Gulf and Caribbean Research Vlume 15 ssue 1 January 23 Ppulatin Bilgy f the Ghst Shrimp Sergi trilbata (Biffar 197) (Crustacea: Decapda: Thalassinidea) J.L. Crsetti University f Tampa K.M. Strasser Ferris State University DO: 1.18785/gcr.151.3 Fllw this and additinal wrks at: http://aquila.usm.edu/gcr Part f the Marine Bilgy Cmmns Recmmended Citatin Crsetti, J. and K. Strasser. 23. Ppulatin Bilgy f the Ghst Shrimp Sergi trilbata (Biffar 197) (Crustacea: Decapda: Thalassinidea). Gulf and Caribbean Research 15 (1): 13-19. Retrieved frm http://aquila.usm.edu/gcr/vl15/iss1/3 This Article is brught t yu fr free and pen access by The Aquila Digital Cmmunity. t has been accepted fr inclusin in Gulf and Caribbean Research by an authrized editr f The Aquila Digital Cmmunity. Fr mre infrmatin, please cntact Jshua.Crmwell@usm.edu.
Gulf and Caribbean Research Vl. 15, 13-19,23 Manuscript received February 25,22; accepted January 2,23 POPULATON BOLOGY OF THE GHOST SHRMP SERGO TRLOBATA (BFEAR 197) (CRUSTACEA: DECAPODA: THALASSNDEA) J.L. Crsettil and K. M. Strasser2* ldepartment f Bilgy, (Jniversity f Tampa, 41 W. Kennedy Blvd. Tampa, FL 3366 USA, E-mail j die 97 @ msn.cm 2Department f Bilgical Sciences, Ferris State University, 82 Campus Drive, ASC 24, Big Rapids, M 4937 USA, E-mail Karen_Strasser@ferris.edu ABSTRACT Sergi trilbata is a cmmn burrwing crustacean fund in Tampa Bay, Lemn Bay, and Miami, Flrida, where it inhabits mainly intertidal sft sediments (Biffar 1971, Manning and Lemaitre 1993). Althugh S. trilbata is a dminant member f the benthic cmmunity, very little is knwn abut ppulatin dynamics and reprductin f these thalassinideans. The ppulatin bilgy f this ghst shrimp was examined ver a perid f a year and a half t gain understanding f its life histry. Seasnal variatin in the prprtin f individuals in each size class was bserved during the study, suggesting that there may be envirnmental factrs affecting the ghst shrimp. Additinally, a life span f apprximately tw years is indicated by the seasnal variatin in the prprtin f individuals in each size class. The ppulatin was biased tward females with a 1:1.81 male: female rati, and the mean ttal length f females (68.2 mm) was larger than the males cllected (54.6 mm) (P =.1). Cllectin f vigerus females were directly crrelated with an increasing vary width f females during the preceding mnths, and the ttal length f S. trilbata was psitively crrelated with the number f eggs prduced. Mean number f eggs per female S. trilbatd was 197 with a maximum f 412 ec. NrnucrrN regins f high rganic cntent and cnstruct a thicker burrw wall than d ther species f thalassinidean shrimp in Suth Flrida (Biffar l97). Envirnmental cnditins such as temperature and salinity have been recgnized t impact intertidal decapd crustaceans (Tamaki et al. 1997, Berkenbusch and Rwden 1999). Lw temperatures may cause shrimp t be relatively inactive, resulting in a decrease in sediment turnver rates, decreasing the impact f these Ghst shrimp are decapd crustaceans that cnstruct and inhabit burrws predminantly in shallw intertidal regins cntaining a sft sedimentbase (Felder and Manning 1997, Kevrekidis et al. 1997, Berkenbusch and Rwden 1999). These thalassinideans als cmmnly inhabit regins with a high cntent f rganic matter (Felder and Manning 1997) which has been knwn t trigger settlement in sme species f ghst shrimp (Strasser and Felder 1999). Ghst shrimp are frequently fund in bth trpical and temperate regins rganisms n their envirnment (Berkenbusch and Rwden 1999). Salinity may als be a factr in the distributin f this species, since S. trilbara is usually fund in mid t high salinities and is then replaced by Lepidphthalmus luisianensis in areas f lw r mre fluctuating salinities (Felder l978,felder and Rdrigues (Berkenbusch and Rwden 1999), and are an imprtant cmpnent f the ecsystem due t their biturbating activities, which impact nutrient cycling (Nates and Felder 1998), sediment turnver rates (Felder and Griffis 