Walter Garstang
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Walter Garstang FLS FZS (9 February 1868 – 23 February 1949), a Fellow of
Lincoln College, Oxford Lincoln College (formally, The College of the Blessed Mary and All Saints, Lincoln) is one of the constituent colleges of the University of Oxford, situated on Turl Street in central Oxford. Lincoln was founded in 1427 by Richard Fleming, the ...
and
Professor Professor (commonly abbreviated as Prof.) is an Academy, academic rank at university, universities and other post-secondary education and research institutions in most countries. Literally, ''professor'' derives from Latin as a "person who pr ...
of
Zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal kingdom, including the anatomy, structure, embryology, evolution, Biological clas ...
at the
University of Leeds , mottoeng = And knowledge will be increased , established = 1831 – Leeds School of Medicine1874 – Yorkshire College of Science1884 - Yorkshire College1887 – affiliated to the federal Victoria University1904 – University of Leeds , ...
, was one of the first to study the functional biology of marine
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e. His best known works on marine larvae were his poems published as ''Larval Forms and Other Zoological Verses'', especially ''The Ballad of the Veliger''. They describe the form and function of several marine larvae as well as illustrating some controversies in evolutionary biology of the time. Garstang was known for his vehement opposition to Ernst Haeckel's
Biogenetic Law The theory of recapitulation, also called the biogenetic law or embryological parallelism—often expressed using Ernst Haeckel's phrase "ontogeny recapitulates phylogeny"—is a historical hypothesis that the development of the embryo of an a ...
, now discredited. He is also noted for his hypothesis on
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
evolution, known as Garstang's theory, which suggests an alternative route for chordate evolution from
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s.


Early life

Walter Garstang was born on 9 February 1868 as the eldest son of Dr Walter Garstang of Blackburn and his wife Matilda Mary Wardley, and older brother of the archaeologist
John Garstang John Garstang (5 May 1876 – 12 September 1956) was a British archaeologist of the Ancient Near East, especially Egypt, Sudan, Anatolia and the southern Levant. He was the younger brother of Professor Walter Garstang, FRS, a marine biol ...
. In 1895 he married Lucy Ackroyd; they had one son, Walter Lucian Garstang,. and five daughters.


