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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 Extended parental care in two endobenthic amphipods M. Thiel a; S. Sampson a; L. Watling a a Darling Marine Center, University of Maine. Walpole, ME. USA

To cite this Article: Thiel, M., Sampson, S. and Watling, L. , 'Extended parental care in two endobenthic amphipods', Journal of Natural History, 31:5, 713 - 725 To link to this article: DOI: 10.1080/00222939700770351 URL: http://dx.doi.org/10.1080/00222939700770351

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JOURNAL Extended Astacilla Introduction polychaetes more some care , environment delIavallei Cedhagen, Heterotanais (Aoki Caprella h tanaids the 1994). extended 1995), The While beyond time energy and The the majority OF aetl care parental scaura arietina Brooding parental in juveniles Rctr 1978b) (Richter, NATURALHISTORY, estuarine Darling Extended M. inhabit clutch species, ulig their building females tasks to after (Accepted in protect eti, otbto, intertidal. soft-bottom, benthic, KEYWORDS: 1989), Kikuchi, marine parental (Dales, the where the Tanais most into occur irrigate L. oerstedi THIEL, juvenile they pinguis, water of of typica of the their small care Dulichia Leptocheirus pinguis female increase (Sars, sophisticated fully marine of Marine of juveniles parental in amphipods soft-bottoms hatch and 1950; dulongii female's 5 1991), column both Reproduction, small September (Btickle-Ramirez, is until S. offspring the offspring care own C. maintain developed stage (Lim L. 0022-2933/97 documented. from 1899), SAMPSON the bigelowi species Bartels-Hardege macrobenthic pinguis peracarids. falcata Center, care juveniles a Peramphithoe ml tubes small in Dyopedos burrow juvenile size before in 1997, (Johnson is the and the is from along the rare. Arcturus 1996) brooding by two of leave brood described and burrow and small Alexander, deep $12'00 their University (Kanneworff simultaneously, 31, xedd aetl care, parental extended releasing settling pbnhc . epibenthic and Most is and stage the C. endobenthic the other The at The pouch. 713--725 known burrows and monacanthus burrow. ueie hatch juveniles bigelowi 1997 © Casco bigelowi, raim have organisms the 1965) Gulf female's of L. baffini stypotrupetes cases for two behaviour occurrence sedentaria is and uebro-iig peracarids tube/burrow-living to the marine bottom Attramadal, WATLING not Several advanced of Taylor the of for 1986), It during female. Zeek, are during of Maine, and uncommon is Maine. burrow (Svavarsson bottom. but endobenthic small hypothesized extended & of among isopods consecutive Francis in the Nicolaisen, the has and amphipods C. of 1990) the (Richter, remain Larger juveniles Larger C. from larvae bivalves (Conlan Walpole, at The time planktonic female's extended monoceros bigelowi However, 1982; time , Ltd. about evolved D. those in parental Astacilla and juveniles when amphipods the in porrectus of and or the that the clutches half Borowsky, and xedd parental extended female's (Gallardo 1978a; echinoderms burrow. ME even there is there 1973), peracarids the marine aetl care parental burrow extended Davidsdottir, larvae some in dl size. adult care Chess, juveniles and 04573, of of is , longicornis fully some which of always a both Siphonoecetes Laval, L. Casco (Mattson species of in the in environment, brood L. C. mechanism which 1993), parental pinguis their pinguis 1992), 1983) developed for inhabit The decipiens USA peracarid grow. amphipod (Sewell, only bigelowi marine pouch. among mother 1980), care invest which spend major 1994, endo- some start care The and and and and one can the to is Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 juveniles juveniles parental males environment. produced 714 Study Methods care. there burrow essential. examinations extended several very which iscotta clay. Sampling ae the take parents. those 3"5% station soft openings June inhabitants date ensured which had which telson. were Burrow Results those -1 sampling id six find system. inhabitants burrow The In Leptocheirus and sediments (12 The organic The through different is a is ovigerous in females which area is order inhabit River just All times aeul u quickly but carefully burrows containing