The Condor 89:446-4X 0 The Cooper Ornithological Society 1987

BOOK REVIEWS

MARCY F. LAWTON, EDITOR

Connie Hagar: The life history of a Texas birdwatch- Harry C. Oberholser, then gathering information for er.-Karen Harden McCracken. Texas A&M. Univer- his posthumously published The Bird Life of Texas, sity Press, College Station. xvi + 296 p. $18.95. Ludlow Griscom, leading authority on field identifi- This is the biography of a remarkable woman, an cation, and other experts were understandably skep- amateur who contributed substantiallyto ornithology tical of her many reports of birds far from their re- and conservation. Martha Conger Neblett, who later corded ranges, but nearly always she managed to became Connie Hagar, was born on 14 June 1886, in convince them of the accuracyof her identifications, Corsicana, Texas. From her father, an attorney and often by showing them the very birds whose presence mayor of the small town, she learned the names of in her part of Texas they doubted. native trees,shrubs, and wild flowers.Her mother iden- Mrs. Hagar published only articles in local news- tified the butterflies that frequented her flower garden. papersand periodicalsand short notesin ornithological At night she pointed out the constellations,not as first journals. The Checklistof Birds of the Central Coastof lessons in astronomy, but to teach her daughters to Texas, by Hagar and Packard, was written by Fred M. appreciatereferences to them in poetry and mythology. Packard, using Connie’s Nature Calendar. She is fre- Despite this early introduction to nature, music was quently mentioned in Peterson’s A Field Guide to the Connie’s chief interest. She early learned to play the Birds of Texns, especially in the list of accidentals. piano, and from Forest Park Collegein Saint Louis she Connie probably contributed more to ornithology and received a diploma in voice. Until an advanced age, conservationdirectly through her contactswith people she sangand played for churches. than through her writings. Until the infirmities of age After the dissolution of her first marriage, Connie overtook her, she continued to give frequent talks to returned to Corsicana and, with her younger sister, schools, local Audubon Societies, and other groups. became more interested in nature. They decided to She promoted conservationin Rockport, where a wild- learn the names ofconstellations,butterflies, wild flow- life sanctuarywas named for her. She received many ers, and all the birds in Navarro County. In 1923 the honors,including a plaquefrom the National Audubon sisters helped to organize a nature club which they Society. After a lingering illness, she died on 24 No- affiliated with the National Audubon Society. After vember 1973. marrying her secondhusband, Jack Hagar, when nearly Karen McCracken, herself a birdwatcher of wide ex- 40. Connie startedto band birds for the U.S. Biological perience, was long intimate with Connie Hagar. To Suivey. In 1928 she began, in a blank ledger that she gather information for this Boswellian biography of a found in the family library, a “Nature Calendar” which, woman as unique in her own way as Dr. Johnson was in a successionofmiscellaneous notebooks, often pock- in his, she spent two hours daily, five days a week for et-sized engagementbooklets, she would continue for a number of years, interviewing her subject,impressed more than 35 years, jotting down her daily observa- by Connie’s extraordinary memory. She also consulted tions, chiefly of birds; she kept no other records.Hand- the 25 volumes of the Nature Calendar, letters, news- icapped by lack of contactswith more advanced bird- paper clippings, and photographs.The result is a well- watchers, she did not use a binocular until given one written, leisurely story that endears an exceptionally by her husband when she was 48 years of age. dedicated birdwatcher. At times the year-by-year list- In the summer of 1933, Connie and her sister, both ing of the multitudes of migrants that passedthrough of whom sufferedfrom arthritis, went on their doctor’s Rockport or remained to nest tends to tire the reader advice to spend a month at Rockport, a resort village who has no prospect of seeing them; some of this in- on the Texas Coastal Bend. After a second visit the formation might have been condensed. Intimate following summer, Connie, enchanted by the abun- glimpsesof prominent ornithologistsand amusingep- dance of birds, wished to live there permanently and isodes enliven the narrative. A photographer, sent by study them. Readily agreeing, Jack Hagar bought Life magazineto take picturesfor a story about famous Rockport cottages-eight cabins with simple accom- amateur naturalists, was dismayed when Connie ap- modations for visitors. Thus, late in life, begana happy peared,ready to accompanyhim afield, attired asthough partnership of birdwatching wife and cabin-managing for a fashionabletea party. He thought that she should husband that was to continue until he died 27 years be roughly clad