2013/06/28 by Frank N. Egerton · 1 citation
Arts and Humanities · Environmental Science · #History of Science and Natural History #Polar Research and Ecology #American Environmental and Regional History
paper · pdf · doi:10.1890/0012-9623-94.3.222
Click here for all previous articles in the History of the Ecological Sciences series by F. N. Egerton Ernst Haeckel, one of the world's best-known and most-read zoologists, named and defined a new science, “Oecologie” (1866, II:286). Was he, therefore, an ecologist? Robert Stauffer (my dissertation advisor) said yes (1957), but wrote that what was good in his ecology he got from Darwin, and what was misguided, he got from himself. There is some truth to that, but the story is more complex. (Stauffer was more interested in Darwin than Haeckel.) Haeckel had already absorbed Alexander von Humboldt's ecological perspective long before he read Darwin's Origin, though Humboldt's ecology lacked Darwin's concept of competition. If Haeckel is accepted as one founder of ecology, ecologists will want to know why he remains a controversial figure and how to evaluate the controversies. An introduction to these matters follows at the end. One, arranged on official lines, offered to the sympathetic observer all the species in “typical” specimens, as radically distinct forms, each decked with its pretty label; the other was a private collection, only shown to one trusted friend, and contained only the rejected kinds that Goethe so happily called “the characterless or disorderly races [Geschlechter], which we hardly dare ascribe to a species, as they lose themselves in infinite varieties,”… In this a large number of specimens arranged in a long series, illustrated the direct transition from one good species to another. …among the seaweed there were to be found many of the dearest little animals, specially charming polyps (Tubularia, Eudendrium, Actinia), sea spiders (Pycnogonum littorale), and a very strange member of the tunicata (Amaraecium rubicundum)… we saw a few seals at no great distance from us … The strangest thing we caught was a beautiful green garpike (Belone vulgaris) with green bones, and I have been sitting all day examining the highly peculiar ova under the microscope; besides that we caught swimming crabs, a few shrimps, which are very scarce here, sea devils (Cottus scorpius), sprats, a quantity of different plaice and haddock (torsk), together with other fish. In 1855 Haeckel published his first article in an important new journal edited by Müller (Richards 2008:40). In summer 1856, he accompanied Professor Albert Kölliker to Nice, France, where he found the marine fauna both rich and remarkable (Haeckel 1923:416). He wrote a doctoral dissertation on histology of river crayfish. He returned home to Potsdam in 1858, with an M.D., to practice medicine, and hated it. He spent 1859–1860 in Italy, studying marine life. … what high esteem and profound respect I hold the discoverer of the “Struggle for life” and of “Natural selection”. Of all the books I have ever read, not a single one has come even close to making such an overpowering and lasting impression on me, as your theory of evolution of species. In your book I found all at once the harmonious solution of all the fundamental problems that I had continually tried to solve ever since I had come to know nature as she really is. Since then your theory—I can say so without exaggeration—has occupied my mind every day most pressingly, and whatever I investigate in the life of humans, animals or plants, your theory of descent always offers me a harmonious solution to all problems, however knotty. Thus began a warm friendship that lasted the rest of Darwin's life (Gliboff 2008:155–188). Ernst Haeckel in 1860. [Wikipedia] In 1861 Haeckel published part of his research to qualify for appointment to the faculty of Jena University. He owed the position to Carl Gegenbaur (1826–1903), whom he had known at Würzburg. Gegenbaur became his closest friend and collaborator (Nordenskiöld 1928:499–503, Uschmann 1959:27–33, Coleman 1978). Aside from numerous journeys, Haeckel spent the rest of his life at Jena. In 1862, he was promoted to associate professor and Director of the Zoological Museum, and on 18 August he married his cousin, Anna Sethe (1835–1864)—the great love of his life—who died a year and a half later, on Haeckel's 30th birthday. He continued studying specimens collected in Italy, which led to his monograph, Die Radiolarien 1862; the impressive illustrations in it, which he drew, undoubtedly helped win a gold medal from the Leopold-Caroline Academy. In this work, he speculated on species relationships and genealogy and continued doing so for the rest of his life (Wilson and Doner 1937:57– 58). He sent Darwin a copy. Darwin had already seen Huxley's copy, and Huxley sent Haeckel an enthusiastic letter and some specimens (Di Gregorio 2005:72–73). … constituted the first attempt to apply the general doctrine of development