VULNERABILITY of BRANCHIOPOD CYSTS to CRUSHING S. A. Hath
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/ JO UR N A L O F CRUST.-\C E.-\N BIOLOGY. 16, 3)' .w ~-~ 5 ~ . 1996 VULNERABILITY OF BRANCHIOPOD CYSTS TO CRUSHING S. A. Hath away, D . P. Sh eehan, and M. A. Simovich ABSTRACT Ep hemer al wet lan ds. suc h as verna l pools and pla yas. have become increas ingl y thr eatened by a vari e ty of de structi ve human practices , includi ng off-high way ve hic le (O HV) ac tiv ity. Amo ng the resident ani mals that may be ad versely affec ted are the bra nchiopod crus taceans whi ch produce des iccatio n resi st ant cys ts to withs tand the pools' dry peri od s. We report a qua ntirative investig at ion of the fo rce require d to crush indi vid ua l bran chi op od cysts of 8 spe c ies: Branchin ecta lyn ch i, B. sandie gonen sis. B. ma ckini, St repto cepha lus woouon i, Thamno cephalus platvurus, Linderiella occidentalis, Lep idurus lemmoni, and Tr iops lo ng icaudata . Ver y s ma ll for ces o f less than! new to n crus hed dry cysts. Wet cysts we re even mo re fragile; many were crushed under less tha n 7 x 10-' newtons . Differences in vulnerabi lity were found within and be tween taxa and may be related to cyst morphol ogy. Kn owled ge o f these e ffec ts sho u ld be taken into conside ratio n wh en de ve loping management s trategies for ephe m eral wetlands. Ephem eral w e tla nds , s uc h as vernal found si gnifi cantly more broken cys ts of pools, playas , and dr y lak es, are deli cate Branchinecta mackini De xter and inc reased systems supporting a rich flora and fauna. breakage of cysts of Thamnocephalus p la Man y of the se wetlands are also important tyurus Packar d and Triops longi cau datus to migratory birds and some act as bre eding LeConte in the vehicle tra cks relative to the sites for amphibians. Of the numerou s or undisturbed area. ganisms that inhabit ephemeral pools, prob In our study, we qu antitativel y measured ab ly the mo st ob viou s are bran chiopod the forces required to cru sh branchiopod cru stacean s, including anostrac an s and no cy sts. We also co mpared cyst vulnerability tostracans . These shrimp hatch from desic when wet ver sus dry, and tested whether catio n-tolerant cysts (encysted e mbryos) as there were di fference s in the force s required pools fill from rain or ice melt. If co nditions to crush cysts of different spec ies. permit, the y qu ickly mature, mate , and pro du ce cysts before the pool dries. The cy sts MATERIALS Ai'iD M ETH ODS remain do rm ant in the so il and ha tch in the We determined the force required to crus h cysts of sa me or subseq uent seasons w hen co ndi 6 spe c ies o f an ostracans. Branchinecta lyn chi E ng et tions are again fa vorable. Be cause ephem al., 1990. B . sandiego ne nsis Fugate. 199 3, B . m a ckini Dexter, 1956. St repto cep halu s wOOII!)n ; En g et al., eral we tlands are sea so nal, the y are often 19 90 . Thamno ce phalus p latyurus Pack ard, 1879 . Lin overlooked, and ar e becoming increasingly deriella oc cident alis Dobbs, 1923, and 2 species of rar e du e to de structi on from urbani zation, norostr acans. Lepidurus lem m on; Holmes. J894 , a nd agriculture, recreational ac tivities, and other Triops lon g icaudata L inder, t 95 2, from a population now bei ng described as a new s pec ies o f Tr iops (Sas ca uses (Eng et al ., 1990). sa man. personal co mmunicatio n). Cysts were co llec ted One particular threat to the remaining fro m dri ed soi l an d intac t cysts we re used d ry or wet natural po ols, playas, an d their inhabitants (af ter ap proxim ate ly 6 h of so aki ng in deio nized wa is disturbance fro m off-h ig hwa y vehicles ter). All cysts test ed we re spherical except fo r the 3 species of Branch inecta which had 2 shapes wh