for the outdoors, but she assuredhim later. that shealways watched birds in conventional feminine As Connie’s competence in field identification in- dresses.After seeinga snapshotof herself in boots and creased and Rockport became famous as a spot for baggy denim trousers, she decided that birdwatching watching migrating birds and finding rarities, birders did not require “such desperatemeasures.” came in increasingnumbers. Among guestsat the Cot- Despite the lack of mathematics other than simple tages were many leaders in ornithology and conser- counting of birds, I recommend this book to orni- vation. Always generousof her time and eagerto learn thologists and birdwatchers. It shows how, without as well as to teach, Connie, on almost daily trips afield, shooting a single specimen but giving conscientious guided a successionof amateurs and professionals. attentionto field marks,and freely communicatingone ’s

WI BOOK REVIEWS 441 knowledgeand enthusiasmto others, one can contrib- idea provides biology with its central theme, and bi- ute to a science that has been enriched by men and ologistswith their strongestreason for hoping that their women of diverse temperaments and approaches.- subject will eventually be comprehended among the ALEXANDER F. SKUTCH, Quizarra, 8000 San Isi- exact sciences. Especially after Fisher (1930) recon- dro de El General, Costa Rica. structed Darwin’s theory in terms of Mendelian ge- netics and showed that it was eminently reasonable, evolutionary theorists were anxious to test it further, Unfinished synthesis.-Niles Eldredge. 1985. Ox- and to explore its implications. Why does Eldredge ford University Press, New York. vii + 237 p. ISBN wish to displaceadaptation from the centerof the stage? o-19-503633-6. Eldredge believes (p. 184) that the origin of adap- In this book, Eldredge out!ines what he believes to tation is relatively well understood, and traces exclu- be the content of the “neoDarwinian synthesis”of evo- sively to natural selection within populations. As pa- lutionary theory, explains why he thinks it is incom- leontology deals with changesin higher levels of his plete, and suggestsremedies. To avoid setting up a hierarchies,the replacement of one phylad by another “straw man,” he begins bv discussinnthe four books and the like, Eldredge appears to conclude that pa- which, in his opinion, establishedthe synthesis,to wit, leontology cannot reveal anything about the origin of Dobzhansky (1937, 1941) Mayr (1942) and Simpson adaptation, so that a paleontologicaltheory of evolu- (1944). He outlines the further development of the tion must center on a different theme. Of all the slurs synthesis,describing its growing emphaiis on natural I have ever heard on paleontology,and they have been selectionwithin populationsas the only significantcause many and odious, this one takes the cake. To begin of evolutionary change,which he rightly feels unduly with, it is news to me that the origin of adaptation is narrowed the scope of evolutionary-thought. He sees a solved problem. Considering all the chaos of selfish the climax of this trend in Dawkins’s (1976) The Selfish genes,molecular drives, multiplication of transposable Gene, and quite reasonablywonders how evolutionary elements, and the like, it remains to discoverjust how theorists could possibly treat genesas the only signif- selectionwithin populationsmanages to produce such icant units of selection, or discussevolution as if pa- marvels of organismic adaptation. This question re- leontology were an irrelevance. Finally, Eldredge tains its urgency regardlessof whether these marvels sketchesa hierarchicalview of evolution. He envisions are “optimal.” It also remains to learn what role this a hierarchy of replicators (genes, organisms, species, molecular chaos plays in creating suitable “raw ma- monophyletic lineagesof species),and a hierarchy of terial” for natural selection.It surprisesme rather more interactors (cells, organisms, populations, communi- to hear that paleontologyhas nothing to contribute to ties); relationshipsbetween interactors causedifferen- understandingthe origin of adaptation. I believe that tial replication of replicators. Eldrednehimself remarked on the essentialrole of biot- Eldredgewas perhapsmistaken to base this book on ic crises,those widespread extinctions which punctuate a critique of the neoDarwinian synthesis.A critique of the great eras of the fossil record, in facilitating evo- an articulate and coherentbody of thought can be very lutionary progress.On a more theoretical plane, the stimulating.The neoDarwinian synthesis,however, has outcomes of natural selection within populations are no such coherence:indeed, the phrase means some- unpredictable,and some are better for the speciesthan thing different to everyone who would talk about it. others.Presumably, those species where individual ad- Unavoidably, criticismsof some representativesof the vantage most nearly coincides with the good of the synthesisare grosslyunfair to others. In