golden spot on each gill cover. It feeds on small benthic organisms, detritus, and occasionally on insects and plankton. At various stages of its life cycle it lives in brackish and marine habitats. That is, it is catadromous. It rarely enters full freshwater systems, however, but thrives in lagoons and the lower reaches of estuaries but returns to the sea from July to November, where reproduction takes place. The eggs are oviparous, being released by the females when companion males fertilise them. Following hatching, growing juveniles return to their lagoons and estuaries. In the case of the Chanca de Maspalomas, however, possibly because of the wide variations in salinity that occur over the course of the year, it may be that the fish believes it is physiologically, but not physically, moving in and out of an estuary to breed and, hence, reproduces within it. Individuals become sexually mature after one year and continue breeding for three to four years thereafter, especially as the species may reach a length of 60 centimetres. This explains the huge numbers of _Liza aurata within the Chanca.
Felimare picta (Schultz, 1836) lives in rocky sea beds throughout the Mediterranean Sea and the European Atlantic waters of Spain and Portugal, and the islands of the Eastern North Atlantic Ocean, for example, the Açores. It also occurs, however, in the Gulf of Mexico, south to Brazil. The type locality is Palermo, Sicily. The illustrated individual was collected from intertidal rock pools on La Palma, Canary Islands, Spain, in May 2009.
[Felimare _](formerly[ Hypselodoris_])[_ picta ]is a member of the nudibranch (naked gills or branchiae), shell-less gastropods belonging to the always brightly coloured Chromodorididae[. Felimare picta ]is, actually, a complex of six closely related subspecies, all but two of which are geographically separated. Ortea _et al. (1996) have recently revised this species and suggested that there are five subspecies although Troncoso et al. (1998) have suggested six. These are [Felimare picta picta _](Schultz, 1836), _Felimare picta webbi (d’ Orbigny, 1839),[_ Felimare picta azorica ]Ortea, Valdes & Garcia-Gomez, 1996, [_Felimare picta tema _]Edmunds, 1981, _Felimare picta verdensis Ortea, Valdes & Garcia-Gomez, 1996 and [Felimare picta lajensis _]Troncoso, Garcia & Urgorri, 1998. Because of its wide variation in colour and pattern, however, the systematics of this species is confused even more, as [_Felimare picta _]has been referred to in the literature as [_Felimare elegans _](Cantraine, 1835),[ Felimare webbi ](d’Orbigny, 1839), _Felimare valenciennesi (Cantraine, 1841),[_ Felimare edenticulata _](White, 1952) and [_Felimare tema _]Edmunds, 1981.
Notwithstanding, Felimare picta is the only eastern Atlantic species of Felimare, other than Felimare villafranca, which has more than two parallel yellow lines on the dorsum. [Felimare picta _]is the largest chromodorid in the Mediterranean Sea reaching up to 10 centimetres in length. Juveniles show three bright longitudinal lines dorsally, whereas in adults the pattern is more complicated with more numerous broken or entire lines. A characteristic of this species is that the two more lateral dorsal lines reach as far forward as the rhinophoral sheaths. In adults, the lines form a yellow circle around the edge of the rhinophoral sheath and continue forward. Other distinctive features of _Felimare picta are the uniformly dark blue rhinophores and the external yellow rachis of each branchial leaf.
Regardless of the colour complexity exhibited by Felimare picta, its function is unclear. One possibility is aposematism, or warning coloration. The term aposematic describes a group of anti-predator adaptations where a warning signal is associated with the un-profitability or un-palatability of a prey item to potential predators. It is a form of advertising, possibly taking the form of conspicuous colours in Felimare picta. Certainly, one can only wonder why such predators as fish are not attracted to such an obvious, soft-bodied, colourful potential prey item. Many experiments have, however, shown that when a variety of brightly coloured nudibranchs are dropped through the water column of a aquaria containing sympatric fish, the latter may snap at the former but quickly spit it out. This suggests that such nudibranchs are either distasteful or noxious and their colours advertise this fact. Such aposematic signals are, thus, beneficial for both the predator and the prey, for they both avoid potential harm.[_ Felimare picta_] is itself a grazing predator, Ortea [et al. _](1996) recording that the species[ ]feeds on the sponge _Dysidea fragilis (Montagu, 1818) (Demospongiae) in the Mediterranean.
One of the strangest coastal plants I have ever encountered is the broom-rape [Cistanche phelypaea _](L.) Cout.. The species is restricted to arid and semi-arid regions of southern Spain, the Mediterranean coastline of North Africa to as far east as Egypt and the Sudan and, as herein described, the Cape Verde Islands and, presumably, West Africa. Of the Cape Verde islands, Sal is the driest with rain falling perhaps only twice each year. As a consequence, the island’s landscape is lunar with brown rocky plains and deserts of sand that are shifted by the winds that lash it seemingly interminably. The Monte Grande Volcano on the northeast is some 400 metres high – the remaining land flat, overgrazed by goats, and seemingly desolate. In places, subterranean salt water appears on the land’s surface, evaporates and creates dazzling patches of salt – hence the island’s name. The parched hinterland of Sal grades slowly into the intertidal, which is characterised by halophytic plants. Among these are the bushy beancaper, _Zygophyllum fontanesii Webb & Berthelot, a perennial small shrub with highly succulent, ovoid leaves, and the prostrate sea purslane, [_Sesuvium portulacastrum _](L.) L.. This, near global, halophyte is widespread throughout much of the semi-tropical and arid intertidal flats of the Atlantic coastline. I have seen it before on shores of the Gulf of Mexico and the Açores. On Sal, it grows as a sprawling perennial herb up to 30 centimetres high, with thick, smooth stems almost a metre long. It has smooth, fleshy, glossy green, lanceolate leaves. Its little star-shaped, violet and white flowers (sometimes pink or purple) open to the sun each day against the backdrop of the mesh of fleshy, leaves. The leaves of [_Zygophyllum fontanesii _]are similarly fleshy and the five-petalled flowers are white with a yellow core of anthers and stamens.
Periodically and either next to the shrubs of [Zygophyllum fontanesii _]or amongst the mats of[ Sesuvium portulacastrum_], could be seen tall, bright yellow, flower spikes, here growing up to 50 centimetres tall. This is the broom-rape [Cistanche phelypaea, _]parasitic on its neighbouring plants.[ _]This plant has no leaves and, hence, no mechanism for sustenance through photosynthesis, and its fine roots intermesh and fuse with those of its hosts. It is an obligate root parasite, wholly dependent upon its hosts for water, minerals and nutrients. Removing the flower spike (A) from the sand reveals a bulb-like tuber from which it arises. Occasionally, each tuber has a subsidiary one (B), which has the appearance of a scaly onion. Clearly, the plant can reproduce vegetatively. As the spike grows ©, the flowers begin to develop, appearing as a swollen paler head to each yellow stalk and its receptacle.
A single flower is illustrated in D. There are two pairs of ventrally-located stamens, the anthers of each comprising a capsule, covered in fine, short, curly hairs, that splits lengthwise to divide into two halves (E) thereby exposing the pollen to the single white, spatula-shaped, stigma located atop a long, curved, style (F). At the base of the flower is a receptacle (G) containing two pairs of ovaries each of which comprises a mass of pearl-like ovules.
Elsewhere, Cistanche phelypaea parasitises a wider variety of halophytic plants, including species of Arthrocnemum, Anabasis, Salso, Seidlitzia and [Suaeda, _]plus the two plant species, _Zygophyllum fontanesii and Sesuvium portulacastrum, here encountered on Sal. Cistanche phelypaea, and the various other species of broom-rapes (Orobancheaceae), can parasitise commercial plant crops and are, hence, serious agricultural pests.
Tellina tenuis da Costa, 1778, is a species of the Tellinidae. It occurs off the coasts of north west Europe and Morocco and in the Mediterranean and the Baltic Seas. It is widely distributed and common around the coasts of the British Isles. The shell of Tellina tenuis is brittle, laterally flattened and grows to a length of 25 millimetres. The shell is oval in outline but the valves are asymmetrical with the right larger and more convex than the left. The posterior end is also shortened, wedge-shaped and curves to the right when viewed from the dorsal aspect. The anterior end is gently sloping and broadly curved at the anterior margin. The shell is covered with fine concentric lines and is variably coloured from creamy-white, yellow, shades of pink, orange and brown, often in bands, Internally, the shell is coloured similarly, but somewhat fainter. The mantle is creamy-white and fringed with tentacles.
Tellina tenuis is found from the middle component of the intertidal down to a depth of about seven metres. It inhabits fine, clean sand, with low percentages of silt. and usually lies slightly on its left side. To burrow, Tellina tenuis has a large, digging, foot. It also has two long siphons, which can be extended to the surface of the sediment. Water and food particles are drawn into the through the inhalant siphon while water is expelled through the exhalant. [Tellina tenuis _]is a deposit-feeder taking in fine detritus and diatoms, which it sorts and selects on large labial palps inside the mantle cavity (Yonge 1949). When the tide is out _Tellina tenuis descends into the sand to a depth of about ten centimetres but ascends to nearer the surface when the tide rises and it recommences feeding. In some locations, at the low tide level, it may be the most abundant organism in the intertidal reaching a density of 3,000 individuals per square metre. A population of Tellina tenuis in the Exe Estuary was shown by Holme (1950) to be distributed uniformly, indicating a significant degree of over-dispersion. Holme suggested that the bivalve’s spacing was correlated with the rotational foraging activities of the inhalant siphon on the sand surface. That is, each individual was distributed in their own little territories circumscribed by the activities of the inhalant siphon. Young flatfish, however, sometimes feed on the tips of the protruding siphons but they are able to regenerate (Trevallion et al. 1970).
Individuals of Tellina tenuis are either male or female and spawned gametes are liberated into the water column from June-August where they are fertilised. The resulting larvae are free swimming and form part of the temporary zooplankton, serttling onto sandy beaches after a few weeks. Since individuals in these beaches may have a lifespan of up to five years, their territories are kept stable over long periods (Stephen 1928). Adult [_Tellina tenuis _]are, however, sensitive to low temperatures and numbers are reduced after cold winters, opening up new ground for recently-recruited juveniles to re-populate.
The Greek species name of[_ pes-pelecani ]means ‘pelican foot’ – with good reason. The outer lip of the shell is grossly enlarged in the adult shell to resemble a webbed pelican’s foot. It is even palmate with distinctive ‘toes’ or, more correctly, shell rays. _Aporrhais [pes-pelecani _](Linnaeus, 1758) was first encountered while attending the Easter undergraduate marine field course at the laboratories of the Marine Biological Association of the United Kingdom’s headquarters at Plymouth in 1964. The institute had a small research boat – the R.V. [_Calanus _]that could trawl and dredge and a few students boarded it one day to get the experience. It was a stormy day and not the most pleasant of experiences – though we did trawl up a few[ Aporrhais. _]Back in the laboratory in a tray of seawater, it was one of the strangest animals we students had ever seen.
The animal was also important for another reason, it had been researched by (Sir) Maurice Yonge at the marine station at Herdla in Norway and from this had arisen his seminal paper on it (Yonge 1937). [_Aporrhais pes-pelecani _]lives at depths of between 10-200 metres on firm, muddy, gravel bottoms and into which it buries itself. It occurs from Norway south to the Mediterranean and Black Sea although few people will ever see it, except occasionally as beach drift.
In August 2009, however, there had been a storm in the Kattegat on Jutland’s east coast and many [Aporrhais pes-pelecani _] were washed up alive and, fortunately, of many different sizes. They were kept in a small aquarium containing sand and seawater and they crawled around happily and some even buried themselves and lived long enough for me to draw them. The juvenile shell (A) is ~10 mm in height, does not have the palmate outer lip and has a more simple long, fusiform, shape with a sharply pointed siphonal canal. As the shell grows, however (B-F), the outer lip grows disproportionately to the shell. This is an example of hypermorphosis (gerontomorphosis), described by another greatly admired figure – Sir Gavin de Beer (1899-1972) who first elaborated upon this topic in his book _Embryology and Evolution (1930) that was expanded and retitled Embryos and Ancestors, in the 1940’s edition. In such a short book, de Beer brought embryology into the growing orthodoxy on the relationship between ontogeny and phylogeny. In the book, de Beer suggested that the exaggerated antlers of the extinct European elk, Megaloceros, were an example of hypermorphosis. Males of this elk stood two metres tall at the shoulders and were adorned with gigantic antlers that were almost four metres across. The species probably survived until historical times, perhaps being wiped out by Neolithic man. The analogy with [Aporrhais _]is striking except that in the case of _Megaloceros the antlers were for fighting and served as a sexual stimulus for females, a case, therefore, of sexual selection.
In Aporrhais pes-pelecani the greatly expanded outer lip of the shell grows progressively and then accelerates as adulthood, maybe coinciding with this, sets in. It probably, therefore serves to stabilise the adult animal in the sediment and to direct water containing organic particles of food into the mantle cavity located above the head and wherein the ctenidia, or gills, collect them and direct them to the mouth. As described by C.M. Yonge (G). The ciliary-engendered current of water through the burrow also aerates it and provides the animal with oxygen. When [Aporrhais pes-pelecani _]emerges from its burrow at night and moves about (I), one can see the head that is coloured a brilliant orange-red (H). It also has a long snout that is used to collect surface deposits. _Aporrhais pes-pelecani thus has two methods of exploiting different food resources. When the animal retreats into its shell, the head and foot are retracted and the aperture is sealed with the ovally-elongate, yellow, operculum (J).
The halophytic (sea-loving) Cakile maritima[* *]Scop., has been encountered in many parts of the northern hemisphere, even North America where it has been introduced onto both the eastern and western coasts. In its native Europe, including the Mediterranean, however, Cakile maritima occurs at the extreme seaward fringe of sandy beach communities where it can form low mounds up to 50 centimetres in diameter and up to 40 centimetres tall. Like many coastal plants, the long, narrow, leaves of C. maritima are fleshy and divided pinnately into round-tipped and oblong lobes. The leaves often have a non-uniform, tattered, wind-blasted, appearance, and are of varying shades of green, from bright to dark as they age. They are fleshy to conserve water in this arid environment, which is really most like a coastal desert – the water and sea spray being too salty to be refreshing. The plants can also withstand burial by wind blown, aeolian, sand, growing through it again in the early summer as the winter storms abate. Conversely, the spreading roots of the sea rocket help bind the sand together – thereby stabilising the beaches it grows on and creating future habitat for the advancing dune plants.
