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Dry Weight Of Fresh And Preserved Spiders (Araneida: Labidognatha) PDF

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66 ENTOMOLOGICALNEWS DRY WEIGHT OF FRESH AND PRESERVED SPIDERS (ARANEIDA: LABIDOGNATHA)1 RobertL.Edwards2 WendyL.Gabriel3 , ABSTRACT: Dataonthedryweightfor19taxa(suborderLabidognatha)offreshandpreserved spidersarepresented. Thevariationinweightatlengthforindividualspeciesisalsoprovided. WiththenotableexceptionofthegenusTetragnatha,FamilyTetragnathidae,theFamilyTheridi- idae,andmostofthegeneraoftheFamilyThomisidae,thefamiliesexaminedaresimilartoone another in their weight-length relationships and are not readily separable on that basis. Dry weightcanbe approximatedusing eitherthe weightoffresh specimensorpreserved material. The'typical'spider,basedonthematerialexamined,increasesinlengthfasterrelativelythanit increasesinweight,anddry weightdecreasesrelativetofreshweightaslengthincreases. With the exception ofthe papers by Clausen, 1983, that included dataon fresh and dry weight-length relationships fornine species (five families), and Breymeyer, 1967,forthree speciesofthefamilyLycosidae, there isverylittle general information availableonthedry weightofspiders. Thisreportserves to increasethe informationavailableonspiderweightandexploresthedegree ofdifference betweentaxafrom the weight-length perspective. MATERIALAND METHODS Thebulkofpreservedmaterialwascollectedin 1989and 1990intheFrances CraneWildlifeManagementarea,Hatchville,FalmouthTownship,Barnstable County(CapeCod),Massachusetts,inconnectionwithanotherstudy(Edwards 1993). Collection details are provided therein. All were preserved in 75% denatured ethanol and all had their alcohol replaced at least once, typically within48hoursofcollection. Thetotallengthwasmeasuredfromtheclypeus to the distal end of the abdomen using an ocular micrometer for specimens <12mmintotallengthandverniercalipersforthose>12mm. Thetotallength, as described above, was measured to the nearest 0.1 mm, and the specimens damp dried on absorbent paper before weighing. Obviously distorted speci- menswerenotused. Inthosecaseswhere thepedicelhadelongated,thesepa- ration ofthe thorax from the abdomen was measured and the total measure- mentcorrected accordingly. ThefreshmaterialforthisstudywascollectedinthemonthsofJunethrough September, 1996, from the same area and habitats as the preserved material with one exception. The collection ofLeucauge regnyi Simon, Family Tetra- gnathidae, was made in PuertoRico inSeptember, 1996. Allcollectionswere 1Received April9, 1997. AcceptedJuly7, 1997. 2ResearchAssociate,UnitedStatesNational Museum, Box505,WoodsHole,Ma.02543 3WendyL.Gabriel, NortheastFisheriesCenter, 166WaterSt.,WoodsHole,Ma. 02543 ENT.NEWS 109(1): 66-74,January&February, 1998 Vol. 109,No. 1,January&February, 1998 67 made in the afternoon, the spiders were immobilized in an ethyl acetate col- lecting jar, identified and measured that day, refrigerated overnight at 3 C, and weighed the following day on a MettlerA200 balance, accurate to 1 mg. Followingthis,thematerialwasoven-driedatatemperatureof40 Cforseven days. Tocheckthe efficacy ofthe drying regime, three samplesof50ormore mixedspeciessamplesweredriedforanadditional sevendays,withthe great- estadditional lossofweightobservedofless than 3% in all cases. Specimens ofall species used in this study have been deposited in the United States Na- tional Museum. RESULTS AND DISCUSSION The families, genera, number of individuals weighed, and the range of total lengths are provided inTable 1. Atotal of2,315 measurements offresh, dry fresh, and preserved specimens representing 78 genera and 17 families were made (Table 1). Note in Fig. 1 that the various taxa are identified with the firstfourletters ofthe taxon as listed inTable 1. 