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, . awx,.Jt·s · .~~:I~r:~t~ ..; .~••.... ~~.~i.onofthe _r~~ h . .



--.-.excess manganese

LEnERSTONATURE·--associated to the median
""."", ;"."j,,'i,

vall,~.\· ;md does not spill over into the adjacent '~!dlOugn Il may leak out through fracture

. By~aki~g a con~ervative approach and grouping those plumes tn whlr'J ~~~ "'!?""'!? ':'cc~:- ~~ ~h;:;::;~::-;;;~;:;;:~;, ~.;;: ::;;;~.~~~ •.•••• be identified, that is stations 2, 4 and 5; stations 6 and 7; station 8; station 9; and stations 10 and 11. This analysis predicts a frequency of one source per 340 km of ridge; of course the arhlal frequency may be much highel The position ofventing in the rift appears to vary with latitude. Results from stations 8 and 9 are consistent with venting from the east wall, whereas results from slation 10 indicate a west-wall source. Furthermore, these results are consistent with discharge from the valley walls, but they do not eliminate lhe possibility that venting occurs from the valley f1oor. Finally, hydrothermal manganese along this section appears to be confined
Rcccived 21 November 1984: acc;epted 21 Fehruary 1985.

zones. J'in:: anomaues n:'.>(I.-l<:O nere are smatler than those found near venting in th(' P ••dfic4•7• However, this difference is an artefact of the regi:: n.'l' .,ampling strategy of this survey and is not a true indication 01' the relative impact of venting from the MAR on geochemical cycles an,: the heat budget. Such Information can be acquired on!y by site-specific work. We thank Caplaín R. L. Swanson, officers and crew of the NOAA ship Researcher for their suppori and T. A. Nelsen of NOAA and J. Trefry 01' Florida Institute of Technology for their cooperation. This work was supported by the NOAA Vents programme and the NERC; Cambridge University Earth Scíences Series, contribution' 578.
13, Burton, J. D., Maher, W. A. & Stalham. P. J. in nau Metais in Seawater (eds Wong. C. S.• Doyle, E.• Druland, 11: W., kurtun, J. D. & Goldberg, E. D.) 415·426 (Plenum, New York, 1983). 14. Froelich. P. N. tt 01. Geofhim. 'I.\mochim. Arta 43, 1075-1090 (1979). 15. Discholl, J. L. & [)icks~", I,. W. hJrlh plan.,. Sei. Lell. 2~, .185·.197 (197ll.
16. Edmond,

I. Rona, P. A. J. Volcano/. geolherm. Ret. 3, 219·225 11978). 2. Rona, P. A. ti 01 Geo/. Soe. Am. AbsIr. Progm 14 (7),602 (1982). 3. Rona. P. A. Earrh Scí. R.v.20, 1·104 (1984). 4. Klinkhammer. a., Dender, M. & Wei•• , R. F. Na/ur< 269, 319·320 5. Klinkhammer, G. P. Chem. Geo/. 29,211·226 (1980).


6. Luplun, J. E. el g/. E"grrh planei Sd. Leu. 50,115-127 (1980). ,. Jlllltf'l:. C. J., Jnhm.on. fi. P &. Dclaney. J. H. Gmpltv\. Rn, I •• I. M71. k7t. {JlJIiH. " h KhulluHlIluer. (;" l:IJcrludd. 11. &. I'ud"'\lll, A NlJIIJ'i' lOl\, 1'K"i Itt~ (Ilnn,. 9. Klinkhammcr, G. Anuly', (,hem, 52, 117-120 (ltJKO'. 10. KJinkhammer,G. P. & Dender, M. L Earrh planeI. Só. LeI/.46, 361·384 (1980). 11. Dender, M., Klinkhammer. G. & Spencer, P. lJ"p·S.a Re.•.24, 799·812 (1977). 12. Lamber!, C. E., !Iishop, J. K. O.• Discaye. P. E. & Chesselel. R. Earlh plan.,. Scí. LeI/.70, 2)7·248 119841.

J. M.,



1\, 1...


R. E. &:



I. Ncuurr



17. S •.uu, M. R .• ~;"Iln, R. a., !~(l1l,J, P /\., Hutler, I.. W." Nalwiflllr., A. J. Of'oph.vJ. ReJ. Lru, 1,355·358 (1974). 18. Shearme. S., Cranan. D S. &. Ron., P. A. Mar. Geol. ~I. 269·291 (198)). 19. Rona, P. A. Geoph.vs. Re •. Lell. ~, 993·9Qh 1\978). 20. Jenkins, W. J., Rona, P. A. & Edmond, J. M. Earrh planei. Scí. Lell. 49, 39·44 (1980). 21. Rona. P. A. J. geol Soc, i "nd. 1.17,385·402 (1980).