The tiny flowers of Cakile maritima comprise four petals and are occasionally, and confusingly, white but are most often of a delicate, pale lilac hue. The plants flower from July to early autumn with pollination by insects. Once fertilised, the plant produces fruits that comprise swollen, transversely two-jointed pods the upper one of which is the larger but both possessing a single seed. The upper pod breaks off the plant, but the smaller lower one remains on the plant until the seed is shed. In this way, there are two possible means of seed dispersal – the seed from the former, larger, pod can be washed away by waves, whereas the seed from the latter, smaller, pod will remain near the parent plant to develop into a new plant [_in situ. _]The seeds are resistant to seawater for many weeks and are transported by long-shore currents to other beaches. There, they are washed up with other flotsam and, held within piles of stranded seaweed, germinate and therein and thereby find for themselves a ready source of fertiliser. They thus grow quickly on the beach strand.
Cakile maritima is wonderfully adapted for life on wind-blown, often tempestuous, storm-tossed beaches, as on the west beach of the River Arun, at Littlehampton, West Sussex on the south coast of England. The plant pioneers the colonisation of the beach to create new land in the form of sand dunes that other plants can colonise and further stabilise. It thus represents an amazing example of not just adaptation to life on one of the most extreme environments on the planet (for a plant) but also has to be admired for its tenacious grip on a harsh world that allows others to follow where it leads.
The English botanist Joseph Dalton Hooker (1817-1911) had noted floristic similarities between the Falkland Islands and Iceland, and South America and Europe, respectively, neither having hardly any indigenous species. One well-known example at the time was the beach pea, Lathyrus japonicus[* *]Willd., which was then thought to have a North American and rare British occurrence. To explain this, the English malacologist Edward Forbes (1815-1854) proposed in 1846 that a great land mass had existed in the Miocene encompassing northern Europe and Spain, and extending out from the Mediterranean far westwards into the Atlantic Ocean virtually to the coast of North America and, thereby, accounting for similarities in the plant and animal fossils of Europe and North America. In Forbes’s eyes, the beach pea provided good evidence of this formerly contiguous Atlantic distribution.
On his return to England following the successful voyage of H.M.S. Beagle from 1831 to 1836, Charles Darwin (1809-1882) became sceptical of Forbes’s lost land and sent seeds of the Western Atlantic fabaceans, the sea-heart Entada gigas (L.) Fawc. & Rendle and Mucuna urens (L.) Medik. from Açorean beaches to the Royal Botanic Gardens at Kew where they were planted, germinated and produced healthy, mature, vines. Subsequently, Darwin set up his own experiments in the glasshouse at Down House where he immersed the seeds of 87 species of common plants in seawater for a month. He then tried to germinate them and found that over half (64) had survived. By his own calculations, ocean currents could thus have taken such seeds well over half way across the Atlantic Ocean and he concluded that no Forbesian landmass was necessary to explain Hooker’s biogeographic similarities.
Darwin thus set in place an alternative, and more plausible, theory to that of Forbes, which suggested that newly-emergent islands could be colonised naturally by plants and animals from other locations and that, through natural selection, such isolated individuals could evolve into distinct species. The beach pea has thus played a significant role in our modern understanding not just of geology and biogeography but also the concept of evolution through natural selection. In fact, the beach pea, an herbaceous perennial with trailing stems up to one metre long is now known to be circum-polar. That is, it occurs not just in Great Britain and North America but also Japan. The flowers, between 15-20 millimetres broad, have beautiful deep purple petals and paler, sometimes almost whitish, but more often lavender, wing and keel petals. The seed pods can be eaten but may cause a form of paralysis called lathyrism. Lathyrism, or neurolathyrism, is a neurological disease of humans and domestic animals, caused by eating certain legumes of the genus Lathyrus, notably[_ Lathyrus sativus_] L., the grass pea. The seeds, however, unlike Darwin’s domestic sweet pea, Lathyrus odoratus L., seeds, which did not survive in seawater, remain viable in the sea for up to five years and can, hence, be distributed virtually worldwide in cool temperate regions. The seeds germinate when their outer coat is abraded by waves in the surf on gravel and sand beaches.
The beach pea occurs on the west coast of Ireland, on the shingle beaches of southern England, especially at Dungeness and, particularly, at higher latitudes in Europe, for example on the sand dunes facing the Skagarak and Kattegat in North Jutland.
L ittle sister to its more famous neighbour, Madeira, the tiny, nine kilometre long, island of Porto Santo (Holy Port), lies just 50 kilometres to the southwest. The Madeira Archipelago is itself located ~600 kilometres off the northwest coast of West Africa. The island is most famous because Christopher Columbus (1451-1506), was resident on the islands of both Porto Santo and Madeira between ~1479-1482. This came about when in either 1478 or 1479, he met and married Felipa Perestrello e Moniz, the daughter of a respected, but poor, Portuguese nobleman, Bartolomeu Perestrello, and who had been made in 1445 the first governor of Porto Santo. As part of the dowry, Columbus received all of the deceased Perestrello’s charts and wind records of the Portuguese possessions in the Atlantic. Soon after their marriage, the couple moved with the Perestrello family back to Porto Santo where Felipa’s oldest brother took over its governorship.
Porto Santo was officially discovered (although the Madeiran islands were probably known about long before that) in 1418 by João Gonçalves Zarco and Tristão Vaz Teixeira, knights of Prince Henry the Navigator (1394-1460). Henry was the third son of John I (1358-1433), the tenth King of Portugal and the Algarve and who famously married, in 1387, Philippa of Lancaster, daughter of John of Gaunt (1340-1399) thereby consolidating the Anglo-Portuguese Alliance of 1373.
What makes Porto Santo distinct is that the southern coast of the island comprises an almost nine kilometre long sandy beach and to the rear of which Vila Baleira (now simply Porto Santo) is the island’s capital. Since Columbus’s first love was the sea, there can be no doubt that he patrolled this beach. It is thought that he found seeds of the sea heart [_Entada gigas _](L.) Fawc. Rendle, often now called Columbus’s bean, in the drift on the shore of this beach and which since the plant was unknown in Europe, must have come from an un-discovered (then in ignorance of Viking achievements) land to the west – the Americas (or, rather, as he thought, Asia).
On the long sandy beach of Porto Santo, local people stroll, and they go fishing using beach-casting rods and tiny baits. They are here to catch[_ Trachinotus ovatus ](Linnaeus, 1758). Throughout its wide Iberian, Mediterranean, West African, and Macaranesian island range, this fish has a large number of colloquial names, but is variously known as the _pompano in English, [derbia _]or _plombeta in the Açores and rhanosa in Porto Santo. Trachinotus ovatus is essentially a surf species, occurring in shoals and feeding on small crustaceans, molluscs and smaller fishes that either live here as specialised occupants or find themselves washed here as dead or dying flotsam. It attains a length of up to 70 centimetres and is a beautiful, almost totally silver, streamlined, fast-moving predator. It has dark spots on the apices of its dorsal and pelvic fins and the tips of both caudal fins. In addition there are five spots on the anterior flanks. Judging by how many fish the locals were catching each evening, the more experienced among them having caught a dozen or more, there must be large shoals of them in the surf and which all the holidaying swimmers, surfers and paddlers were wholly unaware of.
Here in Western Europe, whether we know it or not, the sprat[_, Sprattus sprattus _](Linnaeus, 1758), with silver-grey scales, is one of our commonest coastal fishes. Fishes that we call sprat occur virtually throughout the world’s oceans and, sometimes, especially in former times, in huge shoals. The sprat belongs to that great fish family, the Clupeidae, which also contains the herrings, sardines and pilchards. Once, herrings were a staple of the British diet but stocks are now badly and sadly depleted. Sardines and pilchards are also caught and canned – being another staple of the European diet. Sometimes, we may eat sprat as whitebait, especially the juveniles – but the colloquial term in reality encompasses any small, surface dwelling, inshore fish.
The true sprat, Sprattus sprattus,[* *]shows a strong tendency to migrate between offshore winter feeding and inshore summer spawning grounds. Offshore it feeds on planktonic crustaceans, moving up to the sea’s surface at night where they can be caught by gillnetting or purse seine fishermen. In summer, they move inshore to collectively spawn at shallow depths of between 10-20 metres. Each female may produce between 6,000 to 14,000 eggs and the young drift inshore where they become the food of other summer fishes such as the mackerel, Scomber scombrus Linnaeus, 1758. Young sprats are commonly known as brisling.
As a boy, I can remember seeing one gigantic shoal of mackerel moving up the mouth of the River Arun in West Sussex and they were almost certainly chasing an equally large shoal of juvenile sprat that often move into estuaries in summer to feed. Indeed, because they are able to tolerate salinities as low as 4‰, sprat can move far upstream in search of microscopic food items. Even today, in summer, one can sometimes see the river water shimmering silver as the shoals of sprat enter it.
The majority of the huge shoals of brisling sardines that are harvested off the coasts of Norway and Scotland, being so small, typically less than 25 millimetres in total length, are simply processed for fish-meal. The flesh of the fish is around 12% fat and is a source of many vitamins. Some of them, however, are canned, smoked or frozen, often for fish bait, but, increasingly, one can find them on the fishmonger’s slab where they are an inexpensive source of fish protein, omega oils and vitamins. As well as canning, they are variously salted, fried, grilled, baked and marinated but pan-fried, with a light dusting of flour, in butter and lemon juice, sprat make a wonderful summer treat. They are, however, the main, but not exclusive, component of whitebait sold in British seafood restaurants fried in a light batter and typically served as a starter to the meal.
Like most rivers in England, especially, the Arun on the south coast of West Sussex has been canalised for all its 19 kilometre length from its source, the gills of St. Leonard’s Forest to the east of Horsham, to its mouth at Littlehampton. Here, its estuary would naturally have spread out to form a huge, two kilometre wide, marsh (Morton 2007b). Following on from its canalisation too, the original Arun’s former marsh would have survived as remnants on the river’s west bank opposite the town of Littlehampton on the east. That is, until the late 19th century when the Littlehampton Golf Club reclaimed the marsh for its own purposes. One of the components of the former marsh and which there are surviving remnants of was the samphire or glasswort, Salicornia europaea Linnaeus, 1753. Associated with this plant throughout much of its European range is the tiny (up to four millimetres long) staphylinid beetle[_ Bledius spectabilis _](Kraatz, 1857).
Bledius spectabilis is characterised by a long ‘horn’ on the front of its head – a little like a rhinoceros. The insect is, actually, a colonial, sub-social, species and on the River Arun’s west bank there is a small colony of the little animals. Each individual female beetle digs for itself and its male partner a burrow that has a distinctive structure in vertical section. At low tide, the beetles emerge to feed on fragments of plant tissues including green algae but when the tide begins to return, each beetle re-finds its burrow and, once inside, seals up the entrance with a plug of sand taken by the insect’s mandibles from the walls of the living chamber of the burrow (Wyatt 1986). The burrow structure has been illustrated by Evans et al. (1971). The sharp curvature to the entrance of the burrow acts to create a surface tension effect, that is, the air trapped inside the burrow acts like a bubble to keep the water out. This is crucial for the animal’s survival because, like most insects, Bledius [_spectabilis _]is an air breather through spiracles on its abdomen and, hence, if immersed by the incoming seawater would drown. Back in its sealed burrow, however, each beetle has a pocket of air to last it over the next high tide period. When the tide recedes, the burrow is un-plugged and each beetle emerges to again forage over the surrounding sand surface.
The burrow serves another purpose, however, which is that, following successful mating, each female Bledius spectabilis, whose thorax turns a rose pink upon maturity, lays her eggs inside it, each one within its own pocket in the burrow wall, and there cares for not just them but the juveniles that hatch out and reside with her inside the living chamber. To feed her growing brood, the mother makes periodic excursions onto the sand surface where she collects more plant food that is stored in special larder chambers in the burrow (Bro Larsen 1952). Over time, each hatchling undergoes a series of molts, which result in an instar that, by this means, grows, step by step, and changes form, to eventually become an adult and leave home. Such maternal care enables this air-breathing insect to colonise what would otherwise be a wholly inappropriate habitat (Wyatt 1986, Wyatt and Foster 1988, 1989a).
One would think too that occupation of the burrow would protect not just adult Bledius spectabilis but also their eggs and larvae from virtually everything. Not so, however, because Wyatt (1986) has shown that eggs and their contained larvae are preyed upon by another, carabid, beetle Dichierotrichus gustavi, while Wyatt and Foster (1989b) have shown that an ichneumid wasp, [Barycnemis blediator _](Aubert, 1970),[* *]parasitises the 1st instar larvae of _Bledius spectabilis by entering the maternal burrows but can only do so when the female is outside foraging, highlighting the importance of parental care.
The Parque Natural da Ria Formosa comprises an extensive lagoon system that 18,400 hectares encompasses 60 kilometres of coastline between Manta Rota and Vale do Lobo in the Algarve region of southern Portugal. Made up of sand dune islands, marshes, saltpans and freshwater lakes, the habitat is a sanctuary for a rich biodiversity of coastal plants and animals, especially migrating waterfowl. A sweeping tract of coastal sand dunes guard the mouth of the estuary are partly held together by the vegetation that has colonised them. In addition, there are coastal conifer woods of the tall, 20-30 metre, maritime pine Pinus pinaster Aiton.
The lagoon system has been exploited from the earliest human history, for example, the stone or umbrella pine, Pinus pinea L., which is native to the Mediterranean has been cultivated for their edible pine nuts since prehistoric times. In the park too, are five Roman salting tanks near the freshwater lagoons. Dating from the 2nd century AD, they were once used for salting fish prior to their distribution all over the Roman empire. There are also tide mills, a late 13th-century invention utilising changes in water levels associated with the tides that were once common in the lagoon and river estuaries all along the Portuguese coastline. Those in the Ria Formosa were the last operational ones along the Algarve coast.
The parque was visited on a hot day in July 2010 in the company of a group of birdwatchers. All hoping to see the purple swamphen or, more accurately, the Mediterranean subspecies Porphyrio porphyrio porphyrio (Linnaeus, 1758[)_]. I was more interested in the vegetation surrounding the lagoons and where the gallinule was likely to be found. Two of the plants making up much of this embankment vegetation were the silver sea stock, _Malcolmia littorea (L.) R.Br. Rank. (A), and the dry purslane, Limoniastrum monopetalum (Linn.) Boiss. (B), neither of which I had seen before.
