Book Reviews

O Patterns! Wherefore art thou

patterns? (with apologies to

Shakespeare)

Woods, C. A. and Sergile, F. E. (eds)(2001) Biogeography of the West Indies:Patterns and Perspectives. Second edn.CRC Press, Boca Raton, Florida, USA.xxiv + 582 pp., figs, tables, line diagrams,halftones, index. Hardback: Price$139.95. ISBN 0-8493-2001-1.

As a postdoctoral researcher at theUniversity of Texas in Austin in 1989, Ihad the pleasure of listening to a visitingcolleague’s seminar on systematics andbiogeography. The colleague did a fine jobat disparaging historical biogeography,with his most memorable comment beingthat anyone who gives a talk on historicalbiogeography should be required to wear aparty hat. After all, what is the differencebetween a story spun by a raconteur at asocial event and a scenario spelled out bya biogeographer? Neither is necessarilyconstrained by facts or even open tofalsification.

But is that always true for biogeogra-phy? In the opening chapter, Woods (oneof the editors) states that the emphasis ofthis book is on patterns. Many historicalbiogeographers would argue that it is inthese patterns that notions of falsificationand corroboration can arise, allowing usto doff our party hats. The patterns Irefer to of course are cladograms (distri-bution tracks are of penultimate import-ance), and the patterns are of relationshipand congruence among taxa and betweentaxa and areas. Does this volume have thecherished trees that would allow us toremove the party hat from its binding?Unfortunately the answer is no. There areexactly three cladograms and only in oneof these chapters (Judd) does the authorconduct further analysis (Brooks Parsi-mony Analysis) on the cladogram to seekinsight into the historical biogeography ofhis study group. The promise of patternswas not satisfied, but I suppose �patterns�can mean something less explicit thantestable hypotheses in the form of clado-grams.

�The lady doth protest too much,methinks.�(Hamlet, Act 3 scene 2)

The second chapter (Hedges) purports tobe an overview of the historical biogeogra-phy of the West Indies. In reality, thechapter is an overtly defensive attempt toward off the severe criticisms that Iturralde-Vinent & MacPhee (1999) laid on theimmunological distance-molecular clockbased hypotheses put forth by Hedges et al.,(1992; Hedges, 1996) and again inthis book (Chapter 11). In fact in thischapter (Chapter 2) Hedges so often claimsinnocence that I could not but help think ofthe above quote from Shakespeare. IfIturallde-Vinent & McPhee were the onlycritics of Hedges’ notions, then perhaps thecase would remain open to debate, but thepoverty of the immunological distance-molecular clock approach to historicalbiogeography had already been revealed byPage & Lydeard (1994), Crother & Guyer(1996), Guyer & Crother (1996), andPregill & Crother (1999). Chapter 13 in thisbook (Portell et al.) fits nicely in thatit supports the Jamaican immunolo-gical comparisons as strawmen tests ofvicariance (Crother & Guyer, 1996).

To return to patterns, Hedges (p. 21)says, �Since it was proposed, the vicariancemodel has been proven difficult to test.�There are at least a few ways to view thisperception. Granted, vicariance is certainlydifficult to test using approaches that lackcladograms and congruence, and insteadsimply employ scenario construction. Per-haps the patterns are so complex thattesting for vicariance is a hopeless cause,even with cladograms. Or maybe Hedgesmeant that vicariance is simply an untest-able proposition. Whatever Hedges meant,only the former is true, i.e. scenario con-struction (including molecular clock basedideas) cannot test vicariance. What is cer-tain is that the data are available, areslowly increasing, and that vicariance istestable, if anything at all is testable inhistorical biogeography (Nelson, 1974).Crother & Guyer (1996) and Warren &Crother (2001) used cladograms from 11and 15 taxonomic groups, respectively, to

test the broad ancient vicariance pattern inthe Greater Antilles. The studies focusedon BPA but also used CAFCA andCOMPONENT to construct a single treeand both studies recovered the expectedvicariant signal as described in Buskirk(1985). The patterns are there and muchmore work is needed to tease out thedetails for each group.

