Walter Garstang


Walter Garstang FLS FZS (9 February 1868 – 23 February 1949), a Fellow of Lincoln College, Oxford and Professor of Zoology at the University of Leeds, was one of the first to study the functional biology of marine invertebrate larvae. 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.[1]

Walter Garstang
Walter Garstang.jpg
Born(1868-02-09)9 February 1868
Died23 February 1949(1949-02-23) (aged 81)
Alma materJesus College, Oxford
Known forChordate evolution
Marine invertebrate larvae
Zoology poems
Spouse(s)Lucy Ackroyd
Scientific career
FieldsMarine Zoology
InstitutionsUniversity of Oxford
University of Leeds

Garstang was known for his vehement opposition to Ernst Haeckel's Biogenetic Law, now discredited. He is also noted for his hypothesis on chordate evolution, known as Garstang's theory, which suggests an alternative route for chordate evolution from echinoderms.[2][3]

Early lifeEdit

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.[4]

In 1895 he married Lucy Ackroyd; they had one son, Walter Lucian Garstang,.[5] and five daughters.

Academic careerEdit

In 1884 at the age of 16, he was awarded a scholarship to Jesus College, 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, Manchester. 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 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.[6]

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 that was later to become the Centre for Environment, Fisheries and Aquaculture Science (Cefas), part of the Ministry of Agriculture, Fisheries and Food (United Kingdom). Garstang instigated a series of detailed fisheries surveys throughout the southern North Sea aboard the RV Huxley, under the auspices of the newly formed International Council for the Exploration of the Sea (ICES).[7]

Garstang was Professor of Zoology at the University of Leeds from 1907 to 1933.[8] The Garstang Building at the university is named in his honour. In 1912 in cooperation with Professor Alfred Denny of the University of Sheffield he established the Robin Hood's Bay Marine Laboratory. The minutes of Sheffield's Faculty of Pure Science on 12 March 1912[9] 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 hypothesisEdit

A. Lancelet, B. Larval tunicate, C. Adult tunicate. 1. Notochord, 2. Neural tube, 5. Pharyngeal slit, 17. Tail

Garstang made the radical suggestion that the chordates evolved from the larvae of another group, whether the larvae of hemichordates or of echinoderms, by progenesis (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 and pharyngeal slits.[10]

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, instead evolving 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 neoteny (also called pedomorphism): retention of juvenile traits or phenotypes 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.[11]


Larval Forms and Other Zoological VersesEdit

First published in 1951, two years after his death, Larval Forms and Other Zoological Verses (ISBN 978-0-226-28423-1) 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, 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 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 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:

Name Theme Speaking parts Lines
The Amphiblastula and the Origin of Sponges comment on theories 64
The Invaginate Gastrula and the Planula comment on theories 24
The Origin of Cnidoblasts and Cnidozoa theory 60
Conaria and Co. How Conaria becomes Velella 18
Mülleria and the Ctenophore theory that ctenophores arose from neotenic larvae of polyclads Mülleria, ctenophore 48
The Onchosphere development 18
The Trochophores development 32
Mitraria's Fan Dance how it feeds 30
The Ballad of the Veliger, or How the Gastropod got its Twist development, theory 40
Echinospira's Double Shell development, comments 126
The Nauplius and the Protaspis theory that the two are related 24
Kentrogon development into Sacculina 16
Isopod Phylogeny comment about ideas a baby woodlouse 12
The Millipede's Egg-tooth theory 36
The Trilobites and After evolution Paradoxides, Lepidocaris, Arthropleura 72
Actinotrocha development 14
Cyphonautes related to Actinotrocha? 4
Echinoderm Larvae and the Origin of Quinqueradial Symmetry development 48
The Pentacrinule comment about larva of Antedon 30
Tornaria's Water-Works description 56
Oikopleura, Jelly-builder description 48
The Ancestry of Vertebrates homologous head parts 8
Leptocephalus brevirostris, the Larva of the Eel development 52
The Axolotl and the Ammocoete theory 16
An Oceanographer's Dream Bermuda as an ideal place for an oceanography base 32
To a Herring Gull description & relationship 84

The Ballad of the VeligerEdit

Veliger larva of sea hare Dolabrifera dolabrifera, with two curved rows of cilia, Garstang's "whirling wheel on either side" and "double screws"
Diagram of veliger larva of sea slug Fiona pinnata

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.[12]

The Ballad of the Veliger or
How the gastropod got its twist[13]

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 abaft,
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 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, 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.


