24 



THE HISTORY OF ECOLOGY 



the behavior of starfish. Darwin contributed 

 his classic and essentially ecological study 

 on the earthworm; although, as usual, his 

 observations were exact, his long-range 

 conclusions on earthworms appear to have 

 been erroneous (cf. Keith, 1942). Fabre. 

 Lubbock, the Peckhams. and many others 

 reported penetrating field observations of 

 insect behavior. 



In animal behavior, as in self-conscious 

 ecology and other phases of biology, the 

 decade and a half centering about 1900 

 showed a remarkable outburst of impor- 

 tant biological work which, while firmly 

 grounded historically, was still unusually 

 original. A mature contribution came from 

 Whitman (1898) in his Woods Hole lec- 

 ture on "Animal Behavior" in which he 

 demonstrated a naturalist's sensitivity re- 

 garding the necessity for full acquaintance 

 with the normal behavior of animals before 

 experimenting on them. He insisted, on the 

 basis of pertinent original observations on 

 the behavior of a leech, of Necturiis, and of 

 pigeons, that often the origin and signifi- 

 cance of a given behavior pattern antedate 

 individual acquisitions and are a part of 

 the problem of the origin and history of 

 organization itself, as well as reveal adjust- 

 ment between the animal and its normal 

 environment. Whitman's work on animal 

 behavior, though many of his results were 

 too long left unpublished (cf. Whitman, 

 1919), still influences current programs for 

 the analysis of ecological and other aspects 

 of behavior.* 



The late 1890's and the early years of 

 the present century were enlivened by the 

 controversy that developed between the 

 forced movement, nonadaptive explana- 

 tions of animal behavior of Loeb and his 

 school and the adherents of the more com- 

 pUcated "trial and error" adaptational sys- 

 tem of Jennings. Happily we can now see 

 that the views are largely complementary, 

 and they have already been knit, notably 

 by Kiihn, along with other elements, into 

 a comprehensive system of orientational be- 

 havior (cf. Fraenkel and Gunn, 1940). 



By 1897, Davenport, in his Experimental 



• Whitman was himself an able naturalist. 

 He brought C. B. Davenport to the recently 

 founded University of Chicago, in part to 

 foster field studies, and he had much to do 

 with the early development of C. C. Adams 

 and V. E. Shelford. 



Morphology, which reviewed a much wider 

 field, was able to summarize a hterature 

 in response physiology almost as extensive 

 as in developmental and toleration physiol- 

 ogy. The topics he treated historically in- 

 clude chemotaxis, hydrotaxis, tonotaxis, 

 thigmotaxis (stereotaxis), rheotaxis, geo- 

 taxis, electrotaxis, phototaxis, photopathy. 

 and thermotaxis. Much of the literature 

 cited is from the decades immediately pre- 

 ceding publication, but Davenport calls 

 attention to early work, such as that of 

 Trembley (1744, p. 66) that Hydra viridis 

 moved toward the light even when the 

 lighted slit is turned toward cooler air. 



Some of the ecological queries that such 

 studies helped to answer are: 



1. Do animals have definite reactions 

 that enable them to find the habitat suit- 

 able to their ecological tolerances? 



2. Are animal reactions adaptive? 



3. Is a given behavior pattern innate or 

 conditioned ( learned ) ? 



4. Do any animals other than man seem 

 to be conscious of their behavior? if so, to 

 what extent? Is there a choice of habitats? 

 Do animals show preferences? 



RELATION OF POPULATIONS TO 

 ENVIRONMENT 



General biologists, and even ecologists, 

 who have read thus far, may ask: Is this 

 the history of ecology? Without referring 

 to the discussion of the rise of self-conscious 

 ecology, which will be considered when 

 the background is adequately prepared, an 

 answer may be quoted from an early eco- 

 logical summary. Adams (1913) said: 



"There are also so many degrees and kinds 

 of work that go by the name ecological, which 

 may or may not be, and so many also which 

 are truly ecological but which do not pass 

 under that name, that it is necessary that the 

 student shall be able to see through its di- 

 verse guises and recognize its essential char- 

 acter. Whenever the question arises as to the 

 ecological character of a fact, inference, or 

 conclusion, its ecological validity may be 

 tested in the following way: Do the facts, in- 

 ferences, or conclusions show a response to 

 the inorganic or organic environment: 



"1. As an individual of a species or kind of 

 animal? 



"2. As a group of taxonomically related 

 animals? 



"3. As an association of interacting animals?" 