Other taxa discussed in this volumeappear perfect for cladogram constructionand hypothesis testing. Rysodine beetles(Bell), butterflies in the genus Calisto(Miller & Miller) and parrots (Williamset al.) are all excellent candidates. Thequality of the capromyid rodent clado-grams is uncertain because the morpholo-gical analyses appear to have been rootedwith ingroup taxa (based on the results)and the molecular data are from themitochondrial genome which may notreflect species history (Taggart et al.,2001; Ballard et al., 2002). Additionalanalyses of the capromyid rodents wouldbe most welcome. Another mammal groupaccompanied by cladistic analysis is themegalonychid sloths (White & MacPhee).The cladogram is not congruent withBuskirk (1985) but this is not surprisinggiven that the age of these sloth taxa pre-cludes their lineage’s participation in theancient Antillean vicariant events. Becauseof the young age of the sloth fossils, it onlyfollows that White and MacPhee rejectearly vicariance and posit a later land-bridge (GAARlandia) to explain the pres-ence of sloths in the West Indies.

To leave patterns aside, the volume is aneclectic collection of papers on the WestIndian region. Six of the chapters are littlemore than expanded abstracts. Thearcheological papers were interesting toread but I can not critically comment onthem.

�Though this be madness, yet thereis method in�t.’(Hamlet Act 2, scene 2)

In summary, it is pleasing to see thatWest Indian biogeography is still worthyof a dedicated volume. My wish would bethat a future volume would focus on the

Journal of Biogeography, 29, 1263–1265

� 2002 Blackwell Science Ltd

historical patterns and involve advancedanalyses of multiple taxonomic groups’cladograms (especially from plants) andfrom these analyses attempt to ascertainthe histories of the individual groups. Isn’tthat the goal?

Brian I. Crother

Department of Biology,Southeastern Louisiana University,

USA

REFERENCES

Ballard, J.W., Chernoff, B. & James, A.C.(2002) Divergence of mitochondrialDNA is not corroborated by nuclearDNA, morphology, or behavior inDrosophila simulans. Evolution, 56,527–545.

Buskirk, R.E. (1985) Zoogeographic pat-terns and tectonic history of Jamaicaand the northern Caribbean. Journal ofBiogeography, 12, 445–461.

Crother, B.I. & Guyer, G. (1996) Carib-bean Historical Biogeography: Was theDispersal-Vicariance Debate Eliminatedby an Extraterrestrial Bolide? Herpeto-logica, 52, 440–465.

Guyer, G. & Crother, B.I. (1996) Addi-tional comments on the origin of theWest Indian Herpetofauna. Herpetolo-gica, 52, 620–622.

Hedges, S.B. (1996) Historical biogeogra-phy of West Indian vertebrates. AnnualReview of Ecology and Systematics, 27,163–196.

Hedges, S.B., Hass, C.A. & Maxson, L.R.(1992) Caribbean biogeography: mole-cular evidence for dispersal in WestIndian terrestrial vertebrates. Proceed-ings of the National Academy of Science(USA), 89, 1909–1913.

Iturralde-Vinent, M.A. & MacPhee, R.D.E.(1999) Plaoegeography of the CaribbeanRegion: implications for Cenozoic bio-geography. Bulletin of the AmericanMuseum of Natural History, 238, 1–95.

Nelson, G. (1974) Historical biogeogra-phy: an alternative formalization. Sys-tematic Zoology, 23, 555–558.

Page, R.D.M. & Lydeard, C. (1994)Towards a cladistic biogeography ofthe Caribbean. Cladistics, 10, 21–41.

Pregill, G.K. & Crother, B.I. (1999)Ecological and Historical Biogeogra-phy of the Caribbean. Caribbeanreptiles and amphibians (ed. by B. I.Crother), pp. 335–356. AcademicPress, San Diego.

Taggart, T., White, M. & Crother, B.I.(2001) Palm-pitviper (Bothriechis) phy-logeny, mtDNA and consilience. Cladis-tics, 17, 355–370.

Warren, B. & Crother, B.I. (2001) Meto-dos en biogeografia cladistica: el ejem-plo del Caribe. Introduccion a labiogeografıa en Latinoamerica: teroıas,concepts, metodos, y aplicaciones (ed.by J. L. Bousquets and J. J. Morrone),pp. 233–243. Las Pensas de Ciencias,UNAM.

The highs and lows of tropical

forest canopies

Linsenmair, K.E., Davis, A.J., Fiala, B. &Speight, M.R., (eds) (2001). Tropicalforest canopies: ecology and management.Kluwer Academic Publishers, Dordrecht,Boston, London. 370 pp., plates, figs,tables, index. Hardback: Price EUR125.00, 79.00, USA $115.00. ISBN079237049X.