The extends structure study the 19°C care December whole For that the whole variety are reared at parental In station by of adult been at the of shallow can amphipods amphipods most did to Estuary, cared only ah female each the We Mean in beginning over independently. of only which was burrows peracarid Leptocheirus (water collect sediment sediment or amphipods in collected two not 3-6 usually the were will pinguis. size matter and hint of care, the not. for the conducted Low the 1993). host subtidal were endobenthic cm aetjvnl relationship parent-juvenile the Maine. restrict by burrows organization content: emersed measurements course indicating individual In of in were inhabitants a of content Water below above e found be juveniles each female surface. orders with other in the detailed rearing are estuarine Our September were adults The order to pinguis Salinities are Salinities (unpublished In our measured following and of in available. of the main objective main 70-80%) marine for (MLW) studies the order suggests sexed of h year. the Lowes burrow discussion individual One juveniles the amphipods, burrows collected and burrow. knowledge in about the at sediment of soft-bottoms. M. find and were occurrence their we of to juveniles. and the 1993. invertebrates sediments h finding the along during one we use the use we Thiel Cove, We us identify one out at at of ht this that used about The data). for burrow, bottom and After in collected then used then parents, what its to the individual L. Casco describe up hour. their their ufc and surface Leptocheirus the et was to was of peracarid spring sediment naiat with inhabitants which hi inhabitants, their a low pinguis 30, to al. Both We size of retrieving this or the varies females optrbsd video-image computer-based In what dorsal burrows, of shaft term reproductive bigelowi and and with extended the ie mark. tide measured document addition, low every juveniles factors these his form s a is the amphipod water burrow between at crustaceans, size numbers in has 'rearing some form fingers of surface tides, it first those each sheltered juveniles the pinguis of was the two and their the were leading a temperatures parental reproductive observations we cove the size the together when sampling U-shaped eiet we sediment the Sampling a determined juveniles in were 'non-rearing of species behaviour weeks 2'5 female' from we spoon. and burrow only examined a numbers ern females rearing and of cove are is grab-like in despite located to the sizes sampled. extended the care. muddy the of collected the released Casco over with can tidal In date exclusively in was stay all rostrum form (Fig. intertidal vary whether behaviour evolution has this of of No the and the be the opened female' individuals how females the we flats at flats manner to with their with done bigelowi, juveniles extended between found fact parental analysis detailed evolved way n the and Damar- into the Burrow 1). sizes burrow tried muddy on period to the to many they may that The at and one the the we for for of or of in to to is a Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 juveniles juveniles juveniles before own Not be F~6. In U-shaped The they large No female mucus-lined stable, and of surface (Fig. some freshly up Casco information they burrow much 1 tubes are juveniles. 1). the typical liquid bottom to it the stays (a) (Fig. frequently cases reaches It 10cm burrow. are established usually inhabit burrows bigelowi. burrow excavated at sediment appears Burrow N wall burrow and the flush 2). at walls immediately the deep Water the is juveniles The bottom the is variously live wall amphipods with which available with very modified that with and bottom reworking female. (Fig. continuous sediments their The at is is the the a soft female of usually drawn the in can firm after rearing Extended structure 1). own the juvenile the surrounding branched of While on and by bottom extend At and retrieval were the female right at seems their how into burrow at bordering modification organized female the the oxygenated, burrow the sampling branch also of tubes parental the down of the bottom inhabitants surface burrows to burrow and the surface. the as burrow Leptocheirus sediment female found occur later fill of