Fractal dimension or vegetation and the distribution or arthropod body lengths
D. R. Morse*, J. H. Lawton*, M. M. Dodsont & M. H. Willialhson*
Departments of *Biology and tMathematics,
Y..,t'k Y0i 3i:Ji:J) Li,,"

University of York,

transects across a C;-i ai r~:.f~. An alternative widely used method (see ref. 1, p. 130), which has some practical advantages, is as follows. Enclose the ~bject (or a subsection of it) in a square of si de S and divide the square inlo (SI A)2 squares of side À. Let N(A) be the number 01' sCl;;an;,· '.he edge of the object enters and plot log N(A) against lo!,! 1,1 A. Suppose that for small values of A, the graph is almost linear with slope D. Then D can be inter;.. ,~~~.j a.s th~~ f,.;·tc:utl dlrnens10n, .• Slnce





Following Mandelbrotl, recent studies~ demonstrate that some nalural surfaccs are fractal. Hcre wc show that transccts across vel?ctation are fractal, and considcr 00(' J}ossible conscqllcncc of this ohscrvation for arthropods (mainly .nsects) Iiving on plant surfaccs. An important future of a fractal curve or surface is that its lenglh or arca, rcspectively, becomes disproportionately large as the unit of measurement l!'l decreascd I. This suggests that if vegetation has a fractal structure, thcre is more usable space for smaller animais Iiving on vegetation than for fllrger animais. Hence, there should be more individuais with a smalI body length Ihan a large body length. We show that this is the case, and that relative numbers of small and large individual arthropods collected from vegetation are broadly conslstent with theoretical predictions originating from the fractal nature of vegetation7 and individual rates of resource utilizatlon. . Mandelbrotl,4 asks, how long is the coast 01' Britain? Measuring its lenglh by stepping round with successively shorter divider slep-lenglhs allows finer and finer resclulion af lhe coas'tline; if the log of divider step-length is plotted against the log of total length measured, a straight line ofnegative slope is obtainedl,4,8. Mandelbroll•4 inlerprets this as evidence lhat coastlines are fraclal and lhat lhe slope of the graph is an estimale of 1- D, where [) is lhe fraclal dimension 01' lhe coastline, The relationship between D, step-Iength r and lhe perceived length of the line I( r) measured with step r is given by




By definition , for a line (that is, a transect across a surface), D must lie in the range 1 ~ lJ ~ 2 and for a surface, 2 ~ D ~ 3. The fractal dimension of vegetation might also be measured hy stepping along a transect over the slIrface wilh a pair 01' dívidas sei to a series 01' step-lenglhs and lIsing cqualion (I). Th:s ':..~~ ~:I'"It~(.thvJ u::H.,J lU liit.:ê:t;,lI(C; ~i1c í"l dt,;ta; dlm~n"lnl) ""

The method is explained in gr.: ••ter detail in Fig. I legend, together with an example. Photographs were tilken of a variety of plants during early spring. The plants were eilher leat1ess, JUSl coming into leaf or evergreen. By careful choice or plant-parts and camera focal length, the resulting photographs were approximately two· dimensional, greatly simplifying their interpretation under the grid ( Two ditlerent arbitrarily sized .• grids (128 mm or 256 mm along one ed;;e) were partitioned into 2n squares along each edge. Depending on the grid size, n varied from 2 to 6 or 7, down to. a square size of 2 mm. The grid was randomly repositioned several tim,~s to give semi-independent estimates 01' the fractal dimer.,ion 01' the same piece 01' vegetation. The slope of the resulting !il1c (for cxample, rig. 1h) was estimated by legres~ion unulysi". There are several points to note in the interpretation of such graphs. One is that when the number 01' divisions is small, it is prohable that ali lht> sqllar.:s will he entered, hence the ·slop. 01' the graph will be n == 2. This is an artefacl of the melhod; consequently, in the prescnt study ali points that fell on fhe Iille y = 2x (on a log/log pIo:) were omitted when estimating the slope 01' the line. At tht: other extreme of resolution, when a very large number 01' sqllares is used, the slope of the graph will in general fali to I. This could be due 10 a lack of resolution in the photograph or pccallse irregularities in the outline of the plant no longer OCCI.'r <il '!Iat scale, Taking a photograph at a much larger scale (higher ma~nification) resolves this prohlem. Thc graph may he convcr.'··1 hClween the Clllrcml:s 01' [> == 2 1I11d [) - I. or it could h,·\'c lwo straight subre/l.ions characterized ")' di!Jere!ll ~;Iopes (hg. 1 iJ), Similar changt:s in fractal