0.25 FRESH O DRY FRESH DRY PRESERVED -f DC 0.2- O O o.i5- t o.H Q O O 0.05- n a a a a a + a a a a THOMTMAR LINY LEUC ARAN LYCO SALT PISA PHOL TETR THEROXYO PHIL ANYPAMAUCLUB CORI AGEL GNAP TAXA Figure 1.ConditionfactorsforthetaxalistedinTable 1,usingtheequation k=(aLh)IL3. The taxaareidentifiedusingthefirstfourlettersofeachtaxon. Therearevaluesforeachtreatment (fresh,dry fresh,drypreserved) withtheexceptionofAMAU,TMAR, PHOL,andOXYO,for whichtherewasnodrypreserveddataavailable. 68 ENTOMOLOGICALNEWS Thelogtransformedleastsquaresequation,Inweight=lna+b(lnlength), was used to estimate weight (mg) at length (mm). The statistical parameters are provided in Table 2. The coefficient of determination (r2) ranged from 0.743 (Oxyopidae, dry fresh) to 0.985 (Corinnidae, fresh). Averages, mini- mum, and maximum values are listed forall parameters. The average values for coefficient In a (intercept) varied considerably and were greatest (-1.976) for fresh and least for dry preserved specimens (-3.580), while the average values for exponent b (slope) varied relatively little, from 2.739 for dry preserved to 2.800fordry fresh specimens (Table 2). Thenumberofindividualsandgeneraobtainedforeachfamilyvariedcon- siderably (Table 1). It is clear from the statistical parameters presented in Table 2 that there was little difference in the weight-length relationships be- tween taxa. To compare weight at length using different taxa, while taking intoconsiderationthe differing length rangesoverwhich theparameters were estimated, we calculated thecondition factorforeach taxon,k = (aLb)IL3 (L = mid-point of lengths (mm) in sample, a = Exponent In a) and the results showninFig. 1. Ofthe 19taxathegenusTetragnathaLatreille(Tetragnathidae), the family Theridiidae, and the rotund crab-like members of the family Thomisidae(generaXysticusC.L.Koch,OzyptilaSimon,MisumenaLatreille, Misumenops F.O.P.-Cambridge, and Misumenoides F.O.P.-Cambridge) stand apart from the rest. The genus Tmarus Simon, family Thomisidae, is rela- tively slender (less crab-like and rotund), differing in this respect from the other genera ofthe family Thomisidae listed above and is plotted separately (TMAR). Similarly,thelesselongategeneraLeucaugeWhite,andPachygnatha Sundevall,familyTetragnathidae,differinbodyformfromtheelongatemem- bers ofthe genus Tetragnatha and are also plotted separately (LEUC). The uniqueness ofTetragnatha was notedby Greenstone, et. al. 1985. Breymeyer (1967) in a study concerning the dry weight ofpreserved spi- ders reported that alcohol dissolves and extracts some parts ofspiderbodies. Thisappearstobethecase inthisstudyaswell. Itisworthnotingthatundried specimenspreservedindenaturedalcoholweighconsiderablymorethanfresh material (Edwards 1996). Clausen (1983, p. 143-144) noted that"the ratioof dry over wetweight increases with decreasing size of specimens", and sug- gested that "With decreasing size, the exocuticle may make up a relatively greater part of the animal's weight because of the relatively greater surface. And, there may be a minimum thickness ofthe cuticle, which, in effect, will givethe sameresult." TotestClausen's (op.cit.) suggestionfurther,theentire data set available was examined by treatment, i.e. fresh, dry fresh and dry preserved. The statistical parametersforeach treatmentare provided inTable 3.Thepercentoffreshweightfordryfreshanddrypreservedmaterial isshown in Fig. 2. The results are consistent, forthe average spider, with the observa- tions of Clausen (op. cit.), with the caveat that more than the cuticle is un- Vol. 109,No. 1,January&February, 1998 69 y 0.35 LJJ X CO LJJ DC 0.25- oLJJ 0.2- DC LJJ Q. 0.15 10 15 20 LENGTH (MM) DRY FRESH DRY PRESERVED Figure 2. Percent offresh weight represented by dry fresh weight and dry preserved weight, basedonaveragedataforeachtreatment(Table3). doubtedly involved since drying does not reduce the specimens to the cu- ticle only. Other tissues are involved which may also vary in the degree to