en dry. (OHVs). As OHVs pass o ver so il, the either co ncave or s pherica l. Bo th sha pe s are via ble and weight of the vehi cle co mpresses the so il. both we re tested . We tested co ncave cysts to determine Th e largest amount of soil compaction which o rienta tion. sphe ric al si de up or down. was mo re takes place during the first few passes o f durable. We use d the mor e durable or ientatio n. sp he r OHVs (Iverson et aI. , 1981), and most com ical sid e up , to cal culate crushing force. To de term ine the force required 10 c rush an ind ivid paction occu rs just be low the su rfac e ual cys t, a cys t an d a piece o f tun gsten wir e of co m (Webb , 1983) Not only is pool hyd rology par able diamete r were pl aced on a m icroscope s lide disrupted by th is act ivity, it is also poten equ ally distant from the edge o f the sl ide (Fi g . I). An tially very damaging to cysts, most of ot her m icr oscop e sli de was the n ge nt ly res ted o n the wire and the cyst , usi ng fo rce ps . Weight was incre wh ich are found in the top few cent ime ters me ntally add ed (us ing for ceps) to the s lide di rectly of so il. Eri kse n et al. (1986) simulated above the cyst unt il it c rus hed. Was hers were used as OHV traffic through a de sert play a and we ights. all owing us to observe cr ushing thr ough the HATHAWA Y IT AL. VULNERABI LITY OF £lRANCHIO PO D CYSTS 44 9 = w e ig h ts RES ULTS Very small forces crushed cysts of all w:JI----=~ species . and there were significant diffe r ences between hydrat ion conditio ns and L --- +-I ~ b -- I !-a- I' I among species (F = 85.8. P < 0.000 I) (Figs. 2, 3). When dry, the maximum forc e Fig. I. Diagram of apparatus used to det ermine for ce required to crush even the most resistant required to crus h bran chiopod cysts. dry branchiopod cys ts was less than I ne w ton. This is less than the we ight of a 100 g mass. The force was sig nificantly less to center hole . If the cy st c rus hed immed iatel y upon set . ting the microscope slide or weight s on the cyst. that crus h wet cy sts of all species (F = 80.7, P test sam ple was thrown out. In case s where more tha n < 0.000 I ) (Figs. 2, 3) . In fac t, 100% of 3 con secutive cysts crushed under the mi cr oscope slide cysts of L. occ ide ntalis, 70% of B . sand ie alo ne. the force of the microscope slide wa s used as a g onensis, B. lynchi, and S. wo otton i, and co nser vative estimate of cru shing fo rce . The crushing force (F,,,,,,) for each cys t is: 30 % of B. mackini, crushed unde r the fo rce of the microscope slide alo ne or less tha n 7 F<rush = m,g + W- I( U2 - b)/(L - a - b»)m.g X I O- ~ N when wet. Thamnoceph alus where rn, = mass of the mi croscope slide ~ . 8 0 X [0 -) platyurus wa s significantly more resi stan t to kg; g = gra vitational acceleration 9.78 m/s-; W = crushing than all othe r anos tracans except weight in newtons (N ) = 1.00 kg.m/s' : L = length of B. mackini (F == 2.3, P < 0.05 ) when wet. the slide 7.60 X 10- 2 rn; a = dis tance from the wire 10 the edge of the slide 5.00 X 10-) rn; and b = di s The remaining species did not differ. W he n tance from the c yst to the edge of the slide 5.00 x wet, notostracans require d signi fic an tly 10-) rn. more fo rce to cru sh than anostracan cysts St atistic al differences wer e dete rmi ned using S ru (r-test == 11.3, P < 0.000l). den t's r-test or anal ysis of vari ance ( P < 0.05). Where significant difference was fou nd; multiple co mparisons When dry, notostracans again required among mea ns wer e made usi ng Sc heffe F test, and the signi ficantly more force than anostracans (r least signi ficant values are presented (O n . 1977). test == 20.3 , P < 0.000 I). Within the gen us 0.5 0.4 z 0.3 r.'71 l::.:J eo T au ost rac an s > 0.2 < T [2] n ot os tracaos 0.1 T u; 0 B.