The former is a member of the Brassicaceae. It is a profusely flowering annual, which grows to about 15–30 centimetres tall. Here, in the Ria Formosa it makes a mass of pink, purple and white four petalled, cruciform, flowers each with a white to yellow centre and some 2-3 centimetres across.. At the time of the visit it was in full flower and the air was full of its fragrance. It is a branched perennial with narrow woolly-white stems and lanceolate leaves giving the plant a silver-grey appearance.
Alongside the stock, equally profuse, was the dry purslane, or salting, Limoniastrum monopetalum, which is a member of the Plumbaginaceae. This is also an evergreen shrub, but growing up to two metres high (perfect for the swamphen). Its branches are more or less upright with green or reddish young twigs bearing lanceolate to obovate-spatulate leaves covered to varying degees in carbonate deposits making them whitish-grey. The flowers, varying in size from 10-20 millimetres in diameter occur in racemose inflorescences. Like the silverstock, they are showy pink to purple, In the centre of each are five stamens and yellow anthers. This purslane blooms virtually year round and the two plants, here in the national park, were both in full flower.
In Hong Kong, large crabs, some eight centimetres in carapace width, annually migrate downstream from their freshwater abode towards the sea. These are the Chinese mitten crab, [_Eriocheir sinensis _](H. Milne Edwards, 1854), characterised by the hair-like covering on the chelae, especially well-developed in male individuals, and giving it its name of mitten crab (A, C). The crab’s colour varies from yellow to brown and, apart from the mittens, is not particularly noteworthy. The Chinese mitten crab is found in the northern hemisphere only and is native to Asia. It occurs in temperate and tropical waters between Vladivostock in Russia and southern China, including Taiwan and Japan. The Yangtze River is a major habitat for the crab in China.
The taxonomy of mitten crabs has been confusingly problematic. The genus Eriocheir was considered to comprise four species (E. japonica, E. sinensis, E. heupensis and E. ogasawaraensis). The Hong Kong species[_ ]was identified as[ Eriocheir japonica ](de Haan, 1835) by Dai and Xu (1995) although modern techniques of DNA analysis have shown that all the above species are synonymous (Tang _et al. 2003, Zhang et al. 2009). If this is the case, then the correct name for the species is [_Eriocheir japonica _]although [_Eriocheir sinensis _]is the one used most widely.
Throughout its life, the Chinese mitten crab occupies different ecosystems depending on its stage. The Chinese mitten crab is catadromous, that is, it moves from freshwater streams where it spends its juvenile years to saltwater habitats in order to reproduce. Adult crabs occur in fresh, brackish and salt waters, but oviparous females are normally found in greatest numbers in the sea. Larval stages are found in the open water of bays and estuaries. Juvenile crabs are typically found in tidally inundated tributaries as they are moving back to freshwater. After reaching approximately 1-2 centimetres in carapace width, male and female crabs can be differentiated by the shape of the abdomen, which in the former is narrow (D) and shaped like an inverted funnel whereas in the latter it is round (E) and occupies most of the area of the thorax.
Mature adults migrate downstream during the autumn to reproduce in brackish or salt waters. Following copulation, male atop the female, she broods her fertilised eggs beneath her abdomen, held in place by her abdominal appendages and the fringe of long chaetae that surrounds the structure. Upon release and hatching, larvae are planktonic for one to two months. During this marine free-swimming phase, larvae pass through a series of developmental stages, that is, a brief non-feeding pre-zoea stage, five zoea stages and one megalopea stage. Following the megalopal stage, the larvae metamorphose into juvenile crabs that settle to the bottom, usually in late summer. Both males and females are thought to die following reproduction.
The Chinese mitten crab is a traditional food resource in China, where it supports an important aquaculture industry with an annual production of 200,000 tonnes in 2000. The preferred crabs are those captured during autumn, as they have full gonads and stored energy in preparation for reproduction and the coming winter. As a consequence, the crabs appear in Hong Kong shops and markets in autumn. They were originally shipped to Hong Kong by wholesalers in bamboo pots and then, if still alive, tied up using bunches of rice grass, [_Oryza sativa _]Linnaeus (B), effectively immobilising the crab and keeping the spiky carapace and claws secure for handling purposes.
The Chinese mitten crab has been introduced into North America and Europe. It was first recorded in Europe from Germany in the River Aller in 1912. Subsequently, the species spread into the Baltic Sea via the Kiel Canal and it now ranges throughout Finland, Sweden, Russia, Poland, Germany, the Czech Republic, the Netherlands and Belgium. Between 1959 and 1968 it spread into and along the French Mediterranean coast although self-sustaining populations have not been reported from here. This is not the case in Great Britain where the species was introduced into the River Thames in about 1935 (Clark et al. 1998). Since its first discovery, the species has spread throughout the Thames, its population size growing apace.
The major problem with the crabs is their burrowing activities, which result in erosion to river embankment causing collapse and posing serious threats to flood control and water supply efforts. In Asia, the crab is also the second intermediate host for the human lung fluke parasite, Paragonimus westermani _]Kerbert, 1878, but because of the absence of the first intermediate snail (species of [_Semisulcospira) host in Europe, adverse effects on human health in its non-native environment are minimal.
Anurida maritima (Guérin-Méneville, 1836) is a near cosmopolitan collembolan insect that occurs in the intertidal zone. It can occur in aggregations of up to several hundred on the surface of rock-pools or on sandy beaches near rocks. Despite it being apparently common everywhere, I had never found this species until examination of a small patch of surviving sand on a beach close to the sea on the west bank of the tidal River Arun in Littlehampton in West Sussex. The sandy beach abuts surviving island on this bank of the river and where there is an ancient embankment, now largely in disrepair, of formed chalk blocks. This is a perfect habitat for the minute insect, which is only up to 3 millimetres long and hence capable of easily retreating between the blocks when the tide returns. Anurida maritima is also to be found on tidal marshes.
Anurida maritima is a wingless dark slate blue insect although it superficially appears black. Its body is roundish, expanding slightly towards the rear. The head bears a pair of eyes and a single pair of antennae. The thorax comprises three body segments, each of which bears one pair of legs, while the abdomen comprises six segments. The entire body is covered with minute hydrophobic hairs which allow the animal to stay afloat on the meniscus of rock pools close to which it spends much of its life.
In the warmer parts of its range, Anurida maritima is active throughout the year, but in cooler temperate regions, as in Great Britain, it is only active in the summer months, overwintering within rock crevices as eggs. When active, it typically emerges during low tide periods and assumes the role of a scavenger of the upper intertidal zone, feeding on carrion, which can include crustaceans, such as barnacles and molluscs, previously predated by dogwhelks, Nucella lapillus (Linnaeus, 1758), for example. At such free meals, it can occur in large numbers. Like many intertidal animals, Anurida maritima has a distinctive endogenous tidal cycle with a period of 12.4 hours (Manica et al. 2000), thereby moving in rhythm with the tides, but using their eyes to orientate themselves during their low tide forays.
Typically, it emerges from the rocks as the tide recedes and wanders over the sand surface to feed on decaying surface matter including carrion when it is present (Joose 1966) and, over time, in increasing numbers. In this, it is aided by the production of an aggregating pheromone, which is an important aspect of collembolan biology, so that when a good source of food is found it can occur in very large numbers of individuals attracted over long (for a three millimetre long insect) distances (Manica et al. 2001). It returns to its rocky home as the tide returns using stored visual clues. Safely back inside its crevice, this air-breathing insect encloses itself in an air bubble. This acts not only as an oxygen store but also as a compressible gas gill, which enables survival while the little creature is submerged by the tide for about three hours (Zinkler et al. 1999). Because it occurs high on the shore, this is sufficient time, therefore, for it to survive being emersed in seawater. One needs a good, practised, eye to spot these tiny insects, individuals impossible to detect, but crowds of them around a dead mussel are more obvious.
Other than taxonomic studies, there is practically no work dealing with the group of decapods of which the genus Athanas is an important and diverse member. Athanas is only found in the Indo-West Pacific with a range extending from as far north as Japan and as far south as Australia (Banner and Banner 1973), eastwards into the islands of the Western Pacific and westwards into the Indian Ocean. From its focus in the Western Pacific, the genus is represented by 28 species whereas in the Indian Ocean there are but eight, ranging from the east coast of Africa to the Indonesian Archipelago (Banner and Banner 1960). Of these eight, only one is known from the mainland coast of the Indian Peninsula (Kemp 1915), the others all occur on remote archipelagos such as the Maldives and Laccadives.
Only one species of Athanas, that is, Athanas dorsalis (Stimpson, 1860) has been recorded from Hong Kong (Morton 1988, Bruce 1990) and here it occurs with the lower intertidal, exposed rocky shore, sea urchin [Anthocidaris crassispina _](Agassiz, 1863). On the Laccadives and the Gulf of Mannar, Thomas (1974) and Sankarankutty (1962), both record [_Athanas dorsalis _]as occurring under the oral surface of the black sea urchin _Stomopneustes variolans (Lamarck). Banner and Banner (1960) record that of the 70 individuals reported upon by them from various locations in the Indo-West Pacific, fifty were living with species of Heterocentrotus. Throughout its huge range in Australia, including Norfolk Island and Lord Howe Island, however, the species lives with six different sea urchins and assumes the colour of its particular host (Hipeau-Jacquotte 1965)
In Hong Kong,[_ ]Morton (1988) showed that its sea urchin host crops the rich band of low intertidal algae it is associated with and that _Athanas dorsalis does too. Typically, the urchin lives in crevices or underneath stable boulders and emerges to feed at night. Its alpheid shrimp emerges with it. It lives beneath the urchin typically in a male-female pair and the two vigorously defend their home urchin against other shrimps. When another pair are introduced into an aquarium with an urchin and its established pair, the two take up opposite sides of the now mutual host with much clicking of the enlarged chelae and dancing to and fro. Eventually, if another urchin is introduced to the four, one pair leaves the disputed territory and adopts the newly arrived host.
The most important aspect of Athanas dorsalis is, however, that it was first described from Hong Kong by William Stimpson (1832-1872) who was the biologist on the United States North Pacific Exploring Expedition and subsequently the director of the Chicago Academy of Sciences. Stimpson stayed in Hong Kong for six months, living aboard the naval sloop Vincennes, assiduously collecting marine animals and describing them, albeit poorly. One of these was [_Athanas dorsalis _]later described by Bruce (1990) also from Cape d’Aguilar, Hong Kong. Cape d’Aguilar and Hong Kong are thus the type localities for this species making the peninsula and its contained marine reserve highly important locally and taxonomically.
The curled picarel, Centracanthus cirrus Rafinesque, 1810, is a member of the[* *]Centracanthidae and occurs in the eastern Atlantic in the waters of the Mediterranean, Portugal, Morocco, Mauritania and southwards to Angola. It also occurs in the Canary Islands, Madeira and the Açores. And, it was in the latter locality that it was encountered, and where it is a minor fishery resource (Santos et al. 1997). It is a neritic species, that is, it lives in coastal waters, and is found over rocks and gravelly bottoms. As such, therefore, it is perfectly adapted to the shallow Açorean offshore rock/gravel seabed. It is also a deep water species generally occurring down to depths of 200 metres. In the eastern Ionian Sea, it occurs at depths from 327-464 metres (Nelson 1984). A maximum length of 34 centimetres has been recorded for the species but, in general, individuals do not grow much beyond 12 centimetres and one of around this size is illustrated. Centracanthus cirrus spends winter months in deeper water and it is thus defined as benthopelagic. Shoals of breeding individuals come inshore in summer to breed, and when they are fished for. Males and females release sperm and eggs into the sea and the fertilised eggs are pelagic. The biggest individuals have been estimated to be five years old and, hence, the curled picarel is a short-lived and fast growing species, attaining 60% of its asymptotic length during the first year of life. Growth is thus very rapid.
The body form of [Centracanthus cirrus _]is elongate with a long single dorsal fin that has thirteen spines and between nine to ten soft rays differentiating the first two-thirds from the last third. The anal fin has three spines and also between nine to ten soft rays. The pectoral fins are much longer than the pelvic fins. The snout of the curled picarel is pointed and the mouth has a protrusible upper jaw and contains a series of rows of small teeth. What _Centracanthus cirrus feeds on is unknown, but from the body form and mouth structure, and bearing in mind that there is no plant life at its inhabited depths, it seems likely that it preys on small invertebrates – most likely crustaceans and polychaete worms.
Because it lives in deep waters where will be little or no light, the curled picarel has very large eyes and the animal is coloured red dorsally whereas the underside is white-yellow. Many deeper water fish are red. This is because light is absorbed progressively with greater and greater depths and in those at which Centracanthus cirrus lives there will be little or no light and red appears black. This will be seen again with the boarfish, [_Capros aper _](page 149), also collected from and illustrated in the Açores.
The Monrovian surgeonfish, or tang, Acanthurus monroviae Steindachner, 1876, is a member of the pan-tropical family of fishes – the Acanthuridae – typically living over coral reefs. Acanthurus monroviae occurs in the Eastern Atlantic from Morocco south to Angola and inhabits the Cape Verde Islands, where the illustrated individual was obtained. Surgeonfishes are characterised by a retractable, scalpel-like, spine on each side of the caudal peduncle or tail base. Surgeonfishes have a deep body and are flattened laterally. They also have a steep forehead with a similarly highly positioned eye. There is a single dorsal fin comprising nine stouter anterior and between 24-27 posterior finer rays. Similarly with the anal fin, which has three anterior stouter and between 23-26 posterior finer rays. The Monrovian surgeonfish varies in colour from brown to dark blue but is characterised by left and right bright patches of yellow on the caudal peduncle. In the centre of each of these patches is a brown horizontal chitinous barb-like spine. Generally, surgeonfishes are peaceable creatures travelling in small schools over coral and other reefs, their diets being based on algae selected from turf communities growing on subtidal rocky surfaces. Some surgeonfishes form schools that migrate daily from their nocturnal refuges in coral reefs to foraging sites on the intertidal, covering distances of more than 500 metres. During summer the main food items are brown and red algae; in winter, lush green algae. This changeover appears to provide the food-base for the accumulation of fat and the recrudescence of gonadial activity. Reproduction occurs in large schools of, sometimes, thousands of fish and over selected sites, where and when they become fiercely territorial.
When attacked, however, surgeonfishes can be not only pugnacious but very aggressive. As they thrash their caudal fin from side to side, retractor muscles connected from the spinal column to each caudal spine are engaged and left and right scalpel-sharp barbs are alternately exposed and retracted. Grabbing or biting it, an unwary natural predator, territorial competitor or careless human would be slashed by the thrashing fish. For this reason, fishermen are careful to de-tail the fish upon capture. The colour yellow is, in the natural world, often associated with danger. It is no coincidence, therefore, that most surgeonfishes identify their caudal areas in such a way. In Australia, the blue-lined surgeonfish, Acanthurus lineatus (Linnaeus, 1758) also has a venom gland associated with each spine, so that any careless handler is not just slashed but also envenomed. It is unknown if Acanthurus monroviae is venomous.