5+ I.68 1. then parts of the samc twigs were rephotographed at a higher magnification. for example. and Zucc. Hedera helix L. Acer pseudoplatanus L. The mean fractal dimension of the samp:es iu Table 1 is· tended to foU:. N.40 1...1~ scales approxiw"'di dS lÍle 0. Taxus baccala L. For ease of calculation. photographed without leaves in early spring.035 0.40 1. Those plants which one might. then 2" squares. Fraxinus excelsior L. ali other things being equal (especial1y the rate of appearance of new resources).3.'.High-magnificationestimateswere derivl'<1rnrn "'n~•• t:t' p~~~~êr::t:h: f :.cctliun estimat.099 0. 0.44. ref. (twigs and leaves) Magnification High Low High Low Low High Low Low High Low Medium Medium Medium Medium No. see rer.39 1.023 3 6 18 3 3 3 3 3 3 Photographs or branehes and twigs of a selection of woody plants.l6-fold increase in surface area for an order-of-magnitude decrease in ruler length. in reality they are orientated at ali anglts with respeet to the grid. the progressivelyfiner divisions are only ilIustrated in one comer of the figure.) Rehd. This lower bound predicts a 3. then for a homogeneous fractal surface having transects with D = 1. ref. W. (twigs and buds) Cotoneaster. In those plants which were photographed at two scales.040 0. This increase in available space for animais of smaller body length may be combined with a consideration of the way Jn which metabolic rate scales with body length 13. estimates of the fraetal dimension are lower at the higher magnific~tion. 365) gives a heuristic upper bound estimate for the expected increase in surface area.75 power of body weight. :":".(evergreen) Virginiaereeper. for the same reference area. IvW-11l3l!'lif.28. outlined above. as in b.11. and in particular when the surface is flat in a direetion transverse to the cross-section( which is dearly not the case for vegetation).79 1. I.that is. 1. Table 1 presents data on the fractal dimension of a selection of pIants. For an order-of-magnitude decrease in body length. (twigs and feaves) Ash. depending on the grid size. 13)..a given decrease in body length.were from pnotographs oftwigs> 50em long. However. typical1y. Berberis vulgaris L.75 = 178-fold increase in the density of individuais. As a first approximation.5.14 to make predictions about the distribution of body lengths of animais living on vegetation. for a close-up photograph of Virginia creeper. Hence.75. permitting O to be estimated at two levclsof resolution.: ••lher ·princip•• axes' in the natural structure of the vegetation.16.5. Single medium estimates were made from speeimens >25 cm long. D. when D= 1. 365). (twigs anJ !eaves) Sycamore. scales as (e)-0.43 1. substitute animai body length (L) for step-Iength (A) in equation (1).a.. Ifthe way in which animais perceive and use their environment is proportional to their body length12. For ease of representation. 1. D = 1. starting with a course grid of two large squares on one side.t·I~.018 0.019 0. Bearing in mind the disconnected character of the surface of vegetation. b. see.) . Estimates of D range from 1.42 1. Then. a 10-fold decrease in body length results in a (103)°. assume that. Next. of squares 00 one side of grid flg.75. as (L'. The metabolic rate of individual animais 13. Also for c1arity. with n varying from 2 to 6 or 7. This could introduce some bias into lhe estimates of i their fraetal dimension. the area perceived by animais 3 mm long may be up to an order of magnitude greater than the area perceived by animais 30 mm long. Parlhenocissus Iricuspidala (Sieb.093 0. Ulmus glabra forma pendula (Loud.009 0.28 1. for. Now consider the implications of the fractal nature of plant surfaces for the animais living on them. with or without leaves. lhe speeies name. (Note lhat ~"mr!e tran!ects !~e·::t. squaring the increase in linear distance (that is.078 0. the plant's leaves in this figure are drawn fiat. such calcula- Species Barberry. The number of squares entered by the outline of the plant were counted. p. were taken from the University of York eampus or SkipwithCommon.5. The difficulties of measuring the fractal dimension of a surface are considerable and no data are yet available. (evergreen) Silverbireh. This holds exactly under some circumstances.):1ional tothe rcdprucal of individual rates of resource utilization (that is.5) across a fraetal surfaee increases hy a factor of jlo = 3. to 1.ftwigs":O em loog. It fol1ows from equation (1) that for an order-of-magnitude decrease in uler ienglh.042 0.47 1. (he expected distance between two points on a linear transect (of dimension D = 1.S5 1.050 0. Data gathered in this way for Virginia creeper. if use of resources per individual is proportional to WO. Belula pendula Roth (lwigs and (eaves) Downy birch. (evergreen) Yew. The state of each plant when photographed and the plant part photographed is indicated in p::!renthesesarte. p.) Planch. 1 a.300 No.31 s.of estimates 3 3 6 3 6 Mean D 1. to prediet reiative numbers of individual animais of different body lengths Iiving on the surface of vegetation. dimension at different scales have been found in other natural objects 1. it fol1ows that population density. (twigs and buds) Weepingelm. Relula pubescens Ehrh. North Yorkshire. adding the fractal dimensions of the orthogonal transects.083 0.I 1001 Jb'155 loi j(1cw magnlflcallon) lJ-:--ii 100 1. a priori. an estimate for the lower bound ofthe fractal dimension is obtained by adding I to the linear fructal dimension (for example.0 for an order-of-magnitude decrease in ruler length. as would be expeeted from the above argument. The twigswere photographed at one scale.d.162 = 10. Therefore. consider to have a more complex growth form have a higher fractal dimension. The slope of the line equals the fractal dimension. Coloneasler horizonJalis De~aisne (twigs and leaves) Ivy.41 1.6" . suppose that population densities are approximately prop.79 for cotoneaslet.042 0.'.111 0. The logarithm of the number of squares entered by the outline of the plant was then plotted against the logarithm of the number of squares along one side of the grid.. the maximum expected increase in surface area is 3.46 1. to metabolic rate-I. This increase in density may be combined with the expeeted increase in the available surface area.35 1. Photographs of plants at various magnifications were placed under a grid.