1994), benthic cmmunity structure (Dittmann 1996) and cmmercial industries such as aquaculture (Davey et al. 199, Lemaitre and Rdrigues 1991, Lafferty and Kuris 1996, Nates and Felder 1998, Berkenbusch and Rwden 2, Felder 21). Sergi trilbata (Biffar 197) is a cmmn burrwing crustacean in Tampa Bay that inhabits mainly shallw intertidal sft sediment habitats (Biffar l97l) and dminates the benthic cmmunity in mid t lwer bay regins. S. trilbatawas described frm Tampa Bay but has als been reprted frm Miami and Lemn Bay, 1993) and higher sedimentary rganic cntent (unpub- lished data). Members f the genus Sergi are cnsidered t be diverse eclgically and may vary with respect t reprductive and life histry characteristics such as develpment and dispersal activities (Nates et al. 1997). t is knwn that varied symbites such as cpepds, pea crabs, caridean shrimp, clams, and gbiid shimp are cmmnly fund in the burrws f ghst shrimp (Jacksn 1996). Althugh Sergi trilbata is a dminant member f the benthic cmmunity, very little is currently knwn abut this species. The present study undertakes bservatins f ppulatin dynamics and reprductive characteristics in this prly knwn but dminant member f the Tampa Bay benthic cmmu- Flrida (Biffar 197, Manning and Lemaitre 1993). Cllectins f this species have been made in Tampa Bay at a water depth f apprximately 2-4 meters (Biffar l97l). S. trilbara is als thught t thrive in nity. t3
CRssrrr an SrnnsssR MlrBnrlLS AND MBrnns classes f 6-89 mm and included nly a very small prprtin f individuals in the smaller size categries f 2-39 mm (Figure 1A). Samples in summer f 2 exhibited a larger prprtin f the ppulatin in the smaller size categries, but mst individuals were still in the 6-19 mm size range (Figure 1B). During the fall, n rganisms were present in the iargest size class (889 mm). Als in this perid, the 6-69 mm size class cntained the highest prprtin f the ppulatin, Mnthly cllectins f Sergi trilbata were made n an intertidal sand flat lcated n the suthwest end f the Gandy Bridge in Tampa Bay, Flrida (27.87N, 82.6W) frm April 2 t August 21. Specimens were btained by extracting animals frm randmly chsen burrw hles in a sub-area f the intertidal sand flat using yabby pumps t extract samples, which were then washed ver 1-mm mesh sieves (Felder 1978). At which may represent individuals recruited int the ppulatin during the previus spring (Fi-uure 1C). n winter 2 and spring 21, there were n individuals present in the smallest size class, and the hi-shest pl'prtin f the ppulatin was again in the 6-19 mm size classes (Figures 1D and 1E). A bimdal distributin is present in bth f these seasns, which ma1' a,eain represent the presence f tw distinct generatins. As in the prer-ius year, in summer 21 the hi-ehest prprtin f individuals was bserved in the 7-79 mm size categry. and there is an increase in the prprtin f lar-eer individuals in the 8-89 mm size cate-qrv tfi-eure 1F1. Althugh cllectins in the fall f 21 u'ere limited t September and Octber, nce a-eain. n r-eanisms in the largest size class f 8-89 mm were represented in the least 15 animals were cllected during each cllectin. Each individual rganism was placed in a clean whirl pack@ plastic bag and returned t the labratry fr examinatin. The prprtin f individuals in each size class (1 mm increments) was examined n a seasnal basis, with three mnths being included in each seasn: spring (March, April, Muy), summer (June, July, August), fall (September, Octber, Nvember), and winter (December, January, February). The sex f the animal, ttal length (TL, measured frm the tip f the rstrum t the psterir end f the telsn in live specimens), and carapaee length (CL, measured frm the tip f the rstrum t the psterir end f the carapace) were recrded fr each individual. A11 measurements were made t the nearest millimeter. Althugh specimens as small as 1.5 mm TL were cllected, individuals belw apprximately 3 mm in ttal length culd nt be sexed and were classified as