Life Cycle and Environmentally Induced Semelparity in the Shore Isopod Ligia cinerascens (Ligiidae) on a Cobble Shore along

Tokyo Bay, Central Japan Author(s): Toshio Furota and Takeshi Ito Reviewed work(s): Source: Journal of Crustacean Biology, Vol. 19, No. 4 (Nov., 1999), pp. 752-761 Published by: The Crustacean Society Stable URL: http://www.jstor.org/stable/1549299 . Accessed: 11/12/2011 10:31
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OF CRUSTACEAN JOURNAL BIOLOGY, 19(4): 752-761, 1999

LIFE CYCLE AND ENVIRONMENTALLYINDUCED SEMELPARITYIN THE SHORE ISOPOD LIGIA CINERASCENS(LIGIIDAE) ON A COBBLE SHORE ALONG TOKYO BAY, CENTRAL JAPAN Toshio Furota and Takeshi Ito
ABSTRACT A 3-year field study and 17-month rearing experiment were conducted on a population of the shore isopod Ligia cinerascens inhabiting a cobble shore in Tokyo Bay, central Japan. The research was designed to clarify aspects of life-cycle and reproductive traits in L. cinerascens. Most males, both in the field and in the rearing experiment, died before the onset of their second winter, but some field males overwintered twice. Most females overwintered once, bred from May through September, then died before the next winter. A small group of females, however, began breeding late in the season of the year of their birth, then most likely died without overwintering. Females in the field produced mostly 1 brood or at most 2 broods before dying, but those in the rearing experiment lived longer, producing up to 3 broods and possibly more. The results of this study indicate that the typical life cycle of L. cinerascens is annual with a 5-month breeding season, but including a shortened life cycle completed within a single breeding season. The data from the rearing experiment, however, suggest that the number of broods produced by a female may be limited by environmental factors rather than by genetic characteristics.

Shoreisopods of the genus Ligia are commonly distributedin coastal environments around the world. Variabilityin life-cycle have been observednot only characteristics among differentspecies, but also among lopopulationsof the same specally separated cies. For example,the longevityof L. oceanica (L.) varies from 1 year to 2.5 years in Europe (Jons, 1965; Willows, 1984, 1987), while thatof L. pallasii Brandtin BritishColumbiais up to 1.5 years (Carefoot,1973a). of broodsproduced The totalnumber by a female also varies,from 1 in L. dilatataBrandt in SouthAfrica (Koop and Field, 1980) to 3 or morein L. oceanica in England(Nicholls, 1931). These data indicate plasticity of the life-cycle patternin Ligia in responseto environmentalconditions. It is still unclear, however,if this plasticity arises from genotypic and/orphenotypicvariabilities. along the easternAsian Ligia is abundant butthe life historiesof Ligia in this seaboard, regionhave not been studied.This studywas conductedto clarify the life-cycle characteristics of a population of L. cinerascens Budde-Lund living on a cobble shore along northern TokyoBay, centralJapan.The lifeof the local field popucycle characteristics lation of L. cinerascensand those rearedunand conditionsare contrasted, der laboratory the results are comparedto data from other species of Ligia in otherpartsof the world.

MATERIALSAND METHODS Four species of Ligia have been recognized along Japanese coasts, and the distribution of each species has been shown to be clearly separated (Nunomura, 1983). Tokyo Bay is located on central Honshu, in the distribution area of L. exotica Roux, but L. cinerascens, which is basically distributedin Hokkaido, northernJapan(Nunomura, 1983), is common. The reason for this segregated distribution of L. cinerascens is not clear, but unintentional introductionis suspected (Itani, personal communication). Study Site.-Shinhama, a man-made lagoon of about 30 ha, is located along the northernmostshore of Tokyo Bay, on the central Pacific coast of Honshu (35?40'N, 139?56'E) (Fig. 1). The lagoon is connected with Tokyo Bay through a 0.5-km long channel. A large population of L. cinerascens is found on a cobble shore bordering the lagoon. The intertidal zone consists of 5-30-cm diameter cobble stones on the surface, with gravel and muddy sediment underneath.The horizontal distance between the highest and lowest tide marks is about 9 m with a maximum range of tidal level of approximately 1.5 m. Distribution of L. cinerascens is restricted to an area between the highest tide mark and the middle of the intertidal zone. Field Study.-Two transect lines (A and B) were established across the intertidalzone from the highest tide mark to the middle intertidal at points A and B in Fig. 1. Five stations (plus a sixth only during the warmerseason) were established along each line at 1-m intervals. The transect lines covered almost the entire width of the local distribution of L. cinerascens. To begin, quantitative samples were collected at each station in March and April 1989. All cobbles and stranded matter in a 25 x 25-cm quadratwere collected along with the surface sediment, using a shovel. The sediment was placed in a plastic box and all isopods in the box were collected.