197Kl U. in general agreement with OUI" predictions (Fig. 1984). 19)13). J. 1.295-296 7.) and Kansas secondary vegetation (O). • \ 0'1. H. Theoretical and emoirical Ii"k~ hl"h''':'P''l ?:·.. & Grren. o o oo \ I.". Str. E. 111(" "'''''aNaf G(!(·tn("ry lJf Na.\ 1'. We know of no oth'~r aitempts to explain patterns in the body size distributions 0(' .mas (eds Mound. \ 733 \ \ \ \ \ \ \ \ \ \ \ I. 121. understorey foliage in cacao plantations in Dominica (. in lhe pFess). e. I: '-. \ \ \ \ 10 I.1I 11" p"·••• . M. c\. Consequently. 6. \ \ \ 100 Body length. Birch ( Betula fluhptN'n<. b. & Schoener. W ll. although previoll'l S.. d.) and 700 (O).ic~/rre4lJl:llcy J ôistributions for animaIs of different sizesl2. in a-d lhe I mm body lenglh dala point and in e the firsl three dala points were omitted. fi. a. & MacArthur.<.'[ic""o"J olBod.t Matltttlh •. Mandelbrot provided helpful criticism of the manuscript.. Rigllul.. (1979).. t:t(J-I~" (1'161).18. \ \ \ I. H. O. I" I". . D. d e 1..• I. r~'cl.l'lalilll1 111' f) -.JH (l"011.. \ \ ~ \ \ i . E.. h. \)'_". R. R. Sí"lt"tll'f.\lem.I N"h"~ . K. C1early. O. New York. &. . ~! Sld~w!t!l ~ Common. refs 13. P. (1984). 1.. b. H. \ \ a o • b \ \ \ \ c \ . Rkhilrds(lO. Moreover. 2. allowing prediction of the expected fractal dimension of the surface of vegetation. London.. 96-110 (1968).•nimal Siu . gratefully ack. Brrviora 454. 655-668 (1983). 21.llowledges the financial support of the NERC. T. t. . Õ 10. \ \ \ \ I.. lhere should be a 560-fold increase in lhe number of individuaIs.) and 530 (O).79. Orford. AlO. 6. and by the biology of associated arthropods.."lrlhl'II." I'a. (il''' . "by. T.16 = 560-fold and 178 x 10= 1. ~.780-fold increase for an order-of-magnitude decrease in body length-is shown by the upper dashed line. approximate agreement betweell data and our predictions does not prove that the fractal nature of vegetation contributes to a steep increase in number of individuaIs as anhropods get smaller.:.}:l':vt\. f'. I. Janzen. 93.I (Edward "'"g. Tabogo primary riparian vegetation (. . 2). \ 1U~ \ \ \ \ 'õ . J. Ihnh)ughs. 12. 16.v af 1o. ~1.•. 5. c. Sca/. O. In a-d. E.800 (.000 00 \ \ \ 100 \ \ \ \ .. n. arthropods were collected bv sweep nel. o. . Ó\b r. 9. B. Finca Tabogo. N.000 z d \ \ \ \ I.01'11.440. 14. Eco/"KY 54. & Whalley. 1__ ---------- . 17.. D. dilferent values for IJ (1..) and Icacos vegetation (0)'6. I. \Vh~' . Regression lines were fitted through the log-transrormed data.' !r•• . e. Rri<hrll. not least of which is the difficulty in sampling very small arthropods. The above heuristic calculations may be reversed. H.. I'feifu. in [)ivtrsi.~~er~~ ir. I. lawlon. D.v Sizt (Cambridge Universily Press..~ .711. R. \ \ . \ -. Understorey foliage in primary forest.Osa secondary vegetation' (. 117-125 (\959). }. D. '. .:'''ft 1 m. & Southwood.'''