juveniles. Female vary width was measured n the secnd abdminal segment at the widest pint visible sample. The ppulatin f S. trilbat r.r'as biased tu-ard females with a rati f 1:1.81. Males ransed in ttal length frm 31 t 82 mm with a mean (+ SEt f 51.6 t 5.4. Females had a mean (t SE) length f 68.2 t 2.2. ranged in ttal length frm 34 t 88 mm. and u'ere significantly larger than males (P =.1) u'hen cmpared with a Mann Whitney U Test. Althu-sh variable. this difference in size was cnsistent ver the entire sampling perid (Figure 2) with the exceptin f Octber 2. thrugh the transparent drsal integument f the secnd abdminal smite, and the number f eggs was als recrded fr each female when vigerus. Females thught t be releasing eggs during the time f cllectin were excluded because it was thught they wuld bias the results. Females classified as thught t be releasing eggs were thse that had a small number f well develped eggs remaining n the plepds and had bviusly released the remainder f the brd. This resulted in the exclusin f a maximum f 1-15 specimens thrughut the entire study. Eggs were als categrized accrding t their clr (red range, range, yellw range, r yellw) and whether r nt they Ovigerus females were cliected frm April thrugh Octber 2, and March thru-eh August 21 (Figure 3A). During the mnths f April and May in bth years, the prprtin f vigerus females cllected was larger than during the rest f the year. The time perid during which vigerus females were present can be directly crrelated with the vary width f the females during the preceding mnths. During the mnths cntained visible eyespts. After measurements were recrded, specimens were archived in 7)V ethanl. f Nvember 2 t February 21, when n vigerus females were cllected, a peak in mean vary width f females was bserved. There was als a decline in mean vary width starting in March and cntinuing thrugh June, when the highest prprtin f vigerus females Rpsur,rs The prprtin f individuals in each size class varied seasnally (Figure 1). n spring 2 the ppulatin was dminated by individuals in the larger size were cllected (Figure 38). The average number f eggs cllected per female was 197 t 18.1 eggs, and a psitive crrelatin existed t4
PpulnrrN Brt-cy SERGT TRTLBATA G t, E tr c a A..6 B.,5.4.3.2.1.6.5.4.3.2.1 Spring 2 N= 66 2-29 3-39 4-49 5-59 6-69 7-79 8-89 t- - 2-29 3-39 4-49 5-59 6-69 7-79 8-89 c..6.5.4.3.2.1 Ttal Length (mm) Summer 2 N= 136 - Ttal Length (mm) Fall 2 N= 74 2-29 3-39 4-49 5-59 6-69 7-79 8-89 Ttal Length (mm) D,.6 E,5.5.4.3.2.'1 F..5.4.3.2.1.6.5.4.3.2.1 Winter 2 N= 8 2-29 3-39 4-49 5-59 6-69 7-79 8-89 Ttal Length (mm) Sprinq 21 N= 6 2-29 3-39 4-49 5-59 6-69 7-79 8-89 Ttal Length (mm) Summer 21 N= 54 2-29 3-39 4-49 5-59 6-69 7-79 8-89 Ttal Length (mm) Figure L. Seasnal variatin in the prprtinf Sergi trilbata in each f six L-mm ttal length size categries: A) Spring 2; B) Summer 2; C) Fall 2; D) Winter 2; E) Spring 21; F) Summer 21. between number f eggs per female and its ttal length (r2-.929, n= 56) (Figure 4). The highest fecundity bserved was 42, althugh this may be an underestimate f ptential fecundity as sme eggs may be lst in the cllectin prcess. Eggs ranged in clr frm redrange immediately after release frm the vary t a pale yellw cntaining distinct eyespts in the days prir t hatching. The smallest female fund t be vigerus was 48 mm in ttal length with 62 eggs, and the smallest female cllected with develped varies was 41 mm in ttal length with varies 2 mm wide. DrscussrN Seasnal variatin in the prprtin f individuals in each size class was bserved during the study suggesting that there may be envirnmental factrs affecting the ghst shrimp. Additinally, a life span f apprximately tw years is indicated by the seasnal variatin in the prprtin f individuals in each size. The bimdal distributin f the ppulatin in winter 2 and spring 21 indicate the presence f tw generatins in the burrwed ppulatin. Maturity f thse recently recruited then fllws the next year as vigerus