A final interesting point about the Monrovian surgeonfish is that it is not native to the Cape Verde Islands, but has somehow made its way across the Atlantic from the Caribbean, but how or when is unknown. In this new environment, however it has become established to such an extent that the wrasse, the blackbar hogfish Bodianus speciosus (Bowdich, 1825), now recognises it as a client and cleans it when the surgeonfish visits its territory (Quimbayo et al. 2012).
The emperor breams or, more simply, the emperors or large-eye breams, belong to the Lethrinidae, and are found in the tropical waters of the Western Pacific and Indian Oceans, and from where many dozens of species have been recorded (Carpenter and Allen 1989). Conversely, Lethrinus atlanticus[* *]Valenciennes,[_ _]1830 is the only species found in the eastern Atlantic Ocean. This is quite strange, but the combination of teeth types, body shape and scale counts for [_Lethrinus atlanticus _]are very different from other species of the genus. Moreover, this unique combination of characters is consistent among all individuals examined from many locations in the tropical Western Atlantic Ocean. There is little doubt that not only is this the only species of this genus present in the eastern Atlantic Ocean, but its populations are readily inter-mixing and inter-breeding.
The geographical distribution of[_ Lethrinus atlanticus_] comprises a narrow range from 20°N to 5°S and where the sea temperature ranges from 20-28°C. It thus encompasses the tropical waters of the west coast of Africa from Senegal to Gabon, and the island groups of Principe, Sao Tomé and Cape Verde. The illustrated individual was obtained from the fish market in Mindelo on the small island of São Vicente in the Cape Verde’s in January 2011. In Portuguese, the official language of the former colony of Cape Verde, the fish is called bica.
Lethrinus atlanticus inhabits shallow coastal waters to depths of some 50 metres and is primarily an associate of coral and other reefs. Here it is an epibenthic predator of bottom-living invertebrates and small fishes. To do this, it has a long, pointed, snout-like, mouth equipped with molariform teeth, which are used to crush shelled invertebrates, such as molluscs and crabs.[_ Lethrinus atlanticus _]grows to a maximum total length of about 50 centimetres although a more common length is 30 centimetres. It is, nevertheless, a pretty impressive fish particularly because it is prettily coloured olive green or light brown with splashes of pink and orange-yellow especially around the head and fins. Below the eye, there is a network of fine reticulations. The species is said to be non-migratory although it is more abundant in the winter months and so may move down to deeper waters in the summer. Like many fish species that occur in such tropical eastern Atlantic waters, little else is known about this beautiful but nevertheless important food fish.
In 2011, I took a cruise up the Amazon River as far as Manaus, capital of the Brazilian state of Amazonas. There many wonderful things of biological as well as historical features of the Amazon that I wished to see, such as, the black caiman, Melanosuchus niger (Spix, 1825), the Amazon river dolphin Inia geoffrensis Blainville, 1817 (although it is not restricted to this river) and the native Amazon water lily, [_Victoria amazonica _](Poepp.) J.C. Sowerby. I was not disappointed by any of these. Each lily pad, for example, is up to three metres in diameter, with a stalk up to eight metres in length. Something I did not know was that the undersurface of each leaf is a maze of razor sharp spines. I also wanted to see the Amazon fishes but knew that to find these, I would not do so from a riverboat, but needed to access the fish markets.
The Amazon has an incredible diversity of fishes and is the centre of diversity for neotropical species, of which more than 5,600 are currently known. The bull shark, Carcharhinus leucas (Müller & Henle, 1839) has been reported 4,000 kilometres upriver at Iquitos in Peru. The pirarucu, [Arapaima gigas _](Schinz, 1822), is one of the largest freshwater fish in the world, with a reported maximum length of three metres and weight of two-hundred kilogrammes. Also present in large numbers are the world famous piranhas. There are estimated to be from 30 to 60 species of piranha. The most notorious of these is the red or red-bellied piranha, _Pygocentrus nattereri Kner, 1858.
The red-bellied piranha is native to South America and is found in the basin of the Amazon River, other rivers of northeastern Brazil, and the basins of the Paraguay and Paraná Rivers. The red-bellied piranha congregates in large schools and has a reputation as a ferocious predator and has been known to attack livestock and even humans, resulting in deaths. Stories of large schools of red piranhas attacking humans, however, are exaggerated. In reality, they are generally timid, their diet typically consisting of other fish, insects, worms and crustaceans. In contrast to their popular reputation of feeding on living prey, therefore, red piranhas are primarily scavengers usually feeding on dead, dying, and injured animals.
A good specimen to draw was found in the fish market at Almeririm, half way between the sea and Manaus. They have a ferocious appearance, which has probably contributed to their reputation although in fact the mouth is small relative to the overall body size. The tiny teeth inside it, however, are razor sharp. As their name suggests, red-bellied piranhas have a reddish tinge to the belly when fully grown, although juveniles are a silver colour with darker spots. They grow to a maximum length of 33 centimetres and a weight of three kilogrammes and, again, contrary to popular legend, are not a highly prized restaurant fish.
The fish usually feed in large schools around dusk and dawn and locate their prey by sight with their large eyes, scent or motion using the lateral line system – a set of mechanreceptors down each side of the body. Red-bellied piranhas will sometimes bite one another, normally on the fins, in a behaviour called fin-nipping. Fish that have had their fins nipped will grow them back surprisingly rapidly. Such behaviour probably results from their schooling behaviour when, excited by the presence of food, this stimulates them to bite at anything in front of them.
Red-bellied piranhas usually spawn around April and May during the rainy season when the swollen river floods the Amazon’s surrounding and gigantic flood plain. The male digs out a nest in rocks and vegetation and awaits an attracted female. Females, with more yellow on the belly than the redder males, can lay up to 1,000 eggs in the nest which the male then fertilises. Males become extremely territorial during spawning, and will aggressively prevent other fish from approaching the nest. After the eggs hatch, both parents guard the broods. Tourist shops dry any species of piranha and imaginatively paint their bellies bright red to sell to the gullible.
At a fish market in Anteririm on a cruise up the Amazon River in 2011 I found my first armoured catfish – Haplosternum littorale (Hancock, 1828). Amazingly, there are, worldwide, over 3,000 species of catfish and of the 2,800 or so species of fishes recorded from the Amazon, almost half (about 1,200) are catfishes. New species are being discovered from this mighty river all the time, for example, the armoured giant Brachyplatystoma capapretum (Lundberg & Akama, 2005). In the Amazon, the piraiba, Brachyplaystoma filamentosum (Lichtenstein, 1819), grows to a length of three metres and can weigh 134 kilogrammes, that is, the weight of two big men.
Haplosternum littorale, however, only grows to a length of 24 centimetres. During the reproductive season, males deposit fat in their pectoral areas and develop a pair of elongated upcurved spines. Haplosternum littorale, is locally called tamuatá and has the widest distribution of any callichthyid catfish. It ranges from Venezuela and Guyana to Argentina and occurs in all sub-tropical South American waters east of the Andes and north of Buenos Aires, including the Orinoco, Trinidad, the coastal rivers of the Guiana’s, the Amazon River drainage, Paraguay, the lower Paraná River, and coastal estuaries in southern Brazil. The species has also been introduced into the Indian River Lagoon of Florida (Nico et al. 1996).
Haplosternum littorale is restricted to swamps where it feeds mainly on benthic invertebrates, such as insects, microcrustaceans and detritus (Winemiller 1987). The catfish uses the two pairs of barbels on its upper and lower lips and the array of sensory structures on its face to detect its prey in the swamps. Such swamps are also characterised by a marked seasonality caused primarily by fluctuations in rainfall, low levels resulting in low dissolved oxygen levels. To cope with this, Haplosternum littorale can breathe with its gills and through its intestinal wall. Juveniles possibly absorb oxygen through their skin, before the armour plates, or scutes, that cover the body develop. Soon, older juveniles develop the capacity to breathe air (Persaud et al. 2006).
Of most interest, however, is the way in which[_ Haplosternum littorale ]reproduces (Andrade and Abe 1997, Hostache and Mol 1998). At the age of one, spawning is triggered by rising temperatures and the first rains. _Haplosternum littorale builds a complex, dome-shaped, bubble nest some 30 centimetres in diameter and six centimetres in height in the peripheral areas of newly-flooded swamps. In the hypoxic waters of these tropical swamps, the nest is rich in oxygen, the bubbles aerating the eggs by lifting them above the water surface while protecting them from desiccation. It may also serve to protect the developing brood against predators, regulate temperature and identify the centre of the male’s territory, which is defended vigorously, using the enlarged pectoral spines
The beginning of nest building is preceded by a courtship ritual. This pair formation consists of the male and female swimming parallel to each other, facing each other, making barbel contact, with the male stimulating the flanks of the female, and swimming to the surface where they produce the first bubbles at the chosen nest site. The male produces most of the foam. Firstly, the pair come to the surface and swim belly-up in small circles. The air-water interface film is swallowed and pumped out through the gills, where it gains mucus. The pelvic fins stir the water and mucus, capturing air bubbles, and making them into a foam. The male often dives to the bottom to collect filamentous plant debris, which he knits together in the nest using his mouth and pectoral fins, and incorporates the plant material into the mass of foam. The end result is a dome-shaped nest made up of loosely interwoven plant material held up to the surface with a layer of foam.
During spawning, the male and female form a T-position, where the female places her mouth over the male’s genital opening and drinks his sperm. Amazingly, fertilisation takes place after the sperm has passed through her digestive tract. Subsequently, the female swims to the foam nest, turns upside down, and lays the eggs in it. The[_ _]females do this up to fourteen times during the seven-month long breeding season, each spawn comprising up to 9,000 eggs. Typically, between two to four females spawn simultaneously, resulting in an average number of 20,000 adhesive eggs per nest. The male maintains and regularly re-supplies the nest with foam and guards it day and night. Once spawning has been completed, the male drives the females away using its large pectoral spines to abrade and slash at them and any and all intruders.
There are approximately[* *]25 species of flying fish and they all occupy tropical, oceanic, waters where the sea surface temperature ranges between 22-24ºC. They belong to a family of bony fishes called the Exocoetidae. The word Exocet was obtained from this family name, that is, supposedly, flying just above the waves. This is incorrect, however, and the term actually means sleeping outside, that is, asleep (or dead) on a ship’s deck. Not of course on modern commercial vessels but on the earliest lower-decked vessels. Flying fish fly just above the surface of the sea and do so for between 40-50 metres – perhaps as much as 100 metres. They do this to escape predators. This is because in the world’s oceans most predators would be other fish such as fast swimming species of tuna. These animals hunt visually only in water and so any fish prey that can leave this environment, even if only briefly, literally disappears for a few seconds. Most human beings only see flying fish from the prow of cruise vessels and are naturally enchanted to see the little creatures fleeing from the ship. They flee because, to them, the ship is simply a large predator.
One of the commonest flying fishes in both the tropical Atlantic and Pacific Oceans is the two-winged flying fish, Exocoetus volitans[* ]Linnaeus, 1758,[ *]which grows up to about 30 centimetres in length. To fly, size is restricted by weight, so that if an animal is too big, it is unable to leave the water. Thus, whales, especially humpbacks, emerge from the sea only to breach and this is in order to crash back in making a sound that travels far. In contrast, the much smaller dolphins are able make spectacular jumps – but not fly. In fact, few marine animals can leave the sea surface. Great white sharks and manta rays can make vertical jumps, the former in pursuit of its similarly leaping prey, seals and sea lions. Salmon can leave water, but do so only to traverse obstacles on their return from the sea as adults to breed in their home freshwaters. In estuaries and shallow coastal seas, shoals of juvenile mullet of many species engage in frenzied attempts to escape larger predators such as mackerel and bass by making emergent jumps but only the flying fishes can actually fly.
To do this, flying fishes are modified remarkably from the basic fish plan. Firstly, they have, like all fast swimming fishes, a bullet-shaped body that offers the least friction to swimming. Exocoetus volitans is, moreover a brilliant blue dorsally and silvery white below so that it is difficult to see against the deep blue ocean waters from above and the sparkling sea surface from below. All flying fishes have typical dorsal fins but the pectoral wings are extremely long and when spread out have a parabolic form, that is, they are ventrally concave and thereby act like lateral parachutes creating lift. Some species of flying fishes only have these wings but, in Exocoetus volitans, the pelvic fins are also enlarged and when expanded create more lift. Seen from the side, the caudal or tailfin is also unusual in that the ventral component is larger than the dorsal. If one looks at the tail fin of a shark, it is organised the other way round and is called heterocercal. That is, when the tail fin beats from side to side, the dorsal component will create the greater force and tend to drive the animal downwards to the sea bed, which is where most sharks feed. Hence the ventrally large tail fin of the flying fish is a reverse heterocercal and tends to drive the animal upwards when in motion. This is, of course, because flying fishes live in the surface waters of the sea and feed on tiny planktonic creatures (Lipskaya 1980).
The reverse heterocercal tail, however, also fulfils another function. It constitutes the propulsive force that drives the animal up and out of the water and once out, keeps the fish moving over the surface by acting as a fast rear paddle. A shark’s heterocercal tail makes it impossible for them to fly – they would simply be driven downwards. Flying is, however, an energy-consuming activity, especially for a fish and Exocoetus volitans can only make up to three consecutive emergences from the water and is then exhausted. But, this alone would take our little fish up to 100 metres away from a pursuing predator. Flying fishes escaping from a ship’s prow are able to steer to left or right by adjusting the angle of their pectoral wings.
In the Caribbean, flying fish constitute a popular component of local people’s diet. The flying fish is also the national symbol of St Lucia, where the species was drawn.