Academic career

In 1884 at the age of 16, he was awarded a scholarship to
Jesus College, Oxford Jesus College (in full: Jesus College in the University of Oxford of Queen Elizabeth's Foundation) is one of the constituent colleges of the University of Oxford in England. It is in the centre of the city, on a site between Turl Street, Ship St ...
and was initially going to study medicine. Under the guidance of
Henry Nottidge Moseley Henry Nottidge Moseley Fellow of the Royal Society, FRS (14 November 1844 – 10 November 1891) was a British natural history, naturalist who sailed on the global scientific expedition of Challenger expedition, HMS ''Challenger'' in 1872 through ...
, he joined the school of
Zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal kingdom, including the anatomy, structure, embryology, evolution, Biological clas ...
and graduated in 1888 at the age of 20. Before graduation, Garstang was offered a position as secretary and assistant to Gilbert C Bourne, the new resident director of the
Marine Biological Association of the United Kingdom The Marine Biological Association of the United Kingdom (MBA) is a learned society with a scientific laboratory that undertakes research in marine biology. The organisation was founded in 1884 and has been based in Plymouth since the Citadel Hi ...
in
Plymouth Plymouth () is a port city and unitary authority in South West England. It is located on the south coast of Devon, approximately south-west of Exeter and south-west of London. It is bordered by Cornwall to the west and south-west. Plymouth ...
. There he met
Ray Lankester Sir Edwin Ray Lankester (15 May 1847 – 13 August 1929) was a British zoologist.New International Encyclopaedia. An invertebrate zoologist and evolutionary biologist, he held chairs at University College London and Oxford University. He was th ...
. In 1891 he left Plymouth and was a Berkley Research Fellow under Milnes Marshall at
Owens College Owens may refer to: Places in the United States *Owens Station, Delaware *Owens Township, St. Louis County, Minnesota *Owens, Missouri *Owens, Ohio *Owens, Virginia People * Owens (surname), including a list of people with the name * Owens Bro ...
,
Manchester Manchester () is a city in Greater Manchester, England. It had a population of 552,000 in 2021. It is bordered by the Cheshire Plain to the south, the Pennines to the north and east, and the neighbouring city of Salford to the west. The t ...
. A year later, Garstang returned to Plymouth as Assistant Naturalist, only to be elected a Fellow of
Lincoln College, Oxford Lincoln College (formally, The College of the Blessed Mary and All Saints, Lincoln) is one of the constituent colleges of the University of Oxford, situated on Turl Street in central Oxford. Lincoln was founded in 1427 by Richard Fleming, the ...
, in 1893. In 1894, while
Ray Lankester Sir Edwin Ray Lankester (15 May 1847 – 13 August 1929) was a British zoologist.New International Encyclopaedia. An invertebrate zoologist and evolutionary biologist, he held chairs at University College London and Oxford University. He was th ...
held the Linacre Chair, he became a lecturer at Lincoln College and in 1895 he started the series of Easter classes in which he took students on week-long field courses to Plymouth. Between 1902 and 1907 Garstang was employed by the MBA as the principal investigator working on North Sea fisheries. He helped to establish a fisheries laboratory in
Lowestoft Lowestoft ( ) is a coastal town and civil parish in the East Suffolk district of Suffolk, England.OS Explorer Map OL40: The Broads: (1:25 000) : . As the most easterly UK settlement, it is north-east of London, north-east of Ipswich and sou ...
that was later to become the
Centre for Environment, Fisheries and Aquaculture Science The Centre for Environment, Fisheries and Aquaculture Science (Cefas) is an executive agency of the United Kingdom government Department for Environment, Food and Rural Affairs (Defra). It carries out a wide range of research, advisory, consulta ...
(Cefas), part of the
Ministry of Agriculture, Fisheries and Food (United Kingdom) The Ministry of Agriculture, Fisheries and Food (MAFF) was a United Kingdom government department created by the Board of Agriculture Act 1889 (52 & 53 Vict. c.30) and at that time called the Board of Agriculture, and then from 1903 the Board ...
. Garstang instigated a series of detailed fisheries surveys throughout the southern North Sea aboard the ''
RV Huxley RV ''Huxley ''was the first research vessel used by the Marine Biological Association of the United Kingdom explicitly for fisheries research and is regarded as the first vessel yielding data for the Ministry of Agriculture, Fisheries and Food (U ...
'', under the auspices of the newly formed
International Council for the Exploration of the Sea The International Council for the Exploration of the Sea (ICES; french: Conseil International de l'Exploration de la Mer, ''CIEM'') is a regional fishery advisory body and the world's oldest intergovernmental science organization. ICES is headqu ...
(ICES). Garstang was Professor of Zoology at the
University of Leeds , mottoeng = And knowledge will be increased , established = 1831 – Leeds School of Medicine1874 – Yorkshire College of Science1884 - Yorkshire College1887 – affiliated to the federal Victoria University1904 – University of Leeds , ...
from 1907 to 1933. The Garstang Building at the university is named in his honour. In 1912 in cooperation with Professor Alfred Denny of the
University of Sheffield , mottoeng = To discover the causes of things , established = – University of SheffieldPredecessor institutions: – Sheffield Medical School – Firth College – Sheffield Technical School – University College of Sheffield , type = Pu ...
he established the Robin Hood's Bay Marine Laboratory. The minutes of Sheffield's Faculty of Pure Science on 12 March 1912 record the following resolution that was carried unanimously: That the Faculty approves of the proposal to extend the work of the Department of Zoology by co-operating with the University of Leeds in establishing a small marine Zoological Laboratory at Robin Hood's Bay.