females are mst cmmnly fund in the spring and summer. These bservatins alng with the apparent mrtality in the largest size classes in fall 2 and 21 suggest the life span f these rganisms may be limited t tw years. The life span f thalassinidean shrimp is variable, but there are sme that have cmparable life spans f tw years, such as with Lepidphthalmus luisianensis (Felder and Lvett 1 989), Callianassa japnica (Tamaki et al. 1997), and Callianassa kraussi (Frbes 1977). Sme ther thalassinidean shrimp exhibit a lnger life span f apprximately 3-4 years such as Upgebia pusilla (Kevrekidis et al. 1997), Sergi mirim (Pezzut 1998), and C al i an a s s a fi h i (B erkenbu s c h and R w den 2 ). The seasnal variatin in the prprtin f individuals in each size class in the ppulatin is als related t the 15
Cnssru AND STRASSER 65 E s Er c 6 (l) J 6 L 55 E) 6 q) Apr- May- Jun- Aug- Sep- ct- Nv- Dec- Jan-1 Feb-1 Mar-1 Apr-1 May-1 Jun-'l Jul-1 Aug-1 Mnth f Cllectin Figure 2. Mean ttal length (t SE) f male (n = 138) and femala (n = 297) Sergi trilbata during each mnth f cllectin (April 2 t August 21). fecundity f the rganisms and recruitment f juveniles int the ppulatin. Fecundity and recruitment may be affected by temperature r salinity fluctuatins, althugh the exact cause f these variatins warrants further study. Salinity was fund t range frm 25 t 38% but des nt seem t impact the reprductive utput f S. trilbata as the fluctuatins in salinity did nt mirrr trends in reprductin. Althugh exact temperatures were nt recrded n a regular basis, seasnal temperature changes were the mst likely cue fr reprductin since data cllected des indicate a seasnal trend in vary size and egg prductin f females. Hwever, even if surface water temperatures were cllected, they might have been misleading since the water temperature in the burrws is ften lwer than the surface water temperature. The ppulatin f ghst shrimp was fund t be biased tward females with a 1: 1.81 male:female rati. Sex ratis have been fund t be variable in ther thalassinidean shrimp; fr example, Sergi mirim (P ezztt 1 998), Lepidphthalmus l uis ianensis (Felder and Lvett 1989, Felder and Griffis 1994) Lepidphthalmus sinuensis Lemaitre (Nates and Felder 1999), Callianassa filhli (Devine 1966, Berkenbusch and Rwden 2), Callianassa japnica Ortmann (Tamaki et al. 1997), Callichirus armatus (Vaugelas et al. 1986), and Callichirus islagrande (Felder and Griffis 1994) exhibit a female biased sex rati. The sex rati has als cmmnly been reprted as male biased in ther species such as with Callichirus kraussi (Frbes 1977) and Callianassa subterranea (Rwden and Jnes 1994). The variability in the sex rati f ghst shrimp can be attributed t many factrs including seasnal variatin and variatin between species (Berkenbusch and Rwden 2). The variability may als have a behaviral basis; fr example, females with eggs are ften fund clser t the burrw entrance than males, which wuld make them mre susceptible t cllectin (Nates and Felder reee). The mean TL f females was larger than the males cllected frm April until Octber 21, with the nted exceptin f Octber 2. Ppulatins in which females are larger than males is cmparable with ther thalassinidean shrimp such as Callianassa filhli (Berkenbusch and Rwden 2) and Sergi mirim (Pezzut 1998) in which the females cllected were als 16
PpulauN BrLGy Ssncr TRTLBATA A. B. -!P (t' E (, tt, c) ED.9 CL -.8.7.6.5.4.3.2.1 e7 E -c E'E '= r4 (E B3 (EZ (u =1 \' A Apr- May- Jun- Jul- Aug- Sep- Oct- Nv- Dec- Jan-1 Feb-1 Mar-1 Apr-1 May-1 r r r --+J ". _Y\ ffiy r Lr \L h. Y Y- Apr- May- Jun- Jul- Aug- Sep- ct- Nv- Dec- Jan-1 Feb-1 Mar-1 Apr-1 May-1 Jun-1 Jul-1 Aug-1 Jul-1 Aug-1 Mnth f Cllectin Figure 3. Reprductive data fr Sergi trilbata frm April 2 t August 21: A) Prprtin f female sample vigerus during each mnth f cllectin; B) Mean vary width (t SE) f females during the cllectin perid. 25 a,^ 2 E,, c,) UJ g 1s ) z G E t a,/ y=53.13x -G4.227 R2 =.9287 5./ ^r/v 4-49 5-59 6-69 7-79 Ttal Length (in 1 mm increments) Figure 4. Mean number f eggs per females f Sergi trilbata in each f six L-mm ttal length (TL) size categry. l7