The solitary carabid beetle Bembidion laterale (Samouelle, 1819) lives towards the rear of rocky but especially sandy beaches (Andrewes 1938) from Germany southwards to the Iberian Peninsula and Morocco. It is widely but locally distributed in Great Britain and Ireland (Good 1998). On the River Arun there is a patch of surviving sand and Bembidion laterale occurs here towards the rear just below a landward fringe of the sea purslane [Halimione portulacoides _](L.) Aellen. _Halimione portulacoides is a small, greyish-green, shrub widely distributed in temperate Eurasia and parts of Africa. It is a halophyte and is characteristic of salt marshes and is usually flooded at high tide. The plant grows to a height of 75 centimetres and is evergreen flowering here on the River Arun from July to September. The flowers are monoecious and are pollinated by wind. The leaves are edible and can be eaten raw in salads or cooked as a potherb. The occurrence of this plant here in such profusion attests to the fact that the River Arun, now canalaised, once here at Littlehampton, possessed a large estuarine marshland system (Morton and Bamber 2012)
Bembidion laterale either builds chambers and galleries in sand or lives in crevices in rocks and from which it ventures at low tide and hunts over the sand surface (Elliott et al. 1983). It is dorso-ventrally flattened and has large prognathous mandibles, which give a clue to its lifestyle. For, on the River Arun beach it feeds on the burrowing amphipod [Corophium arenarium _]Crawford, 1937,[ ]which may reach a length of seven millimetres, but still small enough for [_Bembidion laterale _]to handle. [_Corophium arenarium _]emerges from its burrow that may be between 10 and 60 centimetres deep, as the tide is falling, and leaves long meandering tracks over the sand surface. _Bembidion laterale tracks its prey here on the surface and at these times[_ ]because _Corophium arenarium burrows too deeply to be found there and retreats into its abode as the tide rises. [Bembidion laterale _]and its prey, thus have an endogenous tidal rhythm, the returning tide sending both back to their galleries and burrows, respectively, higher up the shore. Unlike many other littoral insects, however, [_Bembidion laterale _]can withstand long periods of submergence in seawater (Elliott _et al. 1983) – up to 17 hours, so that of all the British intertidal insects it is one of the most highly adapted to such a lifestyle.
The four-horned spider crab Pisa tetraodon (Pennant, 1777) is a member of the Majidae, a family containing some 200 species that also includes the long-legged spider crabs and the decorator crabs. The dorsal carapace of Pisa tetraodon is longer than it is broad, making it roughly sub-triangular in shape, and forming a forceps-like rostrum anteriorly, each distal half diverging outwards, at the front. The tiny eyes are set laterally, partly concealed when retracted, behind the rostrum on stalks. The lateral margins of the lumpy, bumpy, carapace each possess five spines. Typical of spider crabs, there are eight long legs as well as two chelipeds, or claws. The distal end of the first limb segment of each of the walking legs has distinct knobs – giving the appearance of knobbly knees. The chelipeds are of approximately equal size, simple and finger-like to be used for extracting food items out of small places. Unlike the usual view of crabs as scurrying, nipping little monsters, [_Pisa tetraodon _]is ponderous, harmless and not a little whimsical.
The illustrated male individual was collected from the Littlehampton’s East Beach, West Sussex, England, along with other conspecifics, after a storm in February 2012. The offshore sea bed of the English Channel comprises, in places, at depths of around ten to twenty metres, rocky limestone outcrops that are shallow enough to be covered in seaweeds. At these depths, mostly red rhodophytes and corallines. This is a perfect habitat for decorator crabs including Pisa tetraodon. Ovigerous (egg-carrying) female individuals of Pisa tetraodon occur from April-May. Upon release, the fertilised eggs develop into planktonic larvae, the life cycle of which has been described by Rodriguez (1997).
[Pisa tetraodon _]has a narrow latitudinal distribution occurring in the Mediterranean Sea, the Atlantic coast of Portugal and Spain and northwards to the English Channel, encompassing the coastline of western France, the Channel Islands, all of the southern coast of England but only up into the North Sea as far as the coast of Norfolk. The range also includes parts of the west coast of Great Britain, but rarely Wales and even more rarely as far north as the west coast of Scotland. The biology of [_Pisa tetraodon _]in the Mediterranean has been described by[ _]Vernet-Cornubert (1958). There, species may reach a carapace length of 45 millimetres, as it does too on the Channel coast of England.
[Pisa tetraodon _]is a decorator crab. Not all species of the Majidae are decorators but those that are use materials from their environment to hide from predators. _Pisa tetraodon sticks, typically, pieces of seaweed onto its back to effect its camouflage. To achieve this, the dorsal surface of the carapace especially is set with clusters of short hook-shaped setae which hold in place encrusting growths of epiphytes, including pieces of seaweed cut away from parent plants using the chelipeds and epizooites. In the case of the latter, decorator crabs stick mostly sedentary animals to their bodies, some of which may be noxious organisms. Where harmless organisms are used for camouflage, this avoidance of detection is called crypsis. Where there is a tendency to become noticeable using noxious or toxic organisms for defence, the relationship is called aposetism.
Decorator crabs gain concealment by taking on masks of adventitious material. The best example of this is provided by the spider crab [Hyas areneus _](Linnaeus, 1758).[ ]When individuals of this spider crab[ ]were moved from a habitat where they all camouflaged with short pieces of seaweed into different habitats, all of them redecorated themselves with local materials, as quickly as after one night. Other species of decorator crabs are highly specialised in their choices of camouflage. For example, the southern Californian _Pelia tumida (Lockington, 1877) decorates itself only with sponges. Virtually nothing is known about the decorating biology of Pisa tetraodon.
The boarfish [_Capros aper _](Linnaeus, 1758) is member of the Caproidae. [_Capros _]comes from the Latin [_capra _]meaning either goat or wild boar. The species is not un-common in the Açores where, in the summer of 2011, a research workshop was convened in the fishing village of Mosteiros. [_Capros aper _]can reach a maximum length of 30 centimetres. More commonly, however, it only grows to 13 centimetres. It occurs in the Eastern Atlantic from western Norway and the Skagerrak, south to the Shetlands Isles and western Scotland and down to Senegal. It also occurs in the Mediterranean, mainly the western part, and in the Canaries and the Açores. It is especially abundant on the edge of the continental shelf at depths of about 100 metres in the south-western approaches to the English Channel and at similar depths off Ireland and southern Scotland. It is also abundant off Portugal and the Açores but is uncommon in shallow waters. In the Açores, it is not a targetted fish, but is caught, as by-catch, along with other species in gill nets.
Capros aper is wholly marine and demersal, that is, it lives close to the seabed at depths ranging from 40 to 700 metres. It is gregarious, forming schools, or herds, just above the sea bed and is typically found over rocky or coral substrata where it, is said, feeds head down on crustaceans, especially copepods and mysids, worms and molluscs. According to Morato et al. (2003), small shoaling fish, including the boarfish, are major components of the diet of the tope shark, Galeorhinus galeus (Linnaeus, 1758), in waters around the Açores and which is, itself, an important long-line artisanal fishery resource.
The general body form of the little red and silver boarfish is rhomboidal, quite deep and laterally compressed. The dorsal fin comprises two components: a spiky first dorsal and a vibrating second dorsal component and second anal fins. The caudal fin is spatulate with a convex edge. The ventral fins are large and spikey, while the pectoral fins are quite small. The body is orange/red, sometimes with three wide lighter vertical bands. Males are typically smaller than females. During the breeding season there is a striking sexual dimorphism. The male’s body becomes covered with sinuous orange lines and the dorsal and ventral fins turn red. The female becomes orange with a faint dark band in the middle of the body. The belly and basal part of the anal fin become white/silver, while the ends turn dark orange. The anterior component of the dorsal fin has 9-10 spines while the posterior component comprises between twenty-three to twenty-five soft rays while the anal fin has three spines and between 22 to 24 soft rays.
This species was drawn because it appeared strange, such weird animals often being found at great depths. The orange-red body also suggested it had been caught from a great depth. This is because this colour is well suited to deep water as the longer wave length of red light are absorbed by seawater before the blue and greens, which are reflected. The boarfish would thus appear black at the depths it inhabits. These thoughts were supported by the observation that Capros aper has very large eyes in relation to its body. The boarfish, like the John Dory, [_Zeus faber _]Linnaeus, 1758, has a protrusible mouth, forming a short tube when extended, and which is used, like a suction-device, for catching its small prey items. Another feature of the boarfish is that, unusually, it can swim backwards as readily and even more effectively and speedily than it does forwards.
The boarfish gained momentary fame in the European Parliament meeting in Strasbourg on 6 February 2013 to discuss reform of the Union’s common fisheries policy. Amongst other measures, the parliament’s members voted to ban the practice of discarding undersized and unwanted by-catch, fish caught by trawling. With, one exception – the, so called, worthless boarfish. Worthless, because the boarfish has been considered historically rare, with only temporary periodic increases in abundance. Its abundance has, however, increased of late in the North East Atlantic and a modern important industrial fishery has developed around it (Blanchard and Vandermeirsch 2005). Its apparent late age-at-maturity may, however, indicate its susceptibility to over-exploitation (White et al. 2011) and, subsequent to the European Union’s decision, this has become true.
Atherina is a genus of Atherinidae, or the silversides, which are found worldwide in temperate and tropical waters. There are currently five recognised species of Atherina, with the Mediterranean sand smelt Atherina hepsetus Linnaeus, 1758 being widespread in the Mediterranean, Black Sea and Sea of Azov, occurring in lagoons and the lower reaches of estuaries. Silversides are relatively small, with Atherina hepsetus reaching a length of approximately 15 centimetres. The body is elongate. Distinctive characters include two widely-separated dorsal fins of approximately equal size, with the first consisting of flexible spines and the second having one spine followed by soft rays. The anal fin is situated directly below the rearmost dorsal fin, and is of approximately the same size and, similarly, has one spine on the leading edge followed by soft rays. The pectoral fins are located high on the body and are small.
Seen from the side, the ventral component the caudal or tail fin of Atherina hepsetus is slightly larger than the dorsal. This is the opposite to the situation seen in most sharks, where, the upper half is slightly larger than the lower. That is, when the tail fin beats from side to side, the dorsal component will create the greater force and tend to drive the animal downwards to the sea bed, which is where most sharks feed. Such a tail is termed heterocercal. Hence, the ventrally larger tail fin of the sand smelt is a reverse heterocercal and tends to drive the animal upwards when swimming. The advantage of this is that such an upward swimming trend takes the smelt towards the sea surface where at night, especially, its food which comprises tiny planktonic creatures reside. Such little prey move upwards at night, this being termed diurnal vertical migration, and this is when the shoals of sand smelt also swim towards to feed on the zooplankters while they are concentrated here near the surface. To assist in this too, the mouth is directed upwards and the eyes are large. The scales of these small fish are relatively large and the body is silver below and a darkish green above. Such a coloration is typical of shoaling pelagic, surface dwelling, fish but, as the colloquial name suggests, the flanks comprise a broad and distinctive silvery band. Unusually, there is no obvious lateral line.
Atherina hepsetus was drawn from specimens collected from a fish market in Pafos on the western end of the island of Cyprus in December 2011. Because of the long relationship between the United Kingdom and Cyprus, many expatriates live on the island. As a consequence, in the local markets, the fish is advertised as whitebait, a British favourite deep-fried in a light batter. Whitebait itself, however, comprises different species wherever it is sold throughout the world and surveys of whitebait sold in London fish markets have shown it to comprise the mainly juvenile phases of up to twenty-three fish species. The usual constituent of British whitebait is, however, the sprat, Sprattus sprattus illustrated earlier (page 121).
The common cockle,[_ Cerastoderma edule_] (Linnaeus, 1758), occurs in coastal areas of the eastern Atlantic Ocean. Its synonymised name of [Cardium edule _]Linnaeus, 1758[ ]is still often used. The species is distributed widely from Iceland in the west to Norway, through the Baltic and North Sea down to the coast of West Africa to Senegal. It lives all around Great Britain and Ireland. but does not occur in the Mediterranean. The common cockle is one of the most abundant species of intertidal flats in the bays and estuaries of Western Europe. It is an important fishery species but is also a source of food for crustaceans, fish and wading birds, especially eiders, [_Somateria mollissima _](Linnaeus, 1758), and oyster catchers, _Haematopus ostralegus (Linnaeus, 1758), the latter being a significant predator, preferring cockles of 20-30 millimetres shell length. The cockle is fished commercially in Ireland, Great Britain and France by hand raking and suction dredgers. It is also of aquaculture importance such that cockle farming is ongoing in Great Britain, The Netherlands and Portugal. For this reason, studies of its life cycle are especially important and a newly-settled juvenile[_ ]of _Cerastoderma edule from Traeth Melynog, Menai Strait, Anglesey, North Wales, is illustrated.
The adult Cerastoderma edule has a solid, robust, globose, shell usually up to 45 millimetres long, but occasionally over 50 millimetres and somewhat broadly oval in outline. The outer surface of the adult shell is off-white to yellowish-brown. Each valve has between 22-28 thick, prominent, flat-topped and radiating, ribs each with numerous scale-like spines. The ribs create a crenulate and interlocking shell margin. Concentric, annual, growth lines are also prominent, the species living for up to nine years. An important aspect of the near-spherical shell, with interlocking valve margins is that it is difficult to pick up and access by, in particular, wading birds. That is, it is a defensive measure, allowing it to live close to the sediment surface where the shape also means that it is difficult to be dislodged by waves.
Cerastoderma edule typically lives in soft stable sand or mud typically low on the shore but down to a depth of 200 metres. In sheltered bays and outer estuaries, densities of 10,000 per square metre (mostly juveniles) have been recorded where it filters water and collects fine particles of organic matter. It has been calculated that an adult cockle can filter a half litre of water each hour. It, therefore, takes but a brief time for the large cockle populations to filter the incursing tidal waters and, thereby, cleanse them.[* *]The siphons possess tactile tentacles and around the mantle margin are simple eyes. These detect movements in the water or air above causing the shell valves to shut firmly. The tentacles are mechanoreceptors so that when the cockle is touched by a sea star, [Asterias rubens _]Linnaeus, 1758, or the whelk, _Buccinum undatum (Linnaeus, 1758), it can, by quickly extending the foot and undertaking leaping movements, achieve a rapid escape.
Cerastoderma edule reproduces in Great Britain from late February to early July. Sexes are separate and sperm and ova are released into sea where fertilisation occurs. Eventually, after two to three weeks in the sea as, firstly, a trochophore and, secondly, a veliger, these metamorphose and settle onto the substratum as a juvenile. The juvenile shell, barely one millimetre long, is nearly transparent.
The king angelfish, Holacanthus passer Valenciennes, 1834 is a large, up to 35 centimetres in length, member of the Pomacanthidae. It is non-migratory and inhabits tropical reefs in the eastern Pacific Ocean from the coast of Peru north to the Gulf of California, including offshore archipelagos such as the Revillagigedo Islands (Mexico), Malpelo Island (Colombia), Cocos Island (Costa Rica) and as far west as the Galapagos (Ecuador), at depths ranging from 1-80 metres.