burrows to immediately side-branches sediments burrow, and in the the the more care the (Fig. of freeze-dried but which a after in juveniles Laboratory U-shaped and maintains (Fig. burrows two the manner entire simple in surface. no than of 1). first with the burrow pinguis amphipods burrow are tube C. may 2). These diameter 10 behind establishment in bigelowi passes burrow; Single rarely tubes from in larger resembling in cm with the The tubes have observations the the and can juvenile openings below burrow their with burrow field, lab; mucus-lined through oxygenated. the juveniles inhabitants in its sediment be multiple of is bottom juveniles; (b) inferred tubes the much mother's a the we tubes a cave while simplified of have shape U-shaped the sediment honeycomb. sometimes have core the of of openings inhabiting less juvenile usually at are right from Females a walls rearing the burrow. female was is burrow. the sharp regular shown built drawing relatively juveniles branch frozen mounds burrow. bottom. surface. is live to found edge, built. at tubes these from their with The The that 715 and the the in of of in Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 716 F~. 3. juveniles its Photograph juveniles: Frc. visible 2. female of Burrow the in horizontal interior transporting of rearing of burrow a burrow mud M. female Thiel gallery; bolus of Casco Casco et (left) scale al. bigelowi; bigelowi, and bar female = lcm. scale showing irrigating bar the = lcm. female the burrow and some (right); of Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 pinguis juveniles, Reproductive The between females Numbers females December by non-rearing other (Fig. rearing burrow burrow modify #1). females two length found, towards of female (Fig. were developing at sediment The between burrows) found 5 smallest Juveniles estuarine Discussion size. ram, different sampling. the Leptocheirus From Leptocheirus Casco Extended Casco clutches size smallest Between In 4). times 4a). ovigerous in their and female. L. and female. by the of no no at were the (Fig. juveniles the of 13.5 their have and pinguis, the soft-bottoms. is grew but The bigelowi. their 21 an the burrow burrow bigelowi. clear (June juveniles positions the 1993 numbers at females two beginning also of juvenile limited and parental stage non-ovigerous. adult later sizes The July 7). June mothers' been largest 1 non-rearing same The juveniles In to mother but species, September temporal pinguis. and deposited 21.8 (mean pinguis. At but > June which were we (Fig. shape average and of female. and found size after 0.5 was in information in the trend in August) juveniles found there were Also mm. care did of Between The the the the (Fig. and burrows the the size 4b). but two of 23-02mm end always can continuously Extended August we this not in trend female's On burrow. beginning can females was in about were In brood the Up within early structure adult Juveniles in number at we of the clutches 3). In easily found of there find C. date one both average, that females rearing be in to found July found burrow July/beginning below were is also bigelowi 35 on July (Fig. size females' 10 pouch large case 144 observed the date recognizable. was the cases ± The parental varied be in burrows the mm no of and and females of 0-85 of during most > in of distinguished (Fig. the 6). juveniles burrow. proportion only the females always a juveniles 15 juveniles was males July, size numbers of 5 juveniles two only (Fig. (Fig. male maintenance L. mm 58 the mm At between burrows rearing one burrows s.e.; rearing 3). care 3-6 one of for pinguis clearly the endobenthic the there juveniles which females care 4a). 