752

FUROTA AND ITO: LIFE CYCLE OF LIGIA I I I

753

SHINHAMA -_ LAGOON
TOKYOBAY

36

bricks rand fragmentsn-d, 49~~~~~~~o"---~

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brackish water I

PACIFIC OCEAN
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139

140

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Fig. 2. Rearing system used in the experiment. A 5-cm thick layer of sand was placed in a 35-cm long x 25-cm wide x 30-cm deep plastic container, with holes in the bottom to drain water. This container was placed on a shallow tray which held brackish water sprayed regularly over the system.

Fig. 1. Map of Shinhama Lagoon. Dashed lines show the maximum extent of low tide, and A and B indicate the points where the study line was established.

Mesh basket sampling was then implemented from May 1989 to September 1991. A plastic mesh basket (30 x 40 cm wide, 10 cm deep, with about 2-mm mesh) was buried at each station. The tops of the baskets were set level with the surface. All baskets were filled with sediment, and cobbles and stranded material (if present) were added to provide hiding spaces for L. cinerascens. The habitat structurein the basket was thus similar to the natural habitat. During winter, an additional basket was buried at the highest station where overwintering L. cinerascens aggregated under stranded fragments of vegetation. At each sampling, the basket was removed without disturbing the contents and placed in a plastic container. All L. cinerascens were then collected by hand or with an insect aspirator,and fixed in 70% ethyl alcohol. Samples were collected monthly for most of the period, but more frequently during warmer seasons. Sampling was not implemented during January and February 1991. In the laboratory,the isopods collected were placed on a 1-mm scaled plastic film. Body length (BL) (distance between the top of the head and the end of the telson) was estimated to the nearest 0.1 mm under a stereomicroscope. Since a preliminary observation showed that the maximum error of BL was 1.23% from a mean of 10 replicate estimations among 5 individuals, estimation error caused by handling was considered negligible. Sex was determined only for individuals of 10.0-mm BL or more, by inspecting for the presence of the penis. All overwintered individuals, however, were sexed regardless of size. In gravid females, the number of eggs or juveniles in the brood pouch were counted. Rearing Experiment.-The container utilized for rearing isopods is shown in Fig. 2. Two sets of this apparatus

were used for the experiment. Ten females and 12 males collected at the study site on 16 May 1990 were placed in one container,and 10 females and 11 males in the other. None of the females were gravid when introduced. The experiment was continued to October 1991 without addition of new individuals. The rearing containers were set outside without protection from rain and sunlight. Every 3 or 4 days, the containers were showered with 0.5 1 of brackish water (1:1 fresh and sea water). The water was allowed to remain in the tray, keeping the rearing habitat wet. Goldfish food (consisting mainly of fish powder) and vegetables, such as pear, kiwi, cucumber, and lettuce, were used as food items instead of their natural diet, which consists mainly of intertidal diatoms and algae (Nicholls, 1931; Koop and Field, 1980, 1981; Willows, 1987). The pouches of the females were checked for the presence of eggs or juveniles every 2-4 days during the breeding season (May to September) of 1990. All gravid females were marked with individual color patterns. Maximum and minimum temperaturesduring each observation interval were recorded with a maximum/minimum thermometer placed on the sand surface in the containers. RESULTS