turn, reaction species (in terms of higher speciation rate or lower to such criticisms can distract one ever further from extinction rate: Stanley 1979) are the ones whose de- the proper subject of evolutionary theory, the actual scendantspopulate later geologicstrata. If so, selection processesinvolved in “descent with modification” and between speciesplays an integral role in shaping ad- the diversity of forms they yield. aptation (Leigh 1977) and evolutionary theorists can- Eldredge’s critique, moreover, is based on assump- not afford to neglect the fossil record. tions which guaranteethat he can neither understand What comesof displacingadaptation from the center the synthesisnor convince its proponents of its flaws. of evolutionary theory? Eldredge’s logic has nudged Eldredgeremarks (p. 184) that “. . perhapsthe major him toward an elaborate and sterile formalism, which problem with the synthesishas been that adaptation divorces his evolutionary patterns from the processes was seen as the fundamental theme of the entire evo- which might explain them. Sometimes the problem is lutionary process.” Of course the makers of the syn- merely overmuch faith in the tidiness of his hierarchy. thesis focussedon adaptation! . . however much they Eldredgeposits organismsas the fundamental units of differed in other respects.After all, the apparent pur- natural selection (p. 184) but what are organisms? posivenessof living things, and the aura of design in Consider the implications of the challenge clonal re- their morphology, are their most characteristic fea- production poses for Weismann’s doctrine of the se- tures, as Aristotle well knew. Darwin’s argument that questrationof the germ line (Buss 1983). In organisms these featuresare an automatic consequenceof differ- capable of clonal reproduction, somatic cells can give ential replication with heritable variation, and its im- rise to reproductive tissue, so their contribution to the plication that adaptation and diversity can be account- genetics of future generations is no longer identified ed for without appeal to miracle or mystery, has not with the well-being of the individuals of which they ceasedto astonishor even outragelaymen (and many are now part. A similar conflict arises in colonies of biologists as well). To this day, Darwin’s idea is the socialinsects: if workers can lay eggsof their own, their most astonishingto have emerged from biology: this evolutionary future is no longer absolutely linked to 448 BOOK REVIEWS that of the coloniesto which they belong, and colonies swer to the secondquestion will undoubtedly involve may ceaseto be the primary units of selection. Con- selectionbetween species. strutting hierarchies, however plausible, provides no Although evolutionary theory needs to be reformed, licensefor ignoringhow naturalselection actually works. to attempt this reform by abandoningthe quest to un- A more damaging instance of logic run wild is El- derstandthe origin ofadaptation is to make the subject dredge’s criticism (p. 1580 of Ghiselin’s (1974) analogy fearfullv dull, indeed. to cut the heart out of it. El- of species with business firms. Eldredge argues that dredge’s book suffers grievously by abandoning the speciesare not “interactors” becauseone speciesmay theme that would have lent interest and unity to its live in several different communities, and thus may subject.-E. G. LEIGH, Smithsonian Tropical Re- not be decisively affectedby events in any one of them. searchInstitute, Panama. Yet firms may also operatein severaldifferent markets, perhapsbecause the factors which make for successin LITERATURE CITED one market presumably enhance the prospectsof suc- cess in others where the firm has established itself. Buss, L. 1983. Somatic variation and evolution. Pa- Correspondingly, sufficiently widespread crises may leobiology 9: 12-16. extinguish widely ranging species, or multinational DAWKINS,R. 1976. The selfish gene. Oxford Univ. firms, as the casemay be, favoring the more adaptable Press, Oxford. entities.Possibly because his formalism divorcesspecies DOBZHANSKY,TH. 1937. Genetics and the origin of from communities, Eldredge argues(p. 196) that species.Columbia Univ. Press, New York. ‘1 DOBZHANSKY,TH. 1941. Genetics and the origin of . . if the notions of speciesselection are falsified species.2nd. ed. Columbia Univ. Press,New York. by the observationsthat (1) economic adaptations, FISHER,R. A. 1930. The genetical theory of natural at least, solely involve organismic attributes, not selection. Clarendon Press, Oxford. species-levelproperties, rendering speciesselection GHISELIN,M. T. 1974. A radical solutionto the species meaningless. . and (2) in any case, speciesdo not problem. Syst. Zool. 23: 536-544. seem to be active interactorsin the ecologicalarena, GHISELIN,M. T. 1984. Narrow approachesto phy- then so too is the entire adaptivelandscape metaphor logeny:a review of nine books of cladism. Oxford of adaptiveevolution simultaneously falsified-pro- Surveysin Evolutionary