Tropical Forest Canopies (TFC) should beread almost cover-to-cover by anyoneseriously interested in canopy habitats. Thebook includes well-crafted reviews, andyet, in contrast to its predecessor, ForestCanopies (Lowman & Nadkarni, 1995),most chapters also offer new results.Research based in Central America oftenseems to dominate tropical ecology, butthe studies presented in this book bring tomind the comment by Paulo E. Vanzolinithat �the tropics are not a plot of con-venient forest in Costa Rica.� I found itrefreshing to read about projects fromBorneo, French Guiana, Venezuela, andelsewhere. This might reflect the fact thatnine out of 10 TFC authors are from out-side the USA: TFC is a reprint of volume153 of the Dutch-based journal PlantEcology (� 2001) and its contents origin-ated as a series of lectures at Oxford in1998, sponsored by the European ScienceFoundation.

Beyond occasional line-editing prob-lems, the content and style of TFC aregenerally more uniform than in Lowman& Nadkarni (1995) which contains art-icles that vary from those comprehensi-ble to undergraduate biologists to thoseimpenetrable except to authorities in anarrow subdiscipline. TFC is aimed atecologists and is especially useful for

canopy specialists because it takes overwhere Lowman & Nadkarni (1995) leftoff, with chapters surveying the experts orthe literature on selected topics to addresstrends and to discuss problems arisingfrom the climbing methods researchershave used, or the conceptual issues needingfurther study: I heartily recommend theoverview by Martin Barker and MichellePinard and the chapter on invertebrates byYves Basset and on vertebrates by RolandKays and Allen Allison.

There is a widespread tendency for for-est scientists to treat �canopy biology� assynonymous with forest research. Happily,all the chapters of TFC spell out their focuson forests, and thereby explicitly excludeother ecosystems, whether natural oragricultural, terrestrial or marine, forwhich the term canopy can be (and oftenis) applied. Nonetheless, I was disappoin-ted that none of the chapters in TFC tookon the challenge of comparing the cano-pies of forests with those of other (trop-ical) systems. Many concepts and modelsdeveloped for canopies as diverse asmowed lawns, kelp forests, and biofilmsmight be applied to forests (Moffett,2001), but a continued myopia has keptthese literatures almost entirely separate.

Frans Bongers’ assessment of tropicalrain forest canopy structure should havebeen Chapter 1, especially given his is theonly chapter to attempt to clarify thebook’s topic by asking what a �forestcanopy� is. Bongers shows the word is usedin varied ways. He employs the termhimself to describe the �total above groundpart of the forest�, including thereby theherbs down at our feet, the approach I alsoprefer (Moffett, 2000). Bongers doesn’tgive a reason for his choice, but this broaddefinition avoids arbitrary and imprecisedelimitations within above-ground plantparts; reduces semantic entanglementswhen �canopy� is used in combination withterms like �epiphyte� that are applied toorganisms regardless of height or locationon the host or the host’s growth form;includes the more narrow definitions of�canopy� as special cases; and allows forready comparison with non-forest cano-pies.

Throughout the literature on �canopybiology�, few researchers spell out theirviews on the word �canopy� with precision,so that it is often difficult or impossible totell if two articles are in fact discussing thesame thing, and so can be compared.

� 2002 Blackwell Science Ltd, Journal of Biogeography, 29, 1263–1265

1264 Book Reviews

Elsewhere in TFC, Bonger’s definition isused (at least implicitly) only in the chap-ters on throughfall by Calder and byChappell et al. In all other chapters forwhich I could glean information, there arephrases to suggest that, for those authors,canopy biology variously encompasseseither (1) parts of the forest beyondeveryday human reach; (2) all trees (or treecrowns) in combination, regardless of theirheight (but in contrast to Bongers, Calderand Chappell et al. one assumes not herbsand shrubs); or (3) the uppermost treecrowns alone. Other variations, such asusing canopy to describe the outermostleaves (i.e. the �outer canopy�), are absentfrom TFC but common elsewhere. SomeTFC authors try to reduce ambiguity withthe phrase �upper canopy� but never saywhat a �lower canopy� might be. Confusionabounds in the literature: it is not unusualto see �understorey� used to describe astratum that is separate from the canopystratum (or strata), and then, at anotherpoint in the same article, to find the sameword used to describe �part of a canopy� orperhaps equivalently, �one of the severalcanopy layers’. In her chapter in TFC,Margaret Lowman questions the overallvalue of distinguishing the study of thecanopies of forests from the general topicof forest ecology; indeed, in reading TFC,it is an interesting exercise to notice whereit is possible to remove the word �canopy�(or to change the word to �forest�) withoutchanging the meaning. In fact, the accu-racy of many statements appears to beimproved by such an edit. This is notablytrue for the chapter by Nigel Storkon management implications of canopyresearch, where arguments regarding thesignificance of canopies to conservation ofbiodiversity seem ironic given that—asYves Basset points out in his earlier chap-ter—much of Stork’s research and writingshave focused on countering the view thattropical forests harbour an inordinatediversity of canopy specialists, as com-pared to, say, the impressive biodiversityof forest soils (e.g. Stork, 1988).