7). (Fig. were (Fig. was at female They the Rearing mm varied of n non-rearing have in period end rearing The least of of When recognizable females = in of of At rearing were August 8" females found of amphipods 7). small C. decreased hosted found 5). long (see 7). also 1 ovigerous their of were the engaged the size one C. been of their and bigelowi between juveniles of August (mean females always At e.g. juveniles female the bigelowi ventilate and juveniles end in amphipods were male of attending and extended the mother's only 19"8 always were mother found this the burrow females Fig. the the of in (Fig. non-rearing majority (Fig. always mm. found 45.0 some females no move five burrow 19"6 July the with have time either largest rearing 7, were at the in was rearing in accompanied 7). were juveniles care 8 parental females, differs The the + 8). burrows and females no which the with June attending about the burrow 17 been all > In 3.31 larger exclusively ovigerous of of increased was 12th rearing of 15 average females juveniles. burrow about 25'8mm burrows some seven an females no females the the 1993, substantially mm freely both observed s.e.; care. 12"6 December. but than ovigerous, at with frequently decreased ovigerous juveniles. juveniles burrows, a 2 the (Fig. females rearing female female inhabit mm during size of mm or in varied n of in about again small at done were of time The 717 = had one the the the 5). 12 of L. to in in 7 Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 718 100 FIG. 60 20- % % . (a) 4. single paired pinguis September. at sampling 8 August, Reproductive females females between date; No none n=R therefore effort 8 June was stage Imwith []with was and found numbers n=R of made 1 juveniles juveniles rearing content of pouch brood content September at to M. 20 vary n=6 collect females; August Thiel greatly 1993. similar et []with [] and n=13 and between One al. iheg • empty • eggs with only (b) numbers rearing eggs non-rearing n rearing one the n=40 female various of non-rearing was females dates. female n=6 []empty lost during of females was ales I females Leptocheirus found sampling at ] each at ] 1 Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 FIG. burrow Biology FIG. it early (Wildish, .~ __N o O. E e~ z,. .o O E E E pumps 100 10- 20- The 15- To 6. 5. 40- 80" 20 60- 5- summer. between were 1 - our on September. Mean Mean 8June data of 8 through 1980), June several present Leptocheirus counted; numbers suggest size 8 Several June so pinguis; (+standard occasions. ~ its knowledge, I and no each 22 22 (_+ burrow. that June rearing clutches June I standard pinguis September female n Extended L. = error) pinguis Casco numbers ' I Casco females L, error) of and only pinguis 6July 6 of different July 1993; parental bigelowi rearing produces Casco bigelowi of hosted of were juvenile each females filters I I found females 21 bigelowi sizes 21 care females, one sampling July July is suspended several Leptocheirus '6 measure; in a clutch at (and deposit-feeder and amphipods 20 ~ | date in consecutive ages) non-rearing August contrast, of 6 6Aug. see Aug. particles the juveniles pinguis also juveniles were and ~ Fig. I produce • • excavating in found females out only 20/[ug. broods 20 each non-rearing rearing in 4. of Aug. of its six one in the female's only of burrow. rearing in one [ was water Leptocheirus the subsurface one 1 1 female's Sept. found Sept. burrows females spring/ which brood 1 719 at ( Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007

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o. ° ] _.._.1,. - -.'-.' .I 4 6 8 2 4 6 IJ - 2 4 6 " 8 " size classes (mm) size classes (mm) FIG. 7. Numbers of juvenile Leptocheirus pinguis in the size classes found in burrows of each individual female (#1-6) at the respective sampling dates (8 June- 1 September 1993); no rearing females were found at 20 August, and only one was found on 1 September 1993. Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 L. juvenile deposits within FIG. manipulating Juvenile of sediments these their rearing irrigation feeding Extended irrigating the population irrigation. female in which employed there In parents distinguish In species environment pinguis the feeding the at Most sediment 8. own juveniles least may female. following, we following, display female actively burrow which at Number C. references C. parental activities. it. tubes the and The burrows The failed loosened a mode dynamics these bigelowi be The few bigelowi surface, as time sediment Juvenile C. the engage C. start more, its 8 8t 04 might care for at soon of to peracarid bigelowi peracarid diameter bigelowi of common burrow. During burrow. the when juvenile care reveal to by on We 8 0 2 8 0 12 10 8 6 12 10 8 6 during the will of as a as bottom 12 in become also were parental C. her of depth individual within