Biology 1:209-222. vided, of course, that speciesare construed as in- LEIGH,E. G., JR. 1977. How does natural selection dividuals and the literal truth of the two observations reconcile individual advantage with the good of above is agreed upon.” the group?Proc. Natl. Acad. Sci. 74:4542-4546. There are many potential conflicts between individual MAYR, E. 1942. Systematicsand the origin of species. advantageand the good of the species-dispersal rate, Columbia Univ. Press, New York. mutation rate, sexratio, and sexualityitselfbeing among PITTENDRIGH,C. S. 1958. Adaptation, natural selec- the better known. As I mentioned earlier, the resolution tion, and behavior, p. 390-416. In G. G. Simpson of such conflicts is probably a major feature of evo- and A. Roe [eds.], Behavior and evolution. Yale lution. A formalism that complicates or hinders their Univ. Press, New Haven, CT. discussion will inevitably obstruct a deeper under- SIMPSON,G. G. 1944. Tempo and mode in evolution. standingof the origin of adaptation. In sum, an overly Columbia Univ. Press, New York. formalistic approach to evolution and phylogeny is STANLEY,S. M. 1979. Macroevolution. W. H. Free- dangerously constricting, as Ghiselin (1984) so elo- man, San Francisco. quently warned. Given the controversy between population geneti- cists and paleontologists,I had best remark that pos- iting the origin of adaptation as the central problem of Phenetics: evolution, population, trait.-A. V. Ya- evolutionary theory (Pittendrigh 1958) does not imply blokov (trans. M. J. Hall). 1986. Columbia University that natural selection within populations is the only Press, New York. xi + 171 p. relevant causeof evolutionary change.Whatever fault This work is the third edition, heavily revised, of the I may find with his book, Eldredge is quite right to well-known Russian book Fenetika by Alexey V. Ya- insist on the incompletenessof modem evolutionary blokov, a leading Soviet evolutionist and ecologist. thought and to associatethis incompletenesswith ne- “Phenetics” for Yablokov is not the “phenetics” of glect of the various possiblelevels of evolution, which numerical taxonomists, who applied the term loosely can in turn be blamed partly on ignorance of paleon- to a groupof numerical methods. Rather, it is the study tology. Those who think we know all we need to about of “phenes.” Phenes are alternative variants of genet- evolution are welcome to explain (1) how a scientific ically determined traits, thoughthe underlyinggenetics team from another galaxy, with full accessto Earth’s may involve suchcomplexities as pleiotropy and poly- fossilrecord in proper stratigraphicorder, but no access genes.The variation can be continuous,as for size and to living or recently dead organisms,could deduce the color, so long as each individual can be assignedun- principlesofinheritance from the fossil record, and (2), ambiguouslyto one phene-class. how large, and how subdivided, a landmass was re- One should not worry about the introduction of new quired to allow evolution of mammals from reptiles. jargon, since “phene” was first defined, very similarly The absenceof an answer to the first question suggests to Yablokov’s definition, in 1909 by Johannsen, the that the relevance of genetics to paleontology, and Dane who also defined “gene,” “genotype,” and “phe- therefore to evolution, is not fully established,the an- notype.” In a brief historical chapter, Yablokov dis- BOOK REVIEWS 449

cusseswhy study of genes and genotypesrapidly sur- Yablokov conceives of phenetics as primarily a passedstudy of phenes and phenotypes.The essential pointer to interesting situationsfor further genetic and reasonwas that the genewas the centralunit ofgenetics, ecologicalresearch. Most insights to date and for the a sciencethat quickly became exciting and productive near future rest on the geographicaldistribution of var- as the physical nature of genesand chromosomeswas ious phene-classesso it is fitting that by far the largest elucidatedand the rather simple genetic basis of some chapter is on “phenogeography.” Usually one is seek- traits become manifest. However, these latter were ex- ing regions of abrupt changesin the distribution of ceptional and the generalpicture was of a complicated frequencies of phene-classes.For example, an early path from gene to phene. This complication was the study by Serebrovskii focussedon the phenes of color spur to a focus on genes and neglect of phenes-it and comb shape in semiwild chickens in Dagestan seemed as if fruitful study of phenes would require a (North Caucasus). Phenetic frequencies changed clearer understandingof their genetic bases. abruptly at the Avarskoe Koisu River, which flows in Why resurrect the study of phenes now? What in- a gorge several hundred meters wide. Apparently sights can they yield, especially if the underlying ge- chickens cannot cross the gorge, though other gorges netics have not been elaborated?In fact, as Yablokov do not constituteregions of phenetic frequencychange