The National Marine Biological Library at the Marine Biological Association in Plymouth holds some of Garstang's archival material (diaries and photographs) and documents relating to the Easter Classes.[14] Leeds University Special Collections also holds some archival material.


  1. ^ Garstang W. 1951. Larval Forms and Other Zoological Verses. Blackwell, Oxford. Reprint: University of Chicago Press 1985.
  2. ^ Kardong, Kenneth V. 2006. Vertebrates: Comparative Anatomy, Function, Evolution, Fourth Edition, pp. 72–75.
  3. ^ Holland, Nicholas D. (1 December 2011). "Walter Garstang: a retrospective". Theory in Biosciences. 130 (4): 247–258. doi:10.1007/s12064-011-0130-3. ISSN 1611-7530. PMID 21833594. S2CID 207378373.
  4. ^ Gurney, O. R; Freeman, P.W.M (2004). "Garstang, John Burges Eustace (1876–1956)". Oxford Dictionary of National Biography (online ed.). Oxford University Press. doi:10.1093/ref:odnb/33341. (Subscription or UK public library membership required.)
  5. ^ "Garstang, Walter Lucian". Who Was Who. A & C Black, 1920–2008; online edition, Oxford University Press. December 2007.
  6. ^ Hardy, A. C. (1951). "Obituary: Walter Garstang" (PDF). Journal of the Marine Biological Association of the United Kingdom. 29: 561–566. doi:10.1017/s0025315400052772.
  7. ^ MAFF (1992). The Directorate of Fisheries Research: Its Origins and Development. Ministry of Agriculture, Fisheries and Food, Lowestoft. 332pp.
  8. ^ Eastham, L. (1949). "Prof. Walter Garstang". Nature. 163 (4144): 518–519. Bibcode:1949Natur.163..518E. doi:10.1038/163518a0.
  9. ^ University of Sheffield. Minutes and Reports of the Faculty of Pure Science, 1912
  10. ^ Carr, Steven M. (2005). "The Garstang Hypothesis". Memorial University, Newfoundland and Labrador. Archived from the original on 23 January 2018.
  11. ^ Satoh, Noriyuki (2016). Chordate Origins and Evolution The Molecular Evolutionary Road to Vertebrates. Academic Press. pp. 17–30 Chapter 2 – Hypotheses on Chordate Origins. ISBN 978-0128029961. The appearance of chordates has been debated for more than 150 years, and many hypotheses have been offered to explain this evolutionary event. These include the annelid hypothesis, the auricularia hypothesis, the calcichordate hypothesis, the enteropneust hypothesis, the inversion hypothesis, and the aboral dorsalization hypothesis. Each had a theoretical basis at the time it was proposed.
  12. ^ Hardy A.C. 1951. Introduction to Garstang W. Larval Forms. Blackwell, Oxford. pp. 8–9.
  13. ^ Goodhart, C. B. (1987). "Garstang's Hypothesis and Gastropod Torsion". Journal of Molluscan Studies. 53 (1): 33–36. doi:10.1093/mollus/53.1.33. ISSN 0260-1230.
  14. ^ The Archives of the Marine Biological Association of the UK: Archived 26 March 2012 at the Wayback Machine

External linksEdit

  •   Works by or about Walter Garstang at Wikisource
  • Larval Forms and Other Zoological Verses at
  • Abalone: Larval defenses
  • Archival material at Leeds University Library