to the whole range of organic morphology (Anatomy and Biogenesis), and thus to make use of the vast march onwards which the genius of Charles Darwin has effected in all biological science by his reform of the Descent Theory and its establishment through the doctrine of selection. Radiolaria. Haeckel 1899–1904. Lithographer Adolf Gilitsch transformed Haeckel's sketches into forms with somewhat exaggerated colors. Every group of organisms that Haeckel studied became grist in his theoretical mill: he had a good command of Greek and coined many terms from Greek roots, including oecologie, phylum, ontogeny, phylogeny, and protista. He pioneered the construction of phylogenetic charts that indicated which groups of species were most closely related, and their likely ancestors (Dayrat 2003). He thought radiate rhizopods were the lowest level of animals, and so proper subjects for discussing the relationships between plants and animals. He coined the terms protista for all unicellular organisms. Other zoologists offered other names, but Haeckel's has stuck (Rothschild 1989:282–291). He also formulated a biogenetic law (ontogeny recapitulates phylogeny), which was a useful hypothesis to investigate. However, as a Haeckelian pronouncement it went too far and elicited far more disagreement than agreement (Churchill 1980, Rinard 1981, Rasmussen 1991). He provided a chart showing how to organize zoology, with ecology and animal geography placed under a broader heading of Animal Physiology. By ecology, we mean the whole science of the relations of the organism to the environment including, in the broad sense, all the “conditions of existence.” These are partly organic, partly inorganic in nature; both, as we have shown, are of the greatest significance for the form of organisms, for they force them to become adapted. Among the inorganic conditions of existence to which every organism must adapt itself belong, first of all, the physical and chemical properties of its habitat, the climate (light, warmth, atmospheric conditions of humidity and electricity), the inorganic nutrients, nature of the water and of the soil, etc. As organic conditions of existence we consider the entire relations of the organism to all other organisms with which it comes into contact, and of which most contribute either to its advantage or its harm. Each organism has among the other organisms its friends and its enemies, those which favor its existence and those which harm it. The organisms which serve as organic foodstuff for others or which live upon them as parasites also belong in this category of organic conditions of existence. In our discussion of the theory of selection we have shown what enormous importance all these relations have for the entire formation of organisms, and specially how the organic conditions of existence exert a much more profound transforming action on organisms than do the inorganic. The extraordinary significance of these relations does not correspond in the least to their scientific treatment, however. So far physiology, to which this [science] belongs, has, in the most one sided fashion, almost exclusively investigated the conserving functions of organisms (preservation of the individual and the species, nutrition, and reproduction), and among the functions of relationship [investigated] merely those which are produced by the relations of single parts of the organism to each other and to the whole. On the other hand, physiology has largely neglected the relations of the organism to the environment, the place each organism takes in the household of nature, in the economy of all nature, and has abandoned the gathering of the relevant facts to an uncritical “natural history,” without making an attempt to explain them mechanistically. This indubitable and highly important fact is revealed most strikingly in the tendency on average for the absolute number of organic individuals populating our world to remain constant, and for only the relative numbers of the individual species to alter continually in relation to each other. . . . yet Chorology as a whole remained, as far as their labours were concerned, only a desultory knowledge of a mass of individual facts. It could not be called a science as long as the causes for the explanation of these facts were wanting. These causes were first disclosed by the theory of selection and its doctrine of the migrations of animal and vegetable species, and it is only since the works of Darwin and Wallace that we have been able to speak of an independent science of Chorology. Monophyletischer Stammbaum der organismen. The first phylogenetic chart for the evolution of life. Haeckel 1866, II: plate 1. In other words, animal geography could tell us that Darwin's finches on different Galápagos Islands were different species, but not tell us why. Biologists have accepted his new science of ecology but have not considered his chorology a new science. Some biologists adopted Haeckel's term, but