The illustrated adult was drawn from underwater photographs taken in the Galapagos Islands, Ecuador, in July 2012. This is the second-most abundant species in the Galapagos Archipelago, with an average of 13 individuals per 500 square metres (Edgar et al. 2004). It seems to be most abundant over the seabed covered with corals and coralline algae, intermixed with sandy and rocky substrata, and including large crevices.
Sexually mature adults are dark blue-grey with a yellow dorsal fin, blue rimming around the fins and have a vertical white bar on the sides just behind the origin of the pectoral. There is also a crown of bright blue spots on the forehead and a yellow-orange tail fin. Juveniles undergo a progressive change in colour patterning as they mature are primarily yellow, with iridescent-blue-rimmed fins and pale blue-white striping towards the posterior region of their bodies, and an orange mask around the eye, an orange front and tail.
Angelfish are sexually isomorphic, meaning males and females are visually identical externally. Male Holacanthus passer are larger than females although the sexes can be separated because[_ ]females have yellow pelvic fins, unlike the white ones of males, and are more territorial forming pair bonds with the males. Each individual has a strong spike under their lower cheek for defensive purposes and for maintaining a territory, which is performed aggressively. _Holacanthus passer is within-pair monogamous and, over the spawning cycle, will mate daily around sunset. During each spawning event, a pair can produce upwards of ten million fertilised eggs, averaging about 25,000–75,000 daily. The fertilised eggs drift in the water column for about twenty hours, at which point they hatch. After hatching, the fin-less fry live off their yolk sac until it is absorbed, at which point they begin to eat small zooplankton. As adults, king angelfishes are diurnal feeders, cropping algae, sessile invertebrates but seem to specialise on sponges. It has been reported that adult king angelfish also clean parasites from sharks, including hammerheads.
Nephrops norvegicus (Linnaeus, 1758)[* *]is known variously as the Norway lobster, Dublin Bay prawn, langoustine, and scampi in England. It is a slim, orange-pink, lobster-like, wholly marine, crustacean superficially similar to the European crayfish, Astacus astacus (Linnaeus, 1758) but which is freshwater. Both are located in their own families – the Nephropidae and Astacidae, respectively – the former possessing but one species, which grows up to 25 centimetres long, and is the most important commercial crustacean in Europe.[_ ]Around 60,000 tonnes of _Nephrops norvegicus are landed annually, half of it caught in British waters and mostly by trawling. Nephrops norvegicus lives in the north-eastern Atlantic and North Sea as far north as Iceland and northern Norway, and south to Portugal and parts of the Mediterranean, notably the Adriatic, but is absent from the Baltic and Black Seas.
Nephrops norvegicus has the body shape typical of a lobster, albeit narrower than the large European lobster Homarus [gammarus _](Linnaeus, 1758), which can grow to 60 centimetres in total length. It is, overall, a pale orange in colour, but with bright red chelae especially, and grows to a typical length of between 18 to 20 centimetres, including the tail and claws. In the United Kingdom, scampi refers to the tail, or abdomen, after removal of the external, chitinous, skeleton. A carapace covers the cephalothorax, while the abdomen is long and segmented, ending in a broad tail fan or telson. The first three pairs of legs bear claws. Of these, the first are bright red and white, greatly elongate and bear robust ridges and spines. Of the two pairs of antennae, the outer second is the longer and thinner. There is a long, spinous rostrum, and the compound eyes are kidney-shaped, providing the name of the genus, from the Greek roots of _ nephros (kidney) and ops (eye).
Nephrops norvegicus adults prefer to build burrows in muddy seabed sediments, with more than 40% silt and clay. Due to its ecological demands for such particular sediments, Nephrops norvegicus has a very patchy distribution, and is divided into numerous populations separated by inhospitable terrain. As a consequence, adults rarely travel distances greater than a few hundred metres. The burrows are semi-permanent, and are twenty to thirty centimetres deep, with a distance of 50 to 80 centimetres between the front and back entrances. Nephrops norvegicus individuals spend most of their time either in their burrows or by the entrance, only leaving to forage and mate. Adult Nephrops norvegicus are predators and scavengers that emerge from their burrows at night to make short foraging excursions.
The life span of Nephrops norvegicus is usually five to ten years, but perhaps fifteen in exceptional cases. Adult male Nephrops norvegicus moult once or twice a year, usually in late winter or spring, and adult females moult up to once a year, in late winter or spring, after egg hatching. The ovaries mature throughout the spring and summer months, and egg-laying takes place in late summer or early autumn. After spawning, the berried females return to their burrows and remain there until the end of the incubation period. Hatching takes place in late winter or early spring. Soon after hatching, the females moult and mate again. In annual breeding cycles, mating takes place in the spring or winter, when the females are in the soft, post-moult, state. The structure of the reproductive cycle, however, varies depending on geographical position, the spawning timings, incubation length and hatching periods varying with latitude. As a consequence, the breeding cycle changes from annual to biennial from south to north.
In October 2012, on a visit to the School of Biological Sciences of the University of Hong, I was asked by a local Ph.D. student if I could spare some time to examine an animal that had settled on her marine fouling test panels immersed in the sea at Clearwater Bay on the eastern, oceanic, side of Hong Kong’s New Territories. The animal showed me was not one but many, all tiny, no bigger than 200 μm in overall dimensions. Each individual comprised a bulging, basal, component and an erect tube arising at 90o from it and which apically had up to three growth collars. The animal inside the tube was yellow-green and comprised a body located in the swollen basal ampulla. And from which arose a pair of elongate, flat, peristomial lobes both pointed at each apex. Between the bases of the two lobes appeared to be a mouth/anus. When disturbed, the lobes were withdrawn in a flash into the deepest base of the tube, right to the end of the ampulla.
At first it was thought this weird animal must be an exceedingly simple hydrozoan, that is, a sessile animal related to the jellyfishes and anemones. But, unhappy with this diagnosis, a simple illustration was made of it and sent it to some Chinese and British colleagues who might be able to identify it. Two answers soon came back both of whom identified my creature not as belonging to the Kingdom Animalia but as a member of the Kingdom Chromista – the single-celled organisms. Furthermore, it was a member of the Ciliophora (Heterotrichida), the latter containing some 160 species. Narrowing the identification down, it was suggested that the tube-dwelling ciliate belonged to the Folliculinidae, with 72 species and, specifically, the genus [_Folliculina _]with but a handful of species.
There are around three British species of [Folliculina _](Das 1949), one of which mine seemed to match, that is, [_Folliculina viridis _]Wright, 1862. This is because its body is yellowish-blue-green, the transparent tube, or lorica, is angled at 90o to the ampulla and terminates in a series of distinctive collars. Other species of _Folliculina have been recorded from China, notably Folliculina simplex Dons, 1912 (Song et al. 2003, 2009). Huang and Lin (2012) illustrate Folliculina simplex from China and it is clearly different from the species identified from Hong Kong, that is, Folliculina viridis.
Such a discrepancy is not, however, of great significance because it seems highly possible that all of the putative species of this ciliate genus have been introduced and re-introduced all around the world possibly mainly by being carried as larvae in ship ballast water, but also naturally attached to the swimming or floating larvae of other marine creatures, such as those of shells that would give the eventually-formed adult something solid to attach securely to (Scheltema 1973).
The most impressive thing about Folliculina viridis, however, is that this seemingly complex creature actually comprises but one cell the oval nucleus of which can be seen inside the body located inside the basal ampulla. The peristomial lobes of the body possess ciliary tracts, which collect the finest particles of seston suspended in the water column and transport them down to an adoral zone of membranelles and which perform two and one half turns within the buccal cavity to form what is, to all intents and purposes, a mouth. Except this mouth leads directly into the cell’s basal interior. Perhaps weirdly too, the mouth functions as an anus releasing the waste products of intra-cellular digestion back into the water as microscopic, hair-like, threads.
The most fascinating aspect of this protozoan, however, is that this single-celled ciliate, has organelles that mirror the true organs of the animal representatives of the Hydrozoa. The two are, however, wholly unrelated – both having, as single- and multi-celled creatures, respectively, adopted a suspension feeding mode of life in the sea, encased in a protective tube.
The city of Jinhong is situated on the Lancang River that is, itself, a tributary of the Mekong River. Jinhong is the capital of the Xishuangbanna Dai Autonomous Prefecture, of China. The Lancang River arises in southwestern China from the foothills of the southeastern terminus of the Tibetan Himalayas. At Jinghong, the Lancang River is wholly freshwater. From its origins in China, the Mekong River flows along a meandering 4,600-kilometre course toward the South China Sea, passing through Myanmar, Laos, Thailand, Cambodia and Vietnam. The Mekong River Basin is huge (800,000 square kilometres) and its watershed is home to about 65 million people, many of them living in rural areas and relying on subsistence fisheries for their diet. Apart from its important role as a food source, the Mekong is also a biodiversity hotspot for many species but, in particular, fishes. The Mekong River is the second most biodiverse river in the world after the Amazon and 877 fish species (not including estuarine species) have been identified from it. Regional species richness ranges from only 24 species in the Chinese headwaters to the far north to 484 species in the Mekong Delta.
Inspecting the fishes for sale in Jinhong’s market, a most unusual species was encountered and which had obviously been caught locally. At first inspection, it appeared to be a sardine. But that would be impossible so far from the sea. Closer inspection showed it to be a catfish – Clupisoma sinensis Huang, 1981. The species occurs in the main stream of the Mekong as far upstream as Yunnan Province, where Jinhong is located, in China. It is, however, also, reportedly, found in the Pahang River, Malaysia, but I find this a strange disjunction. The catfish belongs to the Schilbidae and the genus [Clupisoma _]was described by the English naturalist W.J.[ ]Swainson in[ ]1838. In Latin, _clupea means sardine, which suggests that Swainson (1789-1855) was also fooled by the fish into thinking it was a sardine.
Wholly unlike the general image of a catfish, [Clupisoma sinensis _]has a sardine-shaped body, that is, it is streamlined and reaches a maximum length of 31 centimetres (Huang, 1981). It seems to occur throughout much of the Mekong being recorded from Cambodia (Rainboth 1996), Laos (Kottelat 2001), Burma (Ferraris 2004) and south-western China (Chen _et al. 2005). The genus comprises but nine species, distributed throughout Asia, the last one to be described being Clupisoma nujiangense Chen et al., 2005, also from south-western China.
Virtually nothing biological is known about these catfishes, but if we examine the animal itself, it can be seen that it has a large, propulsive, tail fin, two dorsal fins, the anterior-most being larger than the posterior, and a long anal fin comprising between 40-50 soft rays. When the head is examined, however, the catfish attributes are realized – there being four pairs of long thin barbels. The first pair is short and arise from the top the head. The second pair is the stoutest and longest and arise from the upper lip of the mouth. The third and fourth pairs arise from the ventral surface of the lower jaw and are easily missed because they are thin and contained within longitudinally aligned slots. The eyes are large and situated on the sides of the head suggesting that the animal swims above the bed of the river seeking bottom-dwelling food. The species is said to be oviparous, that is, females lay eggs on the river-bed, and which males fertilise, but they are not guarded thereafter.
There are almost 200 herring species, all of which belong to the the Clupeidae and, thus, share several distinguishing characteristics. Generally, herring are silvery, fusiform, streamlined, fish with a single dorsal fin starting at the middle of the back, no lateral line and a lower jaw that protrudes beyond their upper lip when the large mouth is closed. Typically, adult Atlantic herrings attain a length of almost one metre and weigh half a kilogramme. The Atlantic herring, Clupea harengus Linnaeus, 1758 is one of the most abundant fish species on Earth. It is deep blue or blue-green above with silver sides and under-body. It is laterally compressed with a deeply forked tail fin and large scales that easily come free when touched.
The Atlantic herring ranges over much of the northern part of the Atlantic Ocean, from the Bay of Biscay northward to Iceland and southern Greenland. It extends north-eastward to Spitzbergen and Novaya Zemlya in the Arctic Ocean, as well as into the Baltic. To the west, it ranges along the east coast of North America, from south-western Greenland and Labrador, down to South Carolina. Around British waters, the Atlantic herring occurs in the English Channel, the Irish Sea and the North Sea. Although Atlantic herring are found in northern waters surrounding the Arctic, they are not considered to be an Arctic species. Rather, they commonly live in coastal and continental shelf waters congregating in large schools. They can grow up to 45 centimetres in length and weigh more than half a kilogramme.
Herrings are considered to be the most important fish group on the planet. They are also the most populous. Atlantic herring are amongst the most spectacular schoolers and when they do it is in groups that consist of thousands to hundreds of thousands or even millions of individuals. Herring schools in the North Atlantic can occupy, it is estimated, up to 4.8 cubic kilometres with densities of between of 0.5 and 1.0 individuals per cubic metre. That is, several billion fish in one school. In such numbers, schools of juvenile herring in particular, ram feed close to the surface and are the dominant converter of zooplankton into fish, consuming copepods, arrow worms chaetognaths, amphipods, mysids, krill and fish larvae in the pelagic zone. Herrings can also filter-feed if there are sufficient densities of its prey to allow this. Gill rakers in their mouths filter incoming water, trapping any zooplankton and phytoplankton
In such numbers, they have their own predators, which include squid, cod and other larger fish, including salmon and tuna, sharks, porpoises, seabirds of many species, dolphins, seals, sea lions and whales, inclding the killer whale, [_Orcinus orca _](Linnaeus, 1758), and the humpback whale, [_Megaptera novaeangliae _]Borowski, 1781. The Atlantic herring has excellent hearing, and a school reacts quickly to evade such predators. Schools also keep a wary distance from a moving predator, like a killer whale, forming a vacuole which looks like a doughnut from above. Schooling, reduces the risk of an individual being found, predator confusion, better orientation, and synchronized hunting. Herring schools can keep up a constant cruising speed of 360 kilometres per hour, and much higher escape speeds.
Clupea harengus is a migratory species, with schools of adults making long migrations to areas where they feed, spawn, and spend the winter. Because of this, schools from an individual stock generally travel in a triangular pattern between their spawning grounds, for example in southern Norway, their feeding grounds in Iceland and their nursery grounds in northern Norway. Individuals reach maturity between the ages of three and nine years and can live for up to 15 years. In any one month, one of the many populations scattered across the species vast range will be spawning. Fish in the North Sea spawn between January and April at a depth of less than 70 metres and a sea temperature of 4-7oC. Female herring can produce between 30,000 to 200,000 eggs each. The eggs are sticky and are laid on marine vegetation or directly onto the seabed. Schools of herring can produce so many eggs that they cover the ocean bottom in a dense carpet. By late spring, larvae grow into juveniles that form large schools in coastal waters during the summer. Because of this, near constant spawning throughout the species’ range, the Atlantic herring is a huge fishery resource with a currently estimating annual catch (for this species alone) of over three million tonnes.