Field Observations Gravid females occurred from early May to early September in 1989, late April to midSeptember in 1990, and mid-May to late August in 1991. Mean size of manca-stage young was 3.0 ? 0.1-mm BL (SD, N = 122). Recruitment of newly released young (less than 4-mm BL) was observed at the site from June to September of all three years. This indicates that the breeding season, as demonstrated by the presence of gravid females, usually lasted from early May to mid-September, and that recruitment began about one month after the onset of gravidity. The three yearly cohorts (1990, 1991, 1992) showed single-size patterns, and the size overlap between yearly cohorts was limited (Fig. 3). This permits estimation of

:::::

40 E E 30 I 20 _. ' 10..C

1989 Mar. 29 -

May 10

June 20

July 31

Aug. 17

, ( 1 40 40 20

[1 ...... . :. :. . f 0 i I i 40 20 0 20 40

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60 40 20 0

........ . I.. . .
20 40 60

20 20

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r I

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No. of Ligia cinerascens

1990 40 [ ,I i I i ' I Mar. 30 May 16

June 13

July 19

Aug. 2

E
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,
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No. of Ligia cinerascens

1991 Mar. 29 May 17 June 14 July 19

40
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. l. 1 : .......... .:: . :!: :. :.:.:.: 20 40

1 31 60

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No. of Ligia cinerascens

Fig. 3. Seasonal changes in size distribution of the shore isopod Ligia cinerascens in Shinhama Lagoon, 1989-1991. Left, by black bars). Shaded bars indicate nonsexed young, which are divided equally on the male and female sides.

40-

A r -,
30-

E
E z 0 z
A

20-

10 -

J JA 1989

J J 1990

MONTH Fig. 4. Changes in mean size of cohorts of the shore isopod Ligia cinerascens in Shinhama Lagoon.

756

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 19, NO. 4, 1999

e-

1989 cohort

-000-r

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6
zo
*i?
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a 1 , ,

10: 4987

A
\
,I

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1989

J
1990

J
1991

Month
Fig. 5. Yearly change of mean density in cohorts of the shore isopod Ligia cinerascens on two observed transect lines in Shinhama Lagoon. Vertical bars show range.

growth (Fig. 4) and change in density (Fig. 5) for each cohort by size-histogram analysis. Growth rate of the 0+ (before first overwinter) isopods differed among the 3 yearly cohorts. Size of the 1+ (after first overwinter) isopods measured in March (about 15-mm BL), however, showed no significant difference either among the 3 yearly cohorts of the same sex, or between sexes in the same year (post-hoc Fisher's PLSD, after normalizing by logarithmic transformation, P > 0.05). During the breeding season, mean size of the 1+ males increased continuously from May (about 18-mm BL) to September (about 30-mm BL). The mean size for females, however, decreased in August (Fig. 4) due to the disappearance of larger females (Fig. 3). All male and female breeders disappeared immediately after the breeding season (Figs. 4, 5), except for the 1989 cohort, for which a small number of 1+ males were collected until November (about 38-mm BL). The largest males (40-mm BL) were found from March to June 1989. These males could possibly have been born in 1987 and experienced two overwinters (2+). Prior to the breeding season, in March and April, the sex ratio (female/male) for 1+ individuals varied among observations (Table 1). Female dominance was found in two of six observations. There was, however, no significant difference in the total ratio for all observations combined. This suggests that the

sex ratio of L. cinerascens at the study site is basically 1:1. Densities estimated at each observation station were converted to the number of L. cinerascens in a 1-m length of shore line (between the highest and lowest observation stations). The density of each cohort was maximum in early summer (June to July), but then gradually decreased until the beginning of the breeding season. An abrupt decrease was recorded during the breeding season (Fig. 5). Females become gravid only after reaching an adult body size of 21-mm BL or more. Only a few females, however, had attained
Table 1. Sex ratio in once-overwintered Ligia cinerascens.
Number of isopods collected Date Male Female Ratio (male/female)

1989 29 March 15 April 1990 30 March 28 April 1991 29 March 30 April Total

75 73 185 106 39 14 492

108 74 153 109 49 31 524

0.69*
0.99n.

1.21n
0.97n.s

0.8Ons

0.45**
0.94n s

Bimodal test: **, P < 0.001; *, P < 0.005; n.s., not significant.