most simply added Darwin's new explanations to the preexisting science of biogeography. Ernst Haeckel and Nikolai Miklucho-Maklai, one of three student assistants who joined Haeckel's private expedition to the Canary Islands, 1866. Ernst-Haeckel-Haus, Friedrich-Schiller-Universität, Jena, Germany. By ecology we mean the body of knowledge concerning the economy of nature—the investigation of the total relations of the animal both to the inorganic and to its organic environment; including, above all, its friendly and inimical relations with those animals and plants with which it comes directly or indirectly into contact—in a word, ecology is the study of all those complex interrelations referred to by Darwin as the conditions of the struggle for existence. Still dissatisfied with the progress of his reform, in 1906 he published a one-volume abridgement of Generelle Morphologie. Repetition and a clear definition possibly helped establish his term, oecologie, for there were alternative suggestions available (Schurig and Nothacker 2001). In 1907 he established a Phyletisches Museum at the university, partly financed with royalties from his books (Uschmann 1959:165–175, Di Greogorio 2005:526–527). When his successor as director of the museum did not run it as Haeckel had expected, he turned his home into a museum—Ernst Haeckel-Haus (photos of both buildings in Smit 1967 and Richards 2008). The first general survey of animal ecology was not by Haeckel, but by Würzburg professor of zoology Karl Gottfried Semper (1832–1893), who had been a student there during some of the time that Haeckel was (Mayr 1975). Semper spent the period December 1857–May 1865 exploring the Philippines and Palau, and afterwards published Reisen im Archipel der Philippinen (10 volumes, 1868–1905; see Johnson 1969). In 1877 he delivered 12 lectures at Lowell Technological Institute in Boston, then prepared them for publication in both German (1880) and English (1881). His Animal Life as Affected by the Natural Conditions of Existence never used Haeckel's term oecologie. He could hardly have been ignorant of Haeckel's Generelle Morphologie, which aspired to reorganize zoology. One Haeckel biographer (Di Gregorio 2005:292) Semper as he saw with remarkable which of the were to one or the other of the and them One of was in der to Animal Life and as He did a Haeckel of a and his on Haeckel was to a long Semper accepted Darwin's his was Haeckel's (Di Gregorio Haeckel defined the new ecological but did he also contribute to the of The of and Darwin He never on a expedition but at Jena he some in to or and Di Gregorio He published of some journeys, which can be for to those by his and Darwin, in their Haeckel's upon and sent an 1859–1860 German was a both Haeckel, and of his to was a and Haeckel returned from with numerous of some of which in his His only became a who in Haeckel in Since was a Haeckel's was into English to It will be our of his It have been by some of his and a However, it is by his for what he found and his of the It was a it a each for the there and by the he was director of the zoology museum and specimens and for it. It was a good is a and He began for this at and spent the summer use of and and he with of including books and (Haeckel He Jena on by to and then by from to the new He had been to the in to study the marine life of his but that had not his for which in the between and at first and then in and (Haeckel He a of at to study the that the were and the were and though the was He spent much of the of his was “the and day of my life I first in a and in at life in and (Haeckel and he had only where he spent a very on the of a German It helped that both the and he went were Haeckel's for plants had never he had become a and his book more to plants than animals, simply the plants he and a for a he had of what his and Darwin had the and the his He also saw and but for to as he have to his of his to he was with one with the and he found time to Haeckel was as much as and Darwin had been on their but that of his book is here, though his of they can be (Haeckel ecologists who the he thought could or its that figure be the of is and remains indicated that the half of was more than it was in by a in he a to on the of where he spent a in a at the of the The was very The term referred to and had more or or with above from the water (Haeckel The and from the river and the also caught and organic and were as of were as the of Haeckel did not where he One Haeckel which his had seen and and large Haeckel's but he did a green a which had a that could on (Haeckel Haeckel by to the in at and on to a in had spent species Haeckel's had been by a German who had died in and had the to Haeckel The who Haeckel, that had at least species the at for the most for warm and the of green in a great of and with and by with and of with and Haeckel had that much as to the of and he continued (Haeckel had already (Haeckel how and green in the of on as as