which they are present indifferent taxa. Thevariability seenin individual species intheratioofdryweighttofresh weightwasexaminedfor 11 speciescollectedinOctoberandNovember, 1996. The individual speciescollectionswereeachmadewithinanhourinrestricted localities to reduce environmental variabilities as much as possible. The col- lectionswere treatedandanalyzedasdescribedearlier(seeTable 4). Onaver- age the ratio ofdry to fresh weight was 0.314 0.038, varying from 0.250 to 0.390. The average slope (b) ofdry on fresh weight was slightly in excess of 1 with two notable exceptions, that of Pardosa lapidicina (b = 1.224) and Phidippusclarus(b= 1.537). Theaverage intercept(Ina)valuesforthesetwo species were also well in excess of the average value, -2.015 and -3.161 re- spectively. Bothofthese collectionswereofimmature individualsthat would have matured the following year, although the Tmarus angulatus collection, alsoofimmature individualsdue to mature the following year, did not have a similar departure from the average values. The average r2 for the weight- lengthregressionsofthese specieswas0.892forfresh and0.864fordryfresh. 70 ENTOMOLOGICALNEWS For studies requiring precision, e.g. those of a single or a set of closely related species, it would be best to use a sclerotized part ofthe body such as the head capsule, to reduce the problems associated with measurement error (cf. Jocque 1981). Whatevermethod is used, it is obvious that the weightof individual spiders is highly variable. Inonesurvey(Edwards 1993)over 12,000specimenswerecollected. Many ofthese were archived against the future. It was encouraging to find out that preservedmaterialalsoservedthepurposeofrealisticallyestimatingdryweight. Table 1. Freshandpreservedspidersexamined. Numberofindividuals=n,numberofgenera- genera,lengths(mm)includedinsample-range. FamilyTetragnathidaeissubdividedintothe genusTetragnathaandasecondcategoryofrounderbodyforms,includingthegeneraLeucauge and Pachygnatha, listed as Leucauge. The family Thomisidae includes all genera sampled exceptthegenusTmaruswhichislistedseparately. Fresh Fresh,dry Preserved,dry Taxon n genera range n genera range n genera range Agelenidae Vol. 109,No. 1,January&February, 1998 71 Table2. Statistical parametersforspiderweight-lengthequations(Inweightug=Ina+b(In lengthmm),forfresh,dryfreshanddrypreservedmaterial. No.ofindividuals-n,standarderror -SE.,coefficientofdetermination-r%exponentofIna=ExpIna. Freshspecimens Taxon 72 ENTOMOLOGICALNEWS Vol. 109,No. 1,January&February, 1998 73 s g -^ 74 ENTOMOLOGICALNEWS ACKNOWLEDGMENTS TheassistanceofEricandAnnabel Edwardsin the fieldwasgreatlyappreciated.Thecom- ments and suggestions ofJess Zimmermann, University ofPuerto Rico, Los Piedras, and Stan Sady, MassachusettsGeneral Hospital, Boston, whokindlycritiquedthe initial manuscriptwere mosthelpful. Wethankthetwoanonymousreviewersfortheircommentsandsuggestions. Nancy McHugh,NortheastFisheriesCenter,WoodsHole,kindlymadelaboratoryfacilitiesavailable. LITERATURECITED Breymeyer,A.1967.Correlationsbetweendryweightofspidersandtheirlengthandfreshweight. Bull.acad.Pol. Sci. CI-II,XV(5):263-265. Clausen, I.H.S. 1983.Weight-length relationsofeight speciesofspiders(Araneae) fromDen- mark.Ent. Meddr50:139-144.Copenhagen,Denmark 1983. Edgar,W. D. 1971. Seasonal weightchanges, age structure, natalityandmortality inthe wolf spiderPardosalugubrisinCentral Scotland. Oikos22:84-92. Edwards,R.L. 1993.Canthespeciesrichnessofspidersbedetermined?Psyche. 100:(3-4)185- 208. Edwards,R. L. 1996. Estimating live spiderweightusing preservedspecimens.J.Arachnol. 24:161-166. Greenstone,M.H.,C.E.MorganandA.Hultsch. 1985. Ballooningmethodology: equations forestimatingmassesofsticky-trappedspiders.J.Arachnol. 13:225-230. Jocque,R. 1981. Sizeandweightvariationsinspidersandtheirecologicalsignificance.Biol. Jb.Dodonaea. 155-165.

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