The research findings of Stork and otherssuggest to me that the best conservationchoices should be based on informationabout forests taken as a whole. This viewappears contrary to a proposal by theInternational Canopy Network, as it isdescribed by Nalini Nadkarni in TFC, to

identify �canopies of international signifi-cance for conservation�, or CIS, none ofthe criteria for which appear to be neces-sarily canopy-specific. How likely is it thata forest will harbour a �significant� canopyflora and fauna and at the same time befound �insignificant� for ground or soilspecies? It is impossible to conserve thetop portion of a system without conser-ving its bottom. To succeed as conserva-tionists, forest canopy biologists need towork as equals with other forest specialiststo develop programmes that make thisexplicit.

A millennium is a time for reflection inall things, canopy biology among them. K.Eduard Linsenmair notes in his forewordhow canopy biology is �maturing� by�becoming more experimental and predic-tive�, an idea echoed by Andrew Mitchell.Stephen Sutton describes the history ofcanopy science as a shift �away from pureexploration (the �Wonderland� phase) totackling the practicalities of rigorous can-opy research (the �Reality� phase), and theunderlying emphasis is now shifting fromaccess to the upper canopy per se to con-ducting replicative and manipulative sci-ence.’ Nadkarni similarly sees a changefrom early studies that �identify phenomenaand document patterns� to group-basedprojects that �address process orientatedquestions to explain the observed patterns�and on from there to the validation of�predictive models.� Yet, in examiningnearly 2700 forest canopy papers, I havefailed to detect any such trends. Numericaland statistical techniques in canopy bio-logy have always been a reflection of theirtime, and canopy ecology has grown asecology has grown. It is worth remem-bering that some of the most creativepapers on the subject remain some of theearliest works. To take one example,many classic studies of epiphytes containnumerous experiments and predictivemodels, superb for their day and still fullof useful insights—consider the work ofColin S. Pittendrigh, Mason E. Hale, Jr.,Takahide Hosokawa, Dick R. Johansson,and others (e.g. see Barkman, 1958).

Sutton, Nadkarni and others stress theneed for standardized research protocols.For conservation endeavours standardiza-tion may have its uses, but in developinga robust science I think that, if overem-phasized, the idea could be deadening. As

ecology has grown, the available pointsof view on issues and the options forattacking a problem have expanded, notshrunk; the range of possibilities is whatattracts the best minds to a field. Bongershits the nail on the head in his section�Canopy structure: what do you want toknow?� Similarly, to allow for the grow-ing possibilities, pivotal terms (e.g. can-opy and stratification) need to be definedbroadly so they can be adapted (explicitly,and with logic and care) to an increasingvariety of viewpoints and situations(Moffett, 2000; Parker & Brown, 2000).At the same time, successful implemen-tation of massive common data sets,advocated by several authors in TFC, willrequire at the minimum a clear expressionof the subject of enquiry. As long as sci-entists are ambiguous even as to whatstrata they have in mind by the term�canopy� (and, further, as to what theymean by �strata�; Parker & Brown, 2000),attempts to manage common data setsor any other form of synthesis will con-tain serious hidden flaws, and canopybiology, regardless of other successes, willnot reach its much-heralded intellectualmaturity.

Mark W. Moffett

Museum of Vertebrate Zoology,University of California at Berkeley,

USA

REFERENCES

Barkman, J.J. (1958) Phytosociology andecology of cryptogamic epiphytes. VanGorcum, Assen, The Netherlands.

Lowman, M.D. & Nadkarni, N.M., eds(1995) Forest Canopies. Academic Press,New York.

Moffett, M.W. (2000) What’s �up�? Acritical look at the basic terms of canopybiology. Biotropica, 32, 569–596.

Moffett, M.W. (2001) The nature andlimits of canopy biology. Selbyana, 22,155–179.

Parker, G.G. & Brown, M.J. (2000) Forestcanopy stratification – is it useful?American Naturalist, 155, 473–484.

Stork, N.E. (1988) Insect diversity: facts,fiction, and speculation. Biology Journalof the Linnean Society, 35, 321–337.

� 2002 Blackwell Science Ltd, Journal of Biogeography, 29, 1263–1265

Book Reviews 1265