at with and of though the Haeckel were He it by selection the of from that of its the will it be seen by its the more can it upon its and the more it is and for the struggle for existence.” Haeckel whom he a and more they with a for He had not his at the but did in and found it very to see marine life as it However, in my life have I been so and as a few of and at and I from the for (Haeckel he thought the was the he had available a with to his and a though there were no on the (Haeckel His revealed many species to those of the of Italy, but also new species. The had shown that species are more than species (Haeckel were great for but they also of into the which its on most parts of the its is and that and of the is which is the first and of life for many marine those of the (Haeckel The of specimens collected at and at were a rich for all his in them (Haeckel and he had time for an expedition into the high with who all the at By and a of the In much of it was home to and but in established a and in than and the and more and The had been a to and for (Haeckel Haeckel at the and they into where with or These so that was the only that could to (Haeckel had which was the of the that The contained that the and had long that from (Haeckel at where the at a and where Haeckel saw He and then a for to the the in had once been but had to river (Haeckel There were large and and large and were a but not in On the to Haeckel arranged and his and In he that on the and on the and he found the more (Haeckel his were more than many but by no all of this live in a relation with unicellular of the group In the they live the in the and on the other hand, the but the in the a form of these unicellular to the in the and to a of the or this life is in very many of the greatest significance for both the for the animal the not only with and but also with and other of for their on the other the vegetable of the the its most important of and as as for it is not only but has been that for a long period in of to the the each other with and are to each other by of the nature of their This is not for the existence of the for in many species the number of is very and in many others they are wanting. Haeckel, on the collected by during the plate by He then this with that of Karl in wrote on the that and he named and the and The concept in the of on the relationship between and in but it was Albert who coined the term in and used the term in the whole by for the is and that the general are not only but also a very on the most important problems of that Haeckel was and Di Gregorio and of have Haeckel's (Stauffer and Haeckel's and other marine biologists to more time than they have without such a into the and of sea Haeckel 1899–1904. This new species of the one of the most beautiful of the was on under the in the It its as a of von the friend of nature, who has the der in numerous by Since was to his of their their relationship was through the between them than a of them were first published without either and into English with his and a for what my friend Gegenbaur had said the book of the to not a He my from first to but has always been of the that these are to which the great has no he too has always the of my By too has always Haeckel in to Haeckel's the with to to This letter and the one are not in the all of their is in the these were edited or he in with what he Haeckel Die was the for who was not the only professor with it (Di Gregorio Richards Haeckel never from the of my in the of the to their and even I as a student to my to the that I all the of science. It was only I had and into the of life and its as a I with all the of and as a student with all the of nature, that I became the most a and a The of his first had also been a to his (Richards have a without it as an but Haeckel did in Die which was a and into (Richards He with a He not have or not the that one can more with than but he have that Darwin was and including to evolution (Richards Haeckel to with a Haeckel was a and and had in numerous that he was by the and the as the of science. on the other hand, an German and The in the of the to which the new doctrine of evolution thus the which between the different world It the of as as that of it the with the theoretical it science with science to form an general or total science. In 1906 Haeckel the An the and the He Haeckel's and to (Richards The German was by Haeckel's I hold these and to be important in German One not that they have always for and the of as as the On the other hand, he had no more for than for and he thought be into German Haeckel was a and that his German However, the his books 1991). to on his of for was to a book with the accepted the in without and Richards his in and who Haeckel as a founder of their science can that and other have not they as that one Haeckel's he remains also an important founder of I and der Ernst-Haeckel-Haus, Friedrich-Schiller-Universität, Jena, for their and for the of Haeckel and