Walter Garstang
Encyclopedia
Walter Garstang a Fellow of Lincoln College, Oxford
, was a marine biologist and zoologist who was one of the first to study the functional biology of marine invertebrate
larva
e. His best known works on marine larvae were his poems which were published together after his death as Larval Forms and Other Zoological Verses, which describe the form and function of several marine larvae as well as illustrate some of the controversies of evolutionary biology of the time. Garstang was known for his vehement opposition to Ernst Haeckel's
Biogenetic Law, now discredited. Garstang is also noted for his hypothesis on chordate evolution, known as Garstang's theory, which suggests an alternative route for chordate evolution from echinoderms. Holland, in Walter Garstang: a Retrospective (Theory in Biosciences 130, in press, 2011) summarizes Garstang’s more important publications with special attention to his evolutionary thoughts—their source, their relation to the biology of his times, and their fate in the twenty-first century.
. In 1884 at the age of 16, he was awarded a scholarship to Jesus College
at the University of Oxford
and was initially going to study medicine. Under the guidance of Henry Nottidge Moseley
, he joined the school of Zoology, 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
in Plymouth
. There he met Ray Lankester
. In 1891 he left Plymouth and was a Berkley
Research Fellow under Milnes Marshall at Owens College. A year later, Garstang returned to Plymouth as Assistant Naturalist
, only to be elected a Fellow of Lincoln College, Oxford
, in 1893. In 1894, While Ray Lankester
was the Linacre Chair, he became a professor 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. The concept of Easter classes was copied at many other schools, and is now considered one of the key courses in any zoologist's training.
There are, of course, many important differences between chordates and echinoderms which must be noted and not ignored. Most importantly, adult echinoderms show little likeness to chordates: echinoderms are radially symmetric, possess calcium carbonate plates in their skin and have tube feet. Garstang made the radical suggestion that perhaps the echinoderm larvae, not adults, had given rise to chordates.
Echinoderm larvae, like chordates, are bilaterally symmetric, and show other shared characteristics with chordates. Especially notable are their similarities to larvae of hemichordates, not true chordates, but rather a step closer to chordates (hemichordates share two of the five most commonly noted chordate characteristics--a hollow dorsal nerve tube and pharyngeal slits).
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 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 and instead could have evolved into chordate ancestors. Species that show this refusal to leave the larval stage include mud puppies and other salamanders, which either partially or completely show pedomorphism (retention of juvenile traits or phenotypes after sexual maturity).
Garstang's Theory is often overlooked, but 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 theory, which Garstang proposed in the early 20th century, seemed far-fetched at the time of its conception and did not receive support until further research had been done after Garstang's death.
, everything in the final publication, including the title and order of the poems, was his own work. The poems included in his final work are:
Of these poems, The Ballad of the Veliger may be his best known 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
wrote on this in the Introduction to Larval Forms:
The great Stensio referred to here was Erik Stensiö
, a Swedish paleozoologist.
Lincoln College, Oxford
Lincoln College is one of the constituent colleges of the University of Oxford in the United Kingdom. It is situated on Turl Street in central Oxford, backing onto Brasenose College and adjacent to Exeter College...
, was a marine biologist and zoologist who was one of the first to study the functional biology of marine invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
larva
Larva
A larva 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...
e. His best known works on marine larvae were his poems which were published together after his death as Larval Forms and Other Zoological Verses, which describe the form and function of several marine larvae as well as illustrate some of the controversies of evolutionary biology of the time. Garstang was known for his vehement opposition to Ernst Haeckel's
Ernst Haeckel
The "European War" became known as "The Great War", and it was not until 1920, in the book "The First World War 1914-1918" by Charles à Court Repington, that the term "First World War" was used as the official name for the conflict.-Research:...
Biogenetic Law, now discredited. Garstang is also noted for his hypothesis on chordate evolution, known as Garstang's theory, which suggests an alternative route for chordate evolution from echinoderms. Holland, in Walter Garstang: a Retrospective (Theory in Biosciences 130, in press, 2011) summarizes Garstang’s more important publications with special attention to his evolutionary thoughts—their source, their relation to the biology of his times, and their fate in the twenty-first century.
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 GarstangJohn Garstang
John Garstang was a British archaeologist of the ancient Near East, especially Anatolia and the southern Levant....
. In 1884 at the age of 16, he was awarded a scholarship to Jesus College
Jesus College, Oxford
Jesus College is one of the colleges of the University of Oxford in England. It is in the centre of the city, on a site between Turl Street, Ship Street, Cornmarket Street and Market Street...
at the University of Oxford
University of Oxford
The University of Oxford is a university located in Oxford, United Kingdom. It is the second-oldest surviving university in the world and the oldest in the English-speaking world. Although its exact date of foundation is unclear, there is evidence of teaching as far back as 1096...
and was initially going to study medicine. Under the guidance of Henry Nottidge Moseley
Henry Nottidge Moseley
Henry Nottidge Moseley, FRS was a British naturalist who sailed on the global scientific expedition of the HMS Challenger in 1872 through 1876....
, he joined the school of Zoology, 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
Marine Biological Association of the United Kingdom
The Marine Biological Association of the United Kingdom 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 laboratory was opened in June 1888...
in Plymouth
Plymouth
Plymouth is a city and unitary authority area on the coast of Devon, England, about south-west of London. It is built between the mouths of the rivers Plym to the east and Tamar to the west, where they join Plymouth Sound...
. There he met Ray Lankester
Ray Lankester
Sir E. Ray Lankester KCB, FRS was a British zoologist, born in London.An invertebrate zoologist and evolutionary biologist, he held chairs at University College London and Oxford University. He was the third Director of the Natural History Museum, and was awarded the Copley Medal of the Royal...