growing in the noticed they juveniles they extended eebr I December e unrecognized, yet Extended refer bigelowi probably their rearing species with multiple with marine burrowing of reproductive Casco bigelowi Casco species his be two observed start hatch of not the true to the study. the extended care juveniles that the amphipods female #7 these achievable females burrows peracarids the building with parental L. filter-feeders. have burrow parental reason the have from female's of medium-sized pinguis time His activities. size juvenile xedd parental extended time species to (#1-7) strategy never openings two parental the intensive after L. in periods why ty close stay is their care. on observations). (own classes L. care pinguis amphipod period major by h sz cass found classes size the female's burrow, which burrow, too from pinguis been at the at they as offspring After Wildish an own where We in big care. those niae increased indicated of tasks: individual study amphipods juveniles hatch of deposit-feeding observed (ram) 12 and #21 speculate tubes extended parental extended for smaller for the brood parental and to eebr 1993. December the This (peracarid, with C. (1980) are provided maintaining from juveniles care their juveniles in C. bigelowi fact probably from extended pouch. L. the to juvenile. bigelowi, from that the care pinguis engage mother They never burrow leads us leads juveniles female's peracarid in important 10 there marine are the or are Not is are h burrows the burrowing a is probably parental extended, grazing born, care, rest The encountered deep in but released necessary started hl she while might #3 of 6-10 surprisingly, to active brood their to of invertebrate) the depth crustaceans. suspect the burrow burrows data peracarids engage cm be care. to building methods ed on feed females mother. activity into burrow of so pouch. on below a of filter- when shift each was 721 that that and any We the the the of in in Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 the other can FIG. histories. isopods, peracafids, environment epibenthic The coastal species 722 Peracarids Extended be 9. two eia order): betical (1947). Conlan Attramadal (1982); Attramadal 18) Mtsn n Cdae (1989); Cedhagen and Mattson (1986); 19) Silkr and Shillaker (1899); deposit-, Tube distinguished: environments, with extended Classification endobenthic amphipods where Among or structure which burrow suspension-, than and parental from the Chess the in include pelagic include iuhabitants; juveniles Aoki the built and pce without species of benthic tube one arnphipods mobile parental where detritus- burrow care Nematotanais Siphonoecetes dellavallei Siphonoecetes eeoaas oerstedi Heterotanais Phronima sedentaria Leptocheirus pilosus Leptocheirus Lembos Peramphithoe stypotupetes Peramphithoe Leptocheirus pinguis Leptocheirus Tanais dulongii Tanais Casco bigelowi Casco Corophium volutator Corophium (1992); peracarids with extended parental care. parental extended with peracarids first Kanneworff n iuh (91; idadHlih 18) BikeRmrz (1965); Btickle-Ramirez (1985); Holdich and Bird (1991); Kikuchi and tanaids by where i Moore on/in is category are the websteri peracarids the or burrow/house reported in the Chess filter-feeding living disturbance the female hard-bottom (Hassack mirabilis Leptocheirus pinguis soft-bottom 18) Saaso ad aisotr (1994, Davidsdottir and Svavarsson (1987); juveniles amphipods and juveniles parental M. (1993); Goodhart (1939); Harrison (1940); Harrison (1939); Goodhart (1993); are exposed from with Nicolaisen (Fig. Thiel almost and McCloskey are potential extended a 9). are care. Rctr 1978a, (Richter, whole et on Holdich, much mostly Epibenthic, exclusively al. sheltered the 17) Lvl (1980); Laval (1973); ~l larger variety and body ~', parental suspension-feeding 17) Rctr 17a 1978b); (1978a, Richter (1970); and 1987) predation juveniles in of Casco before References of species on a the care, Laval, tube, with phylogenetically Arcturus baffini Arcturus arietina Astacilla Iongicornis Astacilla Aeginina longicornis Aeginina ypds porrectus Dyopedos monacanthus Dyopedos Dulichia falcata rhabdoplastis Dulichia Caprella decipiens Caprella Pseudoprotella on monoceros Caprella typica scaura Caprella on structure lifted female bigelowi