Wynken, Blynken, and Nod one night sailed off in a wooden shoe. Sailed on a river of crystal light, into a sea of dew. “Where are you going, and what do you wish?” the old moon asked the three. “We have come to fish for the herring fish that live in this beautiful sea; nets of silver and gold have we!” said Wynken, Blynken, and Nod.
[_Eugene Field (1850-1895) _]
Calappa is a genus of crabs known colloquially as box crabs or, more-commonly, shame-faced crabs. The former name comes from their distinctly bulky carapace and the latter from the way the crab’s chelae fold inwards towards the face, as if hiding it in shame. With good reason. Calappa granulata (Linnaeus, 1758) is a subtropical species restricted to the Mediterranean eastwards to Israel but rarely as far north as the Adriatic Sea. It also occurs in the Eastern Atlantic from northwest Africa to the coast of Portugal, and westwards out to the Açores in the near-mid Atlantic. The species is demersal, with a recorded depth range of 13-400 metres, but more usually occurs from 30-150 metres. The illustrated individual was collected from the fish market at Marsaxlokk, Malta in March 2013 and shows it in dorsal view (A) and from in front (B), to illustrate the chelae.
The carapace of Calappa granulata can reach a width of 11-12 centimetres and is coloured rose-yellow with five rows of dark crimson tubercles. There are four similar rows on each chela. The carapace is squat, rounded and convex with four converging grooves between the rows of spots and flat, rayed, spines around the margin. The claws, or chelae are triangular and heavily spinose dorsally. The eyes and antennae are both small and quite inconspicuous. The four pairs of walking legs are slim and their basal gills are hidden within the carapace.
The shame-faced crab is a lie-in-wait predator and can rest motionless, sometimes for several days, in a temporary burrow in sand. To bury itself, the front surfaces of the chelae are pressed against the sand like planks. The crab then uses these bases to push itself obliquely backwards into the substratum, the upturned spinose rear of the carapace assisting in this backward shovelling. In such buried repose, only the eyes and antennae are visible above the sediment surface. The inner faces of the huge chelae are concave and thereby form a respiratory channel behind them when the animal is buried, thereby preventing sand from clogging the gills. The motionless, buried, camouflaged attitude combined with the dorsal patternation makes for near-perfect invisibility. When prey is detected, Calappa granulata emerges from its burrow and chases it by holding the body upright and running tip-toe on its now straightened legs.
Although Calappa granulata is the only common calappid in the Mediterranean Sea, there are many more species in the Indo-West Pacific and one of the commonest of these is Calappa philargius (Linnaeus, 1758) (Sakai 1976). The adult spotted box crab has a total of six spots: two large dark spots on each chela and a dark ring around each eye. It too lives in sand or broken shell substrata from the intertidal to depths of around 50 metres. Calappa philargius is about the same size as Calappa granulata and an individual was found on Hoi Sing Wan in the New Territories of Hong Kong in 1990. It is known that calappids feed mainly on molluscs and I thus had the opportunity to see this Asian species capturing prey for myself. The individual was kept in an aquarium with a good bed of sand into which it buried quickly. It was left for a few days to acclimate and then offered small gastropods some of which were empty shells with hermit crabs inside. The left chela, which has pointed fingers, is used to hold a potential prey shell. The moveable finger of the bigger right chela has a large tooth fitting between two prominences on the fixed finger. This apparatus is used like a can opener to open up prey shells from the aperture upwards until the retracted inhabitant is reached.
When a prey hermit crab was spotted by my Calappa philargius, it emerged dramatically from its burrow, shook itself like a wet dog and, in this cloud of sediment, grabbed the unsuspecting hermit crab. This animal’s fate was sealed. I could think only of the horror, anthropomorphically, experienced by the hermit crab when attacked in such a nightmarish manner, to retreat deeper and deeper into one’s home but, ultimately, with nowhere else left to go, to be eaten alive.
‘God has an inordinate fondness for beetles’, there being an estimated excess of 30 million species. This quotation is widely attributed to Charles Darwin (1809-1882) although this is incorrect and was spoken by another great British rascal of a philosopher, J.B.S. Haladane (1892-1964), who actually said ‘God has an inordinate fondness for stars and beetles’.
To take just one genus – Dyschirius – it is estimated that there are around an estimated 245 species worldwide although, strangely, it is not recorded from Australasia. They are most diverse in the cooler Palaearctic with around sixty species. Species of Dyschirius are cryptic, subterranean, burrowers and not commonly collected and, thus, species numbers may be much higher. Of the many millions of species of beetles (Coleoptera), – representatives of the Carabidae and Staphylinidae occur on the intertidal zone, the latter being predominant. In northern Europe there may be over 70 such species, most being unstudied. Beetles occupy three rather distinct coastal habitats: the intertidal zone, sandy beaches and salt-mudflats. The species are divided into three major groups, that is halobiontic (restricted to the seashore), haloxenic (tolerant of salty conditions) and halophilous (may be found elsewhere, inland). Most typically coastal beetles are found beneath algal wrack, a few are carnivorous and some phytophagous and most are either very local or rare and but a few are common.
Dyschirius impunctipennis Dawson, 1854 and Dyschirius salinus Schaum, 1843 can be defined as halophilous and halobiontic species, respectively. And so it was unusual to discover two colonies of them living together, sympatrically, on the sandy intertidal bank of a stream draining into Ålbæck Bay, North Jutland, Denmark. Both live in burrows and from which they emerge to feed.[_ Dyschirius salinus ]is[ ]known to[ ]live on such a habitat in Great Britain and Ireland (Luff 2007) while [_Dyschirius impunctipennis _]is similarly known from such habitats in Latvia (Savich 1998). When discovered, species of _Dyschirius are commonly associated with staphylinid beetles of the genus Bledius (also described in this book on page 122-123) and upon which, being predators, they may prey. Bledius spectabilis is an harmless herbivore, eating pieces of algae and other plant matter. Since, however, there are few studies on this relationship it may be merely that they are occupying the same habitats, all of them[_ _]building subterranean burrows in which the females maintain their eggs and larvae.
Species of [_Dyschirius _]are small, cylindrical, beetles with a dark, shiny, and globular pronotum, large eyes and first pair of digging legs. The mouthparts are formidable jaws. The abdomen is similarly shiny and elongate, both species being about six millimetres long and capable of running fast.
In a long-term survey of the estuary of the River Ijzer in Belgium, Desender et al. (2007) identified ninety-six species of coastal beetles. The estuary had, in the past, been subject to extensive damage. Immediately after restoration measures of salt marsh and sand dune habitats, however, there was a marked increase in several target species of high conservation interest, that is, Red-list species. Historical beetle data showed that many species that had disappeared from the area during the past century had not yet been able to re-colonize it, however. This was especially true for salt marsh species. Similarly, many sand dune species re-appeared but did not establish viable populations.
What struck me about the research of Desender et al. (2007), however, was not the details of the effects that habitat restoration had had upon beetle re-colonisation of the estuary, but the diversity of such species. Most studies of intertidal ecology, including those of my own, rarely mention beetles. Yet, it is clear that these are poor representations, in the light of such beetle studies, of biodiversity and of the importance of such species in habitat ecology.
Every Easter, the entire intake of first year biology students and ecology staff members and postgraduates of the departments of Zoology and Botany of the University of Hong Kong decamped to Wu Kwai Sha on the shores of Tolo Harbour in the New Territories of Hong Kong for a week of field studies. A former orphanage, the, now, YMCA camp had accommodation in little houses, a now unused classroom, a lecture theatre and it had a pier. Against this a Tanka fisherman, his wife and boat would sometimes moor each night. In the early 1970’s, the New Territories accommodated a number of ethnic minorities with Hakka farming villages in the valleys, and Hoklo and Tanka, almost gypsy-like, fisher-folk.
One day, in the mid 1970’s, having arrived early at Wu Kwai Sha to prepare for the field camp, I asked the Tanka fisherman if one of my students and I could go out fishing with him. He eventually agreed and we had a great day trawling up and down Tolo Channel for prawns. The front of the vessel also had a powerful, downward pointing, light, charged by a generator itself working off the engine, and I asked what it was for. The fisherman explained that it was for catching squid and promised to show us how. That evening the boat’s light was fired up and hand-lines baited with sprat caught earlier were thrown overboard. We were shown how to wait for a tug on the line and to slowly wind it in towards the light. This was not to attract fish but so that one could see, in its glare, the feeding squid. When close enough, a dip net was plunged into the water behind the squid (because their escape response is in that direction). As they were lifted unceremoniously from the sea, the squids ejected black ink everywhere and changed colour in a dazzling show of melanophoric display of distress and flashed two bright, luminescent green lights inside their bodies.
The squid we caught that night was, I discovered later, [Photololigo duvaucelii _](d’Orbigny, 1835). The, so-called, Indian squid reaches a body length of, generally, fifteen centimetres plus, of course, the head and ten tentacles. They may, however, reach a body length of 30 centimetres. This tropical squid is widely distributed throughout the Indo-West Pacific and in Indian Ocean waters it is fished intensively using trawl nets (Kasim 1985,[ ]Vidyasagar and Deshmukh 1992) and has been intensively studied there.[ _]I could identify the species using a Hong Kong book written by two local fisheries officers Voss and Williamson (1971), both of whom had moved on by the time I arrived there. In these authors’ description of [_Photololigo duvaucelii, _]they explain that each squid has a pair of oval luminous organs situated on each side of its rectum – hence the generic name.
With the passing years, snorkelling in the north-eastern waters of Hong Kong, I would see this squid again from time to time. But this time alive and swimming in small shoals. Here, in the shallow waters of bays and inlets in Hong Kong, which is why I prefer to call it the bay squid, Photololigo duvaucelii breeds. The long left ventral arm (with which it catches its prey) of the male is modified for the transfer of sperm packets from its mantle into that of a female it has embraced face-to-face. The sperm capsules are inserted into the reproductive orifice of each successfully courted female where they fertilise her eggs. She, subsequently, lays egg masses consisting of many egg capsules each of which, in turn, contains between 125-150 eggs. The fertilised eggs are two millimetres in diameter and very yolky. The young squid hatch out after only five days [_(_]Asokan and Kakati 1991) and begin their free life in the shallow bays of their parent’s home. They have to be careful here though because the same parent’s are cannibals as well as predators of fish and crustaceans.
The eel goby, [Trypauchen vagina _](Bloch & Schneider, 1801), is the type species of its genus, which includes only one other, recently identified, taxon – _Trypauchen pelaeos (Murdy, 2006) (Murdy 2006, 2011). Trypauchen vagina occurs throughout the Indo-Pacific Region, including the Indian Ocean, the Persian Gulf, Arabian Sea and the western Pacific Ocean, from Kuwait to New Caledonia. It has also been reported from as far out into the Western Pacific as New Caledonia. The species occurs throughout the Malaysian Archipelago (Cantor 1850), the Philippines and northwards into China. The goby mainly occurs in the shallow marine and brackish waters of estuarine and coastal areas and lives in a self-dug burrow, which it stays close to, in the silty-muddy sea bed. And this is where I first found it, albeit quite by accident.
Soon after arriving in Hong Kong, in co-operation with staff of the Department of Civil Engineering of the University of Hong Kong, I built an experimental raft, called Porpita named after the floating anthomedusan ‘blue-button’, Porpita porpita (Linnaeus, 1758), to test antifouling paints on submerged experimental panels. Situated in Tai Tam Bay on the southern coast of Hong Kong Island, the raft housed the racks of test panels and was reached by a small rowing boat named Hypsypops – a fish. One day, a small, newly-constructed, dredge was thrown into the sea from Hypsypops and I and the technician, Mr Lam Man Hung, in charge of the raft, attempted to haul it back onto this structure. No easy task as it now weighed a seeming tonne! But, with the help of passing local sea scouts, this was achieved and its contents spewed into buckets. And in it was a specimen of Trypauchen vagina. Never having seen such a weird creature, it was taken to the laboratory for closer inspection.
The fish is, however, commonly found for sale in seafood markets in Hong Kong and is used to make a milky soup reputedly good for new mothers to stimulate the production of blood and during breast feeding as the fish is also believed to promote milk production.
Trypauchen vagina has an elongated body up to 22 centimetres in length covered in cycloid scales. It is reddish-purple in colour and on the upper edges of the left and right gill covers are distinctive oval holes that open into pouch-like cavities. These are present in only a few of the genera of the Amblyopinae but their function is unknown. It has reduced eyes situated in an orbital depression and are entirely covered with skin and the anterior portion of its head is protected by thick wrinkly flesh. Both adaptations aid it in digging its burrows. But it also gives the fish a countenence only a mother could love. The anal (with between 39-50 rays), two, united, dorsal fins (with between 46-58 rays), the first with six flexible spines, and the caudal fin (with 17 rays) are fused into a membranous structure forming a continuous margin around the posterior end of the body. The pelvic fins are also fused together to form a cup-shaped suction disk. The pectoral fins (15-20 rays) are small with the upper rays longer than the lower ones. The mouth slants obliquely upwards and has two rows of sharp canine-like teeth lining both jaws. The outer row of the upper jaw has 4-16 teeth, the outer row of the lower jaw 8-13. The teeth on the inner rows are smaller than those on the outer and the species is said to be omnivorous, but mostly preying on small crustaceans that wander near its burrow entrance. Emerging from its burrow, it must frighten them to death.
Overall,[_ Trypauchen vagina_] is a distinctive blotchy purple-red colour. The cheeks, the areas around the eyes and the area behind the gills and above the pectoral fins are bright red. The fins are a translucent rose-pink. Easily identifiable, Trypauchen vagina has recently been introduced into the Mediterranean Sea being first recorded from off the coast of Israel in 2009 (Salameh et al. 2010) and subsequently in 2010 from the Levant Basin off the coast of Turkey (Akamca et al. 2011). The species is well known from the Arabian Sea and is thus thought to be a Lessepsian immigrant having reached the eastern Mediterranean probably through the Suez Canal and bringing the total of such introduced fish species to seventy-four.