. In 1891 he left Plymouth and was a Berkley
Berkley
- Places :United Kingdom* Berkley, SomersetUnited States* Berkley, Colorado* Berkley, Iowa* Berkley, Maryland* Berkley, Massachusetts* Berkley, Michigan* Berkley, Virginia, formerly a town, and now a neighborhood in Norfolk, Virginia...
Research Fellow under Milnes Marshall at Owens College. A year later, Garstang returned to Plymouth as Assistant Naturalist
Naturalist
Naturalist may refer to:* Practitioner of natural history* Conservationist* Advocate of naturalism * Naturalist , autobiography-See also:* The American Naturalist, periodical* Naturalism...
, only to be elected a Fellow of Lincoln College, Oxford
Lincoln College, Oxford
Lincoln College is one of the constituent colleges of the University of Oxford in the United Kingdom. It is situated on Turl Street in central Oxford, backing onto Brasenose College and adjacent to Exeter College...
, in 1893. In 1894, While Ray Lankester
Ray Lankester
Sir E. Ray Lankester KCB, FRS was a British zoologist, born in London.An invertebrate zoologist and evolutionary biologist, he held chairs at University College London and Oxford University. He was the third Director of the Natural History Museum, and was awarded the Copley Medal of the Royal...
was the Linacre Chair, he became a professor 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. The concept of Easter classes was copied at many other schools, and is now considered one of the key courses in any zoologist's training.
Garstang's Theory
Walter Garstang noted after extensively studying marine animals that both echinoderms and chordates are deuterostomes (in which the blastopore forms the anus), while most other possible ancestors of Chordates are protostomes (the blastopore forms the mouth). This inspired Garstang to suggest an alternate route of evolution: from echinoderms to chordates.There are, of course, many important differences between chordates and echinoderms which must be noted and not ignored. Most importantly, adult echinoderms show little likeness to chordates: echinoderms are radially symmetric, possess calcium carbonate plates in their skin and have tube feet. Garstang made the radical suggestion that perhaps the echinoderm larvae, not adults, had given rise to chordates.
Echinoderm larvae, like chordates, are bilaterally symmetric, and show other shared characteristics with chordates. Especially notable are their similarities to larvae of hemichordates, not true chordates, but rather a step closer to chordates (hemichordates share two of the five most commonly noted chordate characteristics--a hollow dorsal nerve tube and pharyngeal slits).
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 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 and instead could have evolved into chordate ancestors. Species that show this refusal to leave the larval stage include mud puppies and other salamanders, which either partially or completely show pedomorphism (retention of juvenile traits or phenotypes after sexual maturity).
Garstang's Theory is often overlooked, but 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 theory, which Garstang proposed in the early 20th century, seemed far-fetched at the time of its conception and did not receive support until further research had been done after Garstang's death.
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 Garstang's poems 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 HardyAlister Hardy
Sir Alister Clavering Hardy, FRS was an English marine biologist, expert on zooplankton and marine ecosystems...
, everything in the final publication, including the title and order of the poems, was his own work. The poems included in his final work are:
- The Amphiblastula and the Origin of Sponges
- The Invaginate Gastrula and the Planula
- The Origin of Cnidoblasts and Cnidozoa
- Conaria and Co.
- Mülleria and the Ctenophore
- The Onchosphere
- The Trochophores
- Mitraria's Fan Dance
- The Ballad fo the Veliger, or How the Gastropod got its Twist
- Echinospira's Double Shell
- The Nauplius and the Protaspis
- Kentrogon
- Isopod Phylogeny
- The Millipede's Egg-tooth
- The Trilobites and After
- Actinotrocha
- Cyphonautes
- Echinoderm Larvae and the Origin of Quinqueradial Symmetry
- The Pentacrinule
- Tornaria's Water-Works
- Oikopleura, Jelly-builder
- The Ancestry of Vertebrates
- Leptocephalus brevirostris, the Larva of the Eel
- The Axolotl and the Ammocoete
- An Oceanographer's Dream
- To a Herring Gull
Of these poems, The Ballad of the Veliger may be his best known 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
Alister Hardy
Sir Alister Clavering Hardy, FRS was an English marine biologist, expert on zooplankton and marine ecosystems...
wrote on this in the Introduction to Larval Forms:
Only a few months before he died Garstang had drafted a communication toNature to put forward his latest suggestion that Amphioxus 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.
The great Stensio referred to here was Erik Stensiö
Erik Stensiö
Erik Helge Osvald Stensiö was a Swedish paleozoologist.Erik Andersson, as his original name was, was born in the village of Stensjö in Döderhult parish in Kalmar County; he later took his new surname from his place of origin and is occasionally referred to with both names...
, a Swedish paleozoologist.
The Ballad of the Veliger
This, his most famous poem, was first published in 1928, privately printed, and copies were handed out at the BA meeting that year where Garstang gave the Presidential Address to the Zoology section.-
- The Ballad of the Veliger or
How the gastropod got its twist
- The Ballad of the Veliger or
The Veliger's a lively tar, 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 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 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 above, a foot below—the queerest little creatures.
But when by chance they brushed against their neighbours in the briny,
Coelenterates with stinging threads and Arthropods 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, you see, amidships, next the cuddy-hole shaft,
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 halliards fixed astern alone supplied the head,
While those set aport were spread abeam and served the back instead.
Predaceous 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 moral 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, 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 in armour cased. with a foot to work the hatch,
And double screws to drive ahead with smartness and despatch.
But when the first new Veligers came home again to shore,
And settled down as Gastropods 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.