/''K. 390.. I. ·lI~. Miatn G 10.1. lhe surface 01' vegetatiôn are scaíCC. G. c. Junirer. Ali the data show a steep increase in the number of individuais for an order-ofmagnitude decrease in body length. Eco/. N') 188· 204 i1l1ackwrll.jiln~. Arnold. 1 •. O.: 10 .o. • \ ~ \ \ \ \ \ . 2 Data on the number of individual arthropods (mainly insects) of different body lengths collected from vegetation. W. North and 980 (O). D.62-0.·1~6.. -_..V 30. • . li remains to oe seen whether detailed differences in the fitted slopes apparent in Fig.dim'.. is indicated by the lower dashed line on each graph.R. Costa Rical5. 1'.040 (.1 IIQH· •• 1.fO Imp()"unl? (Cambridsc ' Univcrsil)' Pre •• .tJJf. Figure 2 shows such data for sweep-net samples of terrestria! arthrüpúJ" 1<_17 alló originai data collected by pyrethrum knockdown of a tree (Refuta pubescens) canopy.11. i \ \ \ \ • o \ \ \ \ \ \ o • \ \ \ \ o \ \ \ \ \ \ \ \ I. 7111-2". R.LETTERSTOATURE--·· N Fia. J.mhropods Iiving on plant surfaces. 1>.780·fold in the number of individual animais 7.l· 49.IUIl&l\.. R. \ o. in Piam Surfw"" (rrl. 2. H. MiI'.··!"2 (~gl). in e by pyrethrum knock-down of the tree canopy..14) lead to dilferent predicted slopes in Fig. given the mean ofthe fitted slopes for the distribution of individuaIs' body lengths shown in Fig. \ \ \ .. H. Na'. r\h. 1. Mar. I. HUlchinson. I. u . This calculation results in a prediction of the fractal dimension of the surface rlf vl'l'.". 2. .\.).o". the anti-Iogarithm ofthe slopes of the Iines were taken and are as follows: a."íi. 14. \ 10 \ \ .. Schmidl-Nielsen.. E. ')ltUmSilflncr. 1983). ~I 61 (l9MI).:E(:u. R. 2. L (rT'r. J. A. o \ \ \ \ \ \ \ \ \ \ I . h2~'e examii ••. Data on the number of individual animais of different body !e:-:s!!':~ ~. Andrews. ~ " . 2 can be explained by differences in the fractal dimension of particular plant surfaces.86.!I I!~ ll~N~I. \ I.).çlvf "pecies and number of individuab n:' different body lengths constitute an important unsolved l:'w'. B. the upper bound prediclion-a 1.0 \ ~ ~ ~ \ \ . .••• lI. Avnir. I. 360 (.281. /'eto'''R.ullrr ~O". 11. !!':: . when filtíng regression lines to the dala.M. l-SI.urt> (Freeman. 1). 15.tUN. Farin.~.J r. d. K. M.. Colleagues at York and ?rofessor B.) and at Finca La Lola. I'. Jan. Tahle I) and the metabolic rate exponent (0. Ilradhury. 18. & Wt:lbd. I.t Walolf. The lower bound prediction that. 659-686 11973). :\ . A.. as will the precise form oflhe relationship between the fractal dimension 01' à surface anel transeClS across that surface. 00 \ \ \ \ \ \ \ I. NUIII'~ 2'1. • o \ \ \ I. 1. for an order-ofmagnilude decrease in body lenglh.õ) I \ I I tions predict an increase of between 178 x .en. The decline in the numbers or individuais of < I mm body length may be due to a variety of factors". I. Costa Rica (O)". \ \ u .(:uiIm. Mandelbrot.