Garstang's hypothesis

Garstang made the radical suggestion that the chordates evolved from the larvae of another group, whether the larvae of
hemichordates Hemichordata is a phylum which consists of triploblastic, enterocoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and includ ...
or of
echinoderms An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea li ...
, by
progenesis Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compare ...
(which Garstang called "neoteny"). Echinoderm larvae, like chordates, are bilaterally symmetric. Especially notable are their similarities to larvae of hemichordates, which are a step closer to chordates as they share two of the five most commonly noted chordate characteristics, namely a hollow
neural tube In the developing chordate (including vertebrates), the neural tube is the embryonic precursor to the central nervous system, which is made up of the brain and spinal cord. The neural groove gradually deepens as the neural fold become elevated, a ...
and
pharyngeal slit Pharyngeal slits are filter-feeding organs found among deuterostomes. Pharyngeal slits are repeated openings that appear along the pharynx caudal to the mouth. With this position, they allow for the movement of water in the mouth and out the phary ...
s. Garstang's idea has been expanded and is supported by many lines of evidence. Perhaps most interesting and compelling is the fact that some
amphibians Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbore ...
can stay in larval form and still reach sexual maturity—this shows that echinoderm larvae could, theoretically, have become sexually mature and simply stopped morphing into adults, instead evolving into chordate ancestors. Species that show this refusal to leave the larval stage include mud puppies and other
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
s, which either partially or completely show
neoteny Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
(also called pedomorphism): retention of juvenile traits or
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological proper ...
s after sexual maturity. Garstang's hypothesis (this term is also used for his proposals on gastropod torsion, as described in ''The Ballad of the Veliger'') was revolutionary for both its time and idea: it suggests that not only may single species evolve, but that single life stages of species may evolve into separate organisms. The hypothesis, which Garstang proposed in the early 20th century, seemed far-fetched at the time of its conception and did not receive support until after Garstang's death.


Poetry


''Larval Forms and Other Zoological Verses''

First published in 1951, two years after his death, ''Larval Forms and Other Zoological Verses'' () is a compilation of 26 poems by Garstang on the form, function and development of various larval invertebrates. Although they were published posthumously, Garstang had had a desire to publish them for many years and never did because he always thought he would add to them. Except for the introduction written by
Sir Alister Hardy Sir Alister Clavering Hardy (10 February 1896 – 22 May 1985) was an English marine biologist, an expert on marine ecosystems spanning organisms from zooplankton to whales. He had the artistic skill to illustrate his books with his own drawing ...
, everything in the final publication, including the title and order of the poems, was his own work. Many of his poems were written to express his views on the scientific theories of the time. Most notable may be ''The Axolotl and the Ammocoete'' which speculates an evolutionary relationship between the Axolotl and the Ammocoete.
Alister Hardy Sir Alister Clavering Hardy (10 February 1896 – 22 May 1985) was an English marine biologist, an expert on marine ecosystems spanning organisms from zooplankton to whales. He had the artistic skill to illustrate his books with his own drawings ...
wrote on this in the Introduction to ''Larval Forms'':
Only a few months before he died Garstang had drafted a communication to ''Nature'' to put forward his latest suggestion that ''
Amphioxus The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...
'' might be regarded as a paedomorphic ammocoete-like larva of a Cyclostome; it was never sent, because the day on which he was to have posted it he found that the whole of his idea had recently and quite independently been published by the great Stensio.
Most of these poems were written before 1922 and reflect the knowledge and theories of that time. In some of the poems, some of the animals, or species of animals, have speaking parts. The poems included in his final work are:


''The Ballad of the Veliger''