being pressure female female two a burrow or from above 1980), wide Lim are major or released ul by built bottom; body shallow-water from antennae attached to attached belong phasma n Alexander and 1995); range is very is and Johnson categories cave, (in into the benthic distinct female Watkin of to alpha- high. Sars this and life and an Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 relatively peracarids Mediterranean. continental might reproductive benthic extended first lei inhabit predation burrow. big and big for their epibenthic predators sheltered expect delivery category this Extended evolution methodological thorough boundary predation peracarids bigelowi as deposit-feeders. mudflats invertebrates important is incomplete. species and Acknowledgements Studies 1985), depicted unknown, (Richter, The Our Our a group Alexander, category mother's caprellid be more be whereas extended deeper boundary other at deep Other particular (Lim classification without are suspension-feeders. which of parental laboratory parental pressure seems to of ih predation high layer) and examination on the clues with by might Maine, hl where shelf The extended 1978b; behaviour. of parent's and (see, category the University particularly burrows. the caprellid and tube-dwellers, obvious tubes Tube-dwelling larger have reached species 1986). burrow/tube extended limitations. M.T. major reason parental care parental for are parental layer offspring that care thanks be e.g. care while Alexander, Conlan environments, such valid, a or burrows. or parental individuals was burrow/tube. However, an of of assumptions in into Duffy, to better In with of in Intraspecific abundant he peracarids of this Extended parental as peracarid pressure, an assistance. go caprellid soft-bottom supported important the water and but favour amphipods this (see was Maine. , We inhabitants. epibenthic predator to sizes e.g. extended category understanding for care within tanaids marine 1996). Chess, other 1986). Larry Mayer Larry Very presocial assume supposed also probably layers are relatively This Peramphithoe this is this care, a of (protection Aoki ground-fish males parental such bu their about sheltered with species hermit by about competition strategy factors reports this small with environment environments would 1992; of and Extended parental with a high that Their and that extended as graduate peracarids have been group more to extended the arcturid extended crabs, 6 juveniles the with care the sufficient Chess, study disturbance Kikuchi to of need our in who life also juveniles from to safe important stocks 12 East reproductive life stypotrupetes Mattson higher of care than in style general brachyuran parental extended and reduce mm parental gave be fellowship the feeding to peracarids. from predation, amphipods history 1993), found is small parental might parental Pacific a than e considered be (1991) true flow predation of the fact isopods. at occur. single grow social M.T. epibenthic potential small the time and conclusions with intraspecific to recruits for also care for care speeds not be of parental attack found much found kelp care live crabs to individual. biology from the Cedhagen, juvenile or reaching above many Thus, and that behaviour care juveniles A respect one live Intuitively leads remarkably within can Siphonoecetes female-offspring freedom of able and forests are very digestive (in the all and and in not predation marine also leaving be in care n this in a in environments shallow species are suspended with to to Center L. very fishes prey future competition engaging important this depths along in Interestingly, the reach above higher pinguis 1989; might or among most counterfeit one to the safety susceptible regard large invertebrates exposed behaviour on the evolution category category h shallow the explore (e.g. the benthic as for within studies. waters) would their dellaval- and female's likely they survival provide particle sizes Marine marine and in within Reise, to group with own Lim life, 723 of this this and can not the the the the by C. or of of of of in to in A is a Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 Aora, M. Aora, 724 BARTELS-HARDEGE, BIRD, References BOROWSKY. BOCKLE-RAMIREZ, Du~v, CHESS, DALES, CONLAN, HASSACK, HARRISON, GALLARDO, GOODHART, JOHNSON. KANNEWORFF, LAVAL, LIM, MATTSON, McCLOSKEY, POCHa'ER, RICHTER, REISE. SEWELL, SARS, . T. S. G. G. monoceros Hydrobiotgia, Biology, phasma the 563-572. pilosus Journal Peramphithoe (Annelida: Polychaeta), amphipods tacean Kr6yer 595-603. Amphithoidae) cean 714-782. 