FUROTA AND ITO:LIFECYCLEOF LIGIA

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Adult female
10, I

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15001

Newly recruitant

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Fig. 6. Short-term change in reproductive condition of adult females and new recruits (smaller than 4 mm BL) of the shore isopod Ligia cinerascens in Shinhama Lagoon, 1990 breeding season.

this adult size by the beginning of the breeding season. Most continued to grow, then achieved adult size and became gravid as the breeding season progressed. At the end of the breeding season (August and September), when most 1+ females had disappeared, some of the largest 0+ females (BL > 20 mm) started gravidity. After the breeding season (late September), however, the number of those 0+ adult-size females,

some of which may have already bred, decreased sharply (Fig. 3). This could indicate that most of these 0+ breeder females died before their first overwintering. Short-interval observations on new recruits (BL < 4 mm) and the reproductive condition of adult females were conducted in 1990 (Fig. 6). Density of all 1+ adult females (BL ? 21 mm) showed a peak in late June, then rapidly decreased through July and August. Density

758

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 19, NO. 4, 1999

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6
p

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0

o 0+ female

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Fig. 7.

10

I l

X l

X |

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I l

10

30 [Body length (mm)] 3 x 10-3

20

40

Relationship between body size (BL3) and brood size for the shore isopod Ligia cinerascens.

of gravid 1+ females, however, showed two peaks, in mid-May and late June. These peaks may be responsible for the two peaks in new recruit density located in mid-June and midJuly. Density of all 0+ adult females (BL > 20 mm) gradually increased after late August. Gravid 0+ females, however, were found only from late August to mid-September. A small peak in new recruit density was observed in late August. Since no gravid 1+ females were found after August, and some of the 0+ females carried fully developed eggs in which eyes had appeared, these late August recruits most likely resulted from release of young by the 0+ females. Size of brood (number of egg or juveniles in a brood pouch) was found to be positively dependent on BL3 (Fig. 7). No significant difference in the relationship was found between the 0+ and 1+ females (one-way ANCOVA, d.f. = 1, F = 1.498, P = 0.222). Average brood size was 94.5 ? 20.6 SD.

Rearing Experiment During the breeding season of May through September 1990, the number of gravid females in the first generation (collected at the study shore on 16 May 1990) showed three peaks, in mid-June, mid-July, and mid-August (Fig. 8). Four clear reductions in the number of surviving females were observed, following the gravid female peaks. The patterns of male mortality from July to September were similar to the female, but the overall male survival ratio was higher. At the end of the breeding season (25 September), four females (27.6-mm BL ? 0.5 SD) and 13 males (31.8mm BL ?+1.0 SD) of the first generation survived. All of these survivors were reconfirmed on 19 October, but none overwintered successfully. During the 1990 breeding season, a total of 39 gravidity cases were found among the 20 introduced females (some of the females be-

AND ITO:LIFECYCLEOF LIGIA FUROTA

759

25 20
C.) (n3

01)
C.z -4
%4.-

15 10 5 0 * 1 i 1 I

c5

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Fig. 8.

! S

Month
Survival curve for the shore isopod Ligia cinerascens reared during the 1990 breeding season.

came gravidmorethanonce). In all of these, except for one in which the female died before brood hatch, young were released. Gravidtime (egg depositionto youngrelease) decreasedwith increase in temperature, and varied from about one month in the early breeding season to about two weeks in the late season (Table2). Since paint markswere lost when the females molted during the interbroodperiod, the interbrood time was estimatedfor only 17 females thatwere remarked duringthe ecdysis. The interbrood time was 17.4 d ? 1.5 SD (N = 9) betweenthe first and second gravidity peaks(June-July),and 12.8 d ? 1.3 SD (N = 8) between the second and thirdgravidity peaks (July-August).Two females produced threebroodsbetweenmid-Mayandlate July, of broodsproduced but the total number over the entireseasoncouldnotbe determined, and may have even exceeded three. About half of the 0+ second generation young (334 males and 309 females) were removed from the rearingsystems on 19 July
Table 2. Gravid period (days) of Ligia cinerascens.
Monthof egg deposition Gravid period (?SD) Mean of medium (+SE) temperature Number observed