Walter Garstang's most famous zoological verse, ''The Ballad of the Veliger'', was first published in 1928 and privately printed. Copies were handed out at that year's BA meeting, where he gave the Presidential Address to the Zoology section. ::''The Ballad of the Veliger'' or
'' How the gastropod got its twist''
The
Veliger A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells. Description The veliger is the characteristic larva of the gastropod, bivalve and scaphopod ...
's a lively
tar Tar is a dark brown or black viscous liquid of hydrocarbons and free carbon, obtained from a wide variety of organic materials through destructive distillation. Tar can be produced from coal, wood, petroleum, or peat. "a dark brown or black bit ...
, the liveliest afloat,
A whirling wheel on either side propels his little boat;
But when the danger signal warns his bustling submarine,
He stops the engine, shuts the port, and drops below unseen.
He's witnessed several changes in
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
motor-craft;
The first he sailed was just a tub, with a tiny cabin aft.
An Archi-mollusk fashioned it, according to his kind,
He'd always stowed his
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
and things in a mantle-sac behind.
Young Archi-mollusks went to sea with nothing but a velum—
A sort of autocycling hoop, instead of pram—to wheel 'em;
And, spinning round, they one by one acquired parental features,
A
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses ** Thin-shell structure Science Biology * Seashell, a hard o ...
above, a foot below—the queerest little creatures.
But when by chance they brushed against their neighbours in the briny,
Coelenterate Radiata or Radiates is a historical taxonomic rank that was used to classify animals with radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of animals that do not form a monophyletic ...
s with stinging threads and
Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s so spiny,
By one weak spot betrayed, alas, they fell an easy prey—
Their soft preoral lobes in front could not be tucked away!
Their
feet The foot ( : feet) is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is a separate organ at the terminal part of the leg made ...
, you see,
amidships This glossary of nautical terms is an alphabetical listing of terms and expressions connected with ships, shipping, seamanship and navigation on water (mostly though not necessarily on the sea). Some remain current, while many date from the 17th t ...
, next the cuddy-hole
abaft This glossary of nautical terms is an alphabetical listing of terms and expressions connected with ships, shipping, seamanship and navigation on water (mostly though not necessarily on the sea). Some remain current, while many date from the 17th t ...
,
Drew in at once, and left their heads exposed to every shaft.
So Archi-mollusks dwindled, and the race was sinking fast,
When by the merest accident salvation came at last.
A fleet of fry turned out one day, eventful in the sequel:
Whose left and right retractors on the two sides were unequal:
Their
starboard Port and starboard are nautical terms for watercraft and aircraft, referring respectively to the left and right sides of the vessel, when aboard and facing the bow (front). Vessels with bilateral symmetry have left and right halves which are ...
halliards fixed
astern This list of ship directions provides succinct definitions for terms applying to spatial orientation in a marine environment or location on a vessel, such as ''fore'', ''aft'', ''astern'', ''aboard'', or ''topside''. Terms * Abaft (preposition ...
alone supplied the head,
While those set aport were spread
abeam This glossary of nautical terms is an alphabetical listing of terms and expressions connected with ships, shipping, seamanship and navigation on water (mostly though not necessarily on the sea). Some remain current, while many date from the 17th t ...
and served the back instead.
Predaceous Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
foes, still drifting by in numbers unabated,
Were baffled now by tactics which their dining plans frustrated.
Their prey upon alarm collapsed, but promptly turned about,
With the tender morsel safe within and the horny foot without!
This manoeuvre (''fide'' Lamark) speeded up with repetition,
Until the parts affected gained a rhythmical condition,
And
torsion Torsion may refer to: Science * Torsion (mechanics), the twisting of an object due to an applied torque * Torsion of spacetime, the field used in Einstein–Cartan theory and ** Alternatives to general relativity * Torsion angle, in chemistry Bi ...
, needing now no more a stimulating stab,
Will take its predetermined course in a watchglass in the lab.
In this way, then, the Veliger, triumphantly askew,
Acquired his cabin for'ard, holding all his sailing crew—
A
Trochophore A trochophore (; also spelled trocophore) is a type of free-swimming planktonic marine larva with several bands of cilia. By moving their cilia rapidly, they make a water eddy, to control their movement, and to bring their food closer, to captur ...
in armour cased, with a foot to work the hatch,
And double
screws A screw and a bolt (see '' Differentiation between bolt and screw'' below) are similar types of fastener typically made of metal and characterized by a helical ridge, called a ''male thread'' (external thread). Screws and bolts are used to fa ...
to drive ahead with smartness and despatch.
But when the first new Veligers came home again to shore,
And settled down as
Gastropods The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
with mantle-sac afore,
The Archi-mollusk sought a cleft his shame and grief to hide,
Crunched horribly his horny teeth, gave up the ghost, and died.


Archives

The National Marine Biological Library at the
Marine Biological Association The Marine Biological Association of the United Kingdom (MBA) is a learned society with a scientific laboratory that undertakes research in marine biology. The organisation was founded in 1884 and has been based in Plymouth since the Citadel H ...
in Plymouth holds some of Garstang's archival material (diaries and photographs) and documents relating to the Easter Classes.The Archives of the Marine Biological Association of the UK: http://www.mba.ac.uk/NMBL/archives/archives_new.htm Leeds University Special Collections
also holds some archival material.


References


External links

*


Abalone: Larval defenses

Archival material at Leeds University Library
{{DEFAULTSORT:Garstang, Walter 1868 births 1949 deaths Evolutionary biologists English marine biologists English zoologists Alumni of Jesus College, Oxford Fellows of Lincoln College, Oxford Academics of the University of Leeds Fellows of the Linnean Society of London Fellows of the Zoological Society of London