Cnserrr en SrnessER TABLE 1 Cmparisn f the mean and maximum number f eggs f female thalassinidean shrimp in selected studies. (- ) indicates n value was prvided in reference. Species Sergi trilbata Lepidphthalmus sinuensis Callianassa tyrrhena Callianassa japnica Le pidphthalmu s ui s ianen s i s Sergi mirim Mean Egg N. MaximumEgg N. 197 4t2 present study 958 Nates and Felder 1999 Thessalu-Legaki and Kirtisis 1997 25t 27 1 33t 188 962 Tamaki et al. 1997 Nates et al. 1997 6586 Pezzut 1998 871 Phl 1946 598 Callichirus majr Reference prduced per female annually, as well as lar-qe e-u-u size (Thessalu-Legaki and Kirtsis 1997. Nates and Felder 1999, Berkenbusch and Rwden 2.1. Althugh 1'ear rcrund reprductive activity is nt likel,r' rn S. trilbata. due t the apparent seasnal trends in reprductin. the pssibility that S. trilbala prduces ilultiple brds in larger in mean size than the males in the ppulatin. There are, hwever, many thalasinidean shrimp in which the males exhibit a larger mean TL than the females such as Upgebia pusilla (Kevrekidis et al. 1997) and Callianassa japnica Oftman (Tamaki et al. 1997).n many cases when males are larger than females in the ppulatin, the size difference can be attributed t male t male cmpetitin. Hwever, in this case males were fund t be smaller than females prviding n evidence f intrasexual selectin r sex-related mrtality which wuld cause the female biased ppulatin. The increased prprtin f vigerus females can be directly crrelated with an increasing vary width f the females during the preceding mnths. During the summer mnths, the prprtin f vigerus females increases as the mean vary width decreases. Crrelatins between the mean vary width and the prprtin f vigerus females in ppulatins f ghst shrimp has als been bserved in ther thalassinidean shrimp such as Lepidphthalmus luisianensls (Felder and Griffis 1994), Lepidphthalmus sinuensis (Nates and Felder 1999), Callianassa japnica (Tamaki et al. 1997) and Netrypaea califrniensis (Dumbauld et al. 1996). The ttal length f S. trilbata was als fund t be psitively crrelated with the number f eggs prduced, which is similar t results fund with many ther ghst shrimp species such as Callianasa tyrrhena (Thessalu- a given breeding seasn cannt be eliminated. Additinally, althugh eg-e size was nt measured in this study, eggs were fund t be much lar-uer than that f a sympatric species f ghst shrimp. Lepidplttltahnus luisianerzsis (persnal bservatin). Eg-s size has been directly crrelated with larval develpment in many species f ghst shrimp in which species u-ith larger eggs exhibit abbreviated develpment in 1arva1 stages (Frbes 1973). The egg size and number f brds prduced annually wuld be f significance in determining the fecundity and the develpment f larval stages f S. trilbata and therefre warrant further study. AcTNwLEDGMENTS We wuld like t thank all thse wh assisted in cllectin f rganisms, namely Mark Driscll, Amanda Kahn, Kelly Graham, and students in the University f Tampa's Eclgy class in 2 and 21. Funding fr this study was prvided t J. Crsetti by the University f Tampa Bilgy Research Fellwship and the University f Tampa Hnrs Research Fellwship and t K.M. Strasser thrugh a University f Tampa Dana Grant. We wuld als like t thank W. Wayne Price and Kevin Legaki and Kirtisis 1997), Callianassa filhli (Berkenbusch and Rwden 2), and Sergi mirim (Rdrigues l97l) (Pezzut 1998). The mean and maximum number f eggs per female S. trilbata (mean 197, max. - 412) was slightly lwer than ther species f ghst shrimp, althugh the mean number f eggs per female in thalassinidean shrimp can be highly variable (Table 1). A lw number f eggs per female is indicative f year rund reprductive activity r multiple brds Beach and three annymus reviewers fr their cmments n this manuscript. 18
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