The blue-rayed limpet Patella pellucida Linnaeus, 1758[* ]used to be ascribed to its own genus, Patina.[ *]Koufopanou et al. (1999) have shown using molecular evidence, however, that it is closely related to the more well known limpets, Patella, of rocky shores. The separate identity was initially proposed because Patella pellucida exists as two forms, which used to be referred to as sub-species – the smaller blue-rayed Patella pellucida pellucida and the larger yellow-brown Patella pellucida leavis. The two forms have only relatively recently been shown to represent distinct phases in the life history of the same species, as first described by Graham and Fretter (1947).
Patella pellucida[* ]is a cold-water species that occurs along the coast of the northeastern Atlantic from Iceland and Norway, south to Portugal. It shuns the silty-sand shores of Belgium and the Netherlands but does occur all around the rocky, exposed, shores of Great Britain and Ireland. The shell of the pellucida form (A) grows to a length of about 12 millimetres and is a thin, translucent, smoothly depressed, cone, with its apex closest to the anterior margin. It is, however, un-mistakably characterised by the distinctive patterning of beautiful, iridescent, blue rays that give it its name. The rays do not develop until the juvenile shell reaches over one millimetre in length and so the latter is exceedingly difficult to discover. Only the pellucida form is active, moving around on its kelp host to feed. The mantle margin beneath and around the shell is fringed with creamy filaments while the two head tentacles are slender with tiny eyespots situated at their bases[.*]
The taller, up to 20 millimetres in length, and more robust shell of the [Patella pellucida laevis _]form (B) and © progressively loses the iridescent blue rays of the smaller _pellucida and is more uniformly cream-brown. It also has prominent concentric growth lines, giving it a more noticeably ledged profile indicating the size at which this individual experiences the species’ change of life. The laevis form also becomes topped by a distinctively eroded apical region, which identifies when this change of life occurs. The laevis form is a highly secretive limpet, living cryptically within the holdfasts of large kelp algae, which it clamps itself firmly into.
To understand the blue-rayed limpet’s life cycle and the two forms that make it up, we need knowledge of the species’ pattern of reproduction. Mature individuals release eggs and sperm into the sea in early spring where the latter fertilise the former. The result is a free-living, planktonic, larval stage which settles in late spring within the carpet of encrusting coralline algae that characterise the lower levels of wave exposed shores and offshore reefs and platforms around the coast of Great Britain. Here, the larvae metamorphose into juveniles that have a high growth rate during their first 4–5 months and a shell length of about 3 millimetres is reached by August (Vahl 1971). As they continue to grow, the juveniles move onto the fronds of tall macrophytic algae. These [pellucida _]juveniles prefer to live, primarily, on the blade-like fronds of the shallow subtidal kelp seaweed _Laminaria saccharina (Linnaeus) Lamouroux (illustrated) but also [Laminaria hyperborea _](Gunnerus) Foslie (Vahl 1972). The juveniles feed on the tissues of its host kelp using the radula inside its mouth, leaving small depressions scraped into the frond. Here, the _pellucida limpet reaches a shell length of 12 millimetres by the following early January and starts to mature sexually and to, later, reproduce and provide the next spring’s cohort of larvae. At this time, however, the limpet undergoes a change of lifestyle and all the pellucida forms begin to migrate down the stipe of the Laminaria kelp towards its holdfast base.
As the limpets grow and move down the stipe of the seaweed towards the holdfast, they manouvre themselves into it where they establish themselves by excavating a depression deep inside this mass of tangled rootlets – themselves gripping some solid object such as a stone. The excavations made by the limpet within the holdfast can weaken its attachment and Kain and Svendsen (1969) suggested that the limpet probably causes considerable loss of plants due to weakening of the holdfast resulting eventually in the seaweed being dislodged by storms. As a consequence, the seaweed is often washed ashore, typically in winter, with the limpets still in place, although juveniles will have been washed off, departed the kelp’s blades or, sometimes, if fortuitously close, hidden themselves within the holdfasts of their host plants. And, hence, usually, only the laevis stage can be found still clamped tightly into its depression within the holdfast.
I was born at a time, grew up in a home and went to a school wherein and whereat there was no room for art in any form. Few pictures adorned our living room walls and no teachers passed on their talents and artistic endeavours because they too, seemingly, had none. As my career in biology began, it so transpired, however, there grew a need for me to draw. At first this was compulsory for higher level biology courses (the old A level examinations) and at university also. Beginning a Ph.D. degree, I was also made to draw, but with an enhanced degree of detail enforced by a meticulous and uncompromising supervisor – Dick Purchon. Subsequently, with my first appointment to the University of Hong Kong, there grew a yet greater need for me to draw and, thereby, illustrate a growing compendium of research papers and popular texts. Slowly, these illustrative skills improved (I think and hope) and the process of drawing became a hobby and then a love. Today, I still draw to illustrate research papers, talks and seminars and, now this book. I have, however, discovered that, in the modern world and among my surviving peers, colleagues and former students, I am an anachronism. Biology and the means to illustrate it have moved on and, today, as I identified in the introduction to this book, where there is a need for it (and this is rare indeed), professional artists fill the void.
There will always be a place in taxonomic biology for illustrations because it is they that provide the detail; for example, the number of hairs or setae (or both) on each limb, or the characteristic, even diagnostic, features of important taxonomic elements that distinguish the taxa, which constitute each of many thousands of plant and animal families and their many millions of species. Additionally, it is typically only when you start drawing that you notice the minute detail of a species you would not otherwise have observed. The best example of this I can give of this was provided by the eminent geneticist Hans Grüneberg (1907-1982) who was Professor of Genetics at University College London from 1956 until his retirement in 1974. Soon after that date, he visited Hong Kong with the intention of studying the inter-population genetics of a small snail, Clithon oualaniensis Lesson, 1831, which I was to help him with by taking him to the dozens of field sights where the species occurred. In discussion, he explained to me how the genetic development of the hairless, laboratory, mouse, had revealed that its face had an arrangement of whiskers hitherto unseen amongst the mat of facial hair, neccesitating his re-description of the species. This paper and illustrations of the revised whisker arrangement of one of the most studied animals in the world was illustrated by his departmental artist. An ancillary post long-vanished from modern universities.
Typically, such illustrative skills were taught in the world’s universities, within the departments of Botany and Zoology. And this is where, at the University of London, I too learnt how to draw the important features of the numerous species presented to us in each practical class. Our slowly developing drawing skills and our interpretation of them, were assessed after each such laboratory sojourn. Not just that, but end-of-year practical examinations also included the assignment of species to sometimes dissect or to examine using a microscope and to then identify, illustrate and interpret. In my case, the bachelor’s degree eventually obtained resulted in being accepted for and undertaking research for a doctoral degree. And this too had a heavy emphasis on illustrative and interpretive skills, as noted above.
Since those days of the 1960’s, however, university degrees in the natural sciences have slowly but progressively removed all such studies of plant and animal form and function from their curricula. Today, the personal, supervised, illustration of a species has almost ceased to exist in undergraduate student biology courses. And because of this, biology students are no longer learning to develop observational and illustrative skills at most, at least western, universities. Thus: where will they learn them?
Today’s scientists lacking illustrative skills use photography as a means to inform readers about biological structures. In recent years, photographic techniques, linked to computer software programmes have greatly facilitated this means of illustration and, accordingly, photographs are now used widely in scientific publications. So much so that, to provide one example I am familiar with, the February 2013 copy of The Malacologist, magazine of the Malacological Society of London, did not contain one original drawing. All the illustrations were either photographs or computer-generated images. In general terms, computer-generated pictures are, at present, acceptable for many scientific purposes although they are not, yet, sophisticated enough for detailed taxonomic purposes.
Photomicrographs obtained using scanning electron microscopy (SEMS) are also used widely and are a particularly useful tool in demonstrating the finest of details that no human eye nor compound microscope could elaborate upon. One failing with this technology, however, is that when describing a new species, it must be of the holotype with the accessory data obtained from paratypes and other specimens. Scanning electron microscopy produces images of gold-film plated cadavers and, thus, any such specimen, if it is to be a holotype, requires access by others and not all researchers will have the appropriate machinery to be able to re-examine such material. A similar example relates to computer-aided drawing, which is being used by some biologists to illustrate their research papers. Technically, in terms of graphs, histograms and maps, the manuscripts that house them have now been almost universally accepted and add near-perfection to such illustrations. Where computer-aided drawing is used to illustrate both living and dead specimens, however, quality is not only lost but is often so stylized as to lack any feeling of a sentient being – more like a cartoon. Doubtless, technology will improve upon the present quality.
Nevertheless, the illustrator will be faced with the same problem – namely that the drawing was created by the utilisation of another’s technological skill in the creation of the programme that facilitated it. Such illustrations, moreover, appear mechanical, which is what they are of course, and lack the human touch that individualises and interprets the subjects. And, moreover, lacks the quality of emphasising the more important features.
In the art world, one part of the importance of, for example, a painting is that it tells a story, and that any such visual image is open to interpretation by each individual observer. Such images thereby become personal and each one tells its own story not only to each person but, collectively, to an entire society. Paintings become a part of our collective psyche, their meanings continually mulled over and often hotly debated by professional critics and members of the art-loving public both. I need only give the example of the Mona Lisa by Leonardo da Vinci in the Louvre, Paris, to justify this. A few photographs by a very few photographers have achieved a similar level of public acclaim and although the images can be both dramatic and revealing, we do not accord them the same level of respect nor, especially, affection. Paintings, even sketches, again I give the example of da Vinci, by great artists assume almost mythical, certainly legendary, significance, whereas photographic images are more fleeting.
There are, of course, levels of greatness ranging from the paintings of the world’s masters to our children’s first squiggles, which we proudly stick on our refrigerator doors. But their meanings are the same – we love the individuality and personality expressed in the piece of art produced by the human hand.
It has been said that the modern use of computer search engines to provide us with information either has destroyed or is destroying our individual and collective capacity for memory. We simply do not need to memorise facts. At the very basic level, we do not even need to memorise an appointment – any modern personal computer or tablet can remind us of our day’s planned activities. Although a memory will always be important individually, there can be made a case for the fact that our collective recollections are shrinking. Instead, it is stored in microchips, which we can, mostly, access through our fingertips and without the need to exercise our mental processes.
Similarly, as I alluded to in the introduction to this book, today, at least in western societies, computer games are now a part of most people’s lives. A 2011 of study of Canadians found that almost 60% of people played electronic games regularly. A 2013 survey in the U.S.A. found that 58% of Americans play games and that most (64%) ‘gamers’ were younger than 36. Of these, half were under 18. Also in the U.S.A., the 2012 sales of video games were worth US$15 billion. The figures are proportionally similar for the United Kingdom and show that a majority of British children and adults regularly find amusement in video games rather than creative pastimes. At one level, an addiction to video games in the young is said to have led, or contributed to, reduced physical exercise and, as a consequence, growing levels of childhood obesity. Almost unbelievably, more women filing for divorce are complaining that their husbands spend too much time playing video games. In 2014, of those British wives who cited unreasonable behaviour for ending their marriage, 15% believed that their partners put video gaming before them.
Our addiction, at all ages, to visual stimulation, notably by the ever-switched on television set, has achieved the same problem again in both children and adults. It has been documented, for example, that the average Briton spends more than four hours each day watching television. Alongside, of course, the associated television dinner of microwave-heated, pre-packaged, food. Not only this, but the average Briton spends half their free time, nine hours each day, in front of a screen – television, computer, tablet and, increasingly, a smart-phone. Such visual distractions, often highly dramatic, even violent, and misogynistic, stifle our heretofore creativity and reduce it to conversation pieces about the previous evening’s viewings and game results.
The advent of visual electronic games and pre-digested television programming has, therefore, achieved not just a reduction in memory by minimalising it to how not why, has also achieved a decline in individual creativity by parasitising the programming skills of others. We seem to be losing and, in extreme cases, have lost, our collective creative compass. The electronic media is no longer a force in our lives, originally perceived to be for good, but is shaping our lives – transforming us from an image of self into a metaphor of what others think and believe about us and judge us on that. Is it any wonder that the suicide rate among the young has never been so high, and is increasing.
I sometimes visit craft and art fares in the villages and smaller towns of southern England. Here, I often find that it is the elderly who, post-retirement, re-discover the pleasure of creative art in all its forms – jewellery, silver-smithing, pottery, carving, sculpting and drawing and painting. Of course, with a family of grown-up children, I am no longer invited to school exhibitions of creative work but I get the feeling it is not over-ridingly prevalent. Local museums and libraries sometimes organise exhibitions by local artists of all categories and I attend these. But, one rarely comes across groups of people, either young or old, sitting together painting at a local beauty spot or drawing an old building. We no longer seem to gain collective pleasure from a mutually appreciated sight and skill. Sharing and, in a sense, immortalising a moment in time and place. One does see this, occasionally, in travels overseas but in our, reputedly, more sophisticated western societies, we seem to have lost so much to episodes of instant electronic gratification not of our own creation.
It is such a pity that fewer and fewer people are not simply picking up pencils, pens, charcoal, watercolours or even oils today. It is almost as if it is considered old-fashioned, almost quaint, to have any kind of artistic abilities or creative talents. For me, the discovery, as a marine biologist, of an unique creature, the examination and study of it and, then, the interpretation and illustration of it, is still a wonderment. It takes one away from the artificial world we live in, back, almost, to a time before time when there was a need to express ourselves in our own unique manner and, thereby, achieve not just instant respect but also a tiny amount of immortality. Just as mind relaxing as the act of drawing is, so it is also fulfilling and I make no apologies for this book of my own illustrated journey through life. Why? Because: it is the self-fulfilment that is important.
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Until relatively recently, hand-drawn illustrations were the backbone of the biological sciences – both for teaching and research. The art of biological illustration has its origins deep in our ancient psyche as evidenced by Neolithic cave wall paintings. Every human society has found the means to express its wonderment at the diversity of life around it culminating in the illustration of our first printed books. And, eventually our scientific and more popular writings. Students of biology used to be taught how to draw. Today - rarely so. In this volume, the author explains how he was taught to draw biological specimens as a university student and which, then, continued with the need to illustrate his own books and research articles. This volume is a compendium of 80 of the author’s drawings, some later ones in colour, nearly all published herein for the first time. It plots the author’s temporal change in his interpretation of marine plant and animal form – permitting an analysis of function and thus a greater appreciation of just how wonderful life is. The author concludes with a plea for the continued teaching of drawing, illustration and art in our schools, colleges and universities if only to broaden our children’s appreciation of life itself. And, because, it is the epitome of the expression of our unique ‘self’ it is, thus, a fulfilment of one’s own being.