demonstrates,phenes have not really been buried from and, by implication, barriers to dispersal. A similar 1909 until the presentand much evolutionary and eco- example for a much more widely distributed bird, the logical researchis actually phenetic: phenotypic traits yellow wagtail, is presentedfrom untranslatedresearch are assumedto have a geneticbasis but the underlying by Timofeeff-Ressovskyet al. geneticsare not worked out. Instead, evolutionary de- In addition to taxonomic uses, such as the delinea- ductionshave often been based directly on phenotypic tion of infraspecific categories,phenogeographic data patterns.One example presentedin detail (studiedphe- can lead to interestingecological questions. A featured neticallyby Timofeeff-Ressovskyand Yablokov in 1973 example is color variation in the rustic autumnal moth in an untranslated Russian paper) is the relation be- in the Shetland Islands, which has a melanistic and a tween precipitation and clinal variation in the hybrid non-melanistic form that were studied by Kettlewell zone between the Baltimore and Bullock orioles de- and Berry. A small valley on ShetlandMainland marks scribedby Rising in 1970. Many other examples,often an abrupt shift from predominantly melanic moths in from the eastern European literature, make the same the north to predominantly non-melanics in the south. point. Mark-and-recapture experimentsshowed that individ- The first goal of Pheneticsis to formalize methods ual moths frequently flew far enoughto crossthe valley for detecting phenes, assigningindividuals to phene- but very rarely did so. Exactly why they do not cross classes,and analyzing resulting phenetic patterns. Ya- the valley, and why melanic moths are favored north blokov gives examples of several graphical and statis- of the valley, are unansweredquestions though several tical methods of presentingphenetic data to facilitate hypotheseshave been proposed in the wake of the detecting patterns and also treats the problem of ana- phenogeographicobservations. lyzing several phenes (say, color and size) simulta- Probably the American edition of Pheneticswill gen- neously for the same group of populations. erate most interest in two ways. First, the focus on the One might have expectedthat the growth of genetics relationship between geographic pattern and micro- and concomitant better understandingof some gene- evolutionary processeswill help to strengtheninterest to-phene pathways would cause phenetics to be sub- in the populational aspectsof biogeography, such as sumed into genetics.In other words, for what specific the significanceof clines and stepped clines. Though questions is analysis of phenes likely to provide an far from moribund, this branch of biogeographyhas answer?Yablokov arguesconvincingly that there are taken a distinct back seat for two decadesto the mul- only a handful of cases for which sufficient genetic tispeciesquestions associated with the equilibrium the- understandingexists so that study of phenotypic traits ory of island biogeographyand with variants of vicar- (phenes) will still be important for years to come in iance biogeography. New methods such as those addressingmany evolutionary questions:the nature of proposedby Yablokov and greatly increaseddata sug- natural selection, the importance of gene flow, muta- gestthat population-oriented biogeographymay be an tions, and population fluctuations,the evolutionary re- extremely fruitful area of researcheven though it is an lationships of different populations and species, etc. old field. Second,a comprehensive(though short) work Ontogenetic noise and phenotypic plasticity bedevil by a respectedRussian scientist must intrigue thought- phenetics,just as they bedevil genetics.However, Ya- ful westernreaders. Especially in evolution and ecology blokov is concernedthat we not overestimate the dif- the traditions have grown sufficiently apart that there ficultiesposed by this indeterminacy. His key argument are somewhat different problems, different perspec- here is that the complexity of interaction of external tives on common problems,and a separate(and usually factors during ontogeny is so great that their effects untranslated)literature. Any American who has worked must somehowcancel one another out, much as do the in an active foreign (especially non-anglophone) lab- movements of particlesin Brownian motion. The evi- oratory knows the excitement and novel insights de- dence he marshals in support of this proposition is rived from discussionswith critical and dedicated sci- quite meagre, given its importance in justifying the entists working on problems similar to one’s own, study of phenes. However, the many cogent phenetic Pheneticsconveys much the samefeeling without one’s examples he presentssuggest that this defect may not having to travel to Moscow.-DANIEL SIMBER- be cruciallydebilitating, but rather a point to be argued LOFF, Department of BiologicalScience, Florida State and perhaps remedied. University, Tallahassee,FL 32306.