483-493. (Bivalvia: Peracarida) with plankton, Biology 311-325. J. K., monacanthus aas cavolinii Tanais 10, Museum) CapreUa vallei, sea (Forskal), tuberculata clarki, R. J. . 1980, P., J. M. R., G., . E. K. S. O., E., G., and E. Rockall . 1950, P., R. 1985, urchin, and 119-129. S. C. A. C. Biology, B., A., B. 1993, and L. 1978b, 1996, J., Natur E. 1899, Biology, Biological 1978a, Zaddach, KIKUCHI, (Crustacea, and . 1993, S., HOLDICH, and Montagu, of B., and and 77, R., 1994, scaura 1983, and Tidal and Oceanography 1940, 270pp. Cyamiidae) HOLDICH, (Amphipoda), the Marine the L. Hyperiid Mayer, Trough, aii Science, Pacific Jassa Effects Boeck . D. H. Eusociality 257-263. 1939, ALEXANOER, Ecology, CEDHAOEN, An 90 A 1970, ATrRAMADAL, The und Einige NICOLAISEN, 223, 13, stypotrupetes, (Metzger) CHESS. F., Small Beobachtungen and fiat On 12, Reproductive account T., (Crustacea: typica, particular D. Journal Museum, Reproductive reproduction falcata 638-646. Bulletin, Journal Notes 1965, of 229-237. (Crustacea: ecology, 1890 Journal ), 410-422. grazing the D. and M., 1991, Beobachtungen amphipods new size, the J. 127, Biological in M., biology Marine Mayer in 1985, A 1985, . 1989, t., Senckenbergiana an C. ZEEK, and R., of stipe-boring and W., of Untersuchungen on and species and brooding of Y. Two a reference 253-272. 187, 108, 1987, estuarine sand gastropods 24, of the G., Tanaidacea), the Ecological coral-reef new the the Marine 1992, G., Caprella 1973, the Amphithoe behavior D. and Amphipoda: 1890, 90-98. zu Biology, habits E., of Crustacea types 259-266. Marine 112-123. 1986, remarkable Marine as Association Zeitschrifi The bionomics of M. Marine 1982, species the porrectus Aspects Entwicklung larval The and Phylogeny 1990, Dulichia zur to Biology, Marine amphipod Thiel tubicolous of Caprellidae. and Reproductive shrimp, on deep-sea decipiens Reproductive Studies, protandry Biological of "Haps", Lebensweise 106, Biological maternal Biological development Marine (Corophioidea: the maritima, flat valida of Reproductive life fiber three-tube-building fiir of tubicolous Podoceridae), et of Norway, Behaviour of Bate, 409-412. (Amphipoda, from kelp an Nature, cycle al. parasitoids the and Peramphithoe the the Morphologie 54, species, und die habit Annual and Biology, Mayer, a in care Growth Laminaria Association the Association frame-supported United behaviour tube-building Association with 191 10, ecology the behaviour of Biologie Verhalten behaviour des the Volume amongst tanaid of South 381, for the and 229-242. apodid (Berlin: Zoologica Review, behavior Amphithoidae), Nereis Flohkrebses comparative 1890 tanaid Journal 71, associated and Kingdom, juveniles Podoceridae) small 512-514. 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Tiere, Dyopedos Caprella Dulichia of Ophelia, oerstedi Crusta- Marine Miiller, Marine Bergen with della- Crus- from zoo- 115 65, 55, the 24, 23, of a Downloaded By: [Smithsonian Trpcl Res Inst] At: 20:49 24 May 2007 WILDISH, SHILLAKER, WATKIN, SVAVARSSON, SVAVARSSON, Ecology, Bate Ecology, estuary, the (Crustacea) 15, E. D. Arctic 569-574. R. E., and J., J. J. O. nentoa Journal International 1947, and and Corophium bonnellii Corophium 10, 110, 1980, and antenna-brooding as DAVIDSDOTTIR, 77-93. DAVIDSDOTTIR, MOORE, The 113-132. indicators Reproductive yearly P. Extended G., of life B. B., 1987, isopod rtrs baffini Arcturus cycle bionomics of Milne 1995, 1994, parental netbae Reproduction, Invertebrate The of aie Biology, behavior? Marine Cibicides Edwards, the biology Foraminiferan amphipod of care w sublittoral two (Crustacea, of spp. Journal in brooding amphipods (Protozoa, Corophium volutator, Journal volutator, Corophium (Protozoa) of Isopoda, Experimental Marine Biology and Biology Marine Experimental in the amphipods 2, Foraminifera) 118, 311-320. amphipods epizoites Valvifera), 239-246. in on Lembos websteri Lembos a as Polar Arctic Bay epizoites of of Biology, isopods Fundy 725 on