May June July August

29.6 + 1.8 22.4 +?1.1 15.9 ?3.9 13.6 ? 5.1

21.3 + 0.8 23.3 + 0.4 26.9 ?0.7 26.9 ?+0.4

6 8 15 6

and 15 August 1990. No gravidfemaleswere found among this group. After this, no observations were made regardingthe gravid conditionof 0+ second generationyoung in the rearingsystems.On 26 April 1991, about 300 overwintered in(1+) second generation dividuals,whichhadreached15-20-mm BL, were confirmedin the rearingsystems. Females of this 1+ second generationstarted gravidity from late May. The mean BL at season (17 June) was 22.3 mm midbreeding ? 3.3 SD (N = 133) for males, and 21.4 mm ? 2.7 SD (N = 127) for females. These sizes were almost equal to those observed at the field study site (Fig. 4). The first occurrence of thirdgeneration youngwas observedon 11 June 1991. During the following month, about500 of these young were removed. At the end of the rearingexperiment(18 October 1991), 23 females (27.0 mm ?+1.1 SD) and 37 males (32.6 mm ? 2.0 SD) of the 1+ second generation born in 1990 were foundin the rearing systems.Meansize of the 0+ third generationyoung (not sexed) was 15.3-mmBL ? 5.5 SD (N = 248), which was almostequal to thatof the 0+ field young as measured duringthe previoustwo years(Fig. 4). These resultssuggestthatgrowthrateand life cycle in the rearingsystems were essentially similarto those in the naturalpopulations, but that the reared L. cinerascens sur-

vived and reproduced longer.

760

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 19, NO. 4, 1999

Month

' /

^t--

^\
I

O
I;;M>:a^

(
~

shortened

route

breeding . main route ......

Fig. 9.

Life cycle of the shore isopod Ligia cinerascens in northern Tokyo Bay.

DISCUSSION

Some of the largest 0+ females of Ligia cinerascens, which were born at the study shore at the beginning of the breeding season (early June), started their first gravidity at the end of the same breeding season, about 2.5 months after birth, but then died before overwintering (Fig. 3). Most females, however, started gravidity after the first overwintering, then disappeared during their first breeding season. Females at the study shore can thus be assumed to experience only one breeding season before dying. Most 1+ females reached adult size (BL > 21 mm) one month after the startof the breeding season (June). The density of the adult females then rapidly decreased in July, and all had disappeared by August (Figs. 3, 6). Most of the largest adult females that started breeding from May died during June (Fig. 3). Furthermore, the 0+ breeders died about one month later (Fig. 6). Adult life time for both the 0+ and 1+ females can be estimated at about one month, during which the female can reproduce only once (Table 2). This sug-

gests that most field females may produce only one brood during their life time. Some long-lived females, however, may live long enough to produce a second brood. A great reduction in field population, as well as high mortality among reared females, were observed during the breeding season (Figs. 3, 5, 8). After the breeding season, mean body size of the surviving females in the rearing experiment was smaller than that of males, despite the fact that sizes of both sexes were similar before the breeding season (Fig. 4). These data suggest that the females invest a great amount of energy in egg production and brood care, causing critical exhaustion which may induce mortality. The life cycle of L. cinerascens at the study shore is portrayed in Fig. 9. The most typical patternis that the population is maintained by yearly generational replacement. A small number of females, however, show a shortened life cycle involving rapid growth and reproduction at the end of the breeding season in which they were born. Since these 0+ breeding females release young only at the very end of the breeding season, their young

AND ITO:LIFECYCLEOF LIGIA FUROTA

761

ACKNOWLEDGEMENTS are unable to reach adult size (21-mm BL) by the beginning of the next year's breeding seaA. Asakura, and We aregratefulto Drs. T. Udagawa, for theirhelpfulsuggestions. Dr.N. Nunoson. These offspring thus wait at least to the M. Hasegawa midbreeding season (July) to release their murakindlyidentifiedthe isopod studied.Ms. M. Itani is thankedfor her comments.Dr. A. lijima assisted in own young. These young, however, in turn do the field observations. Dr. Y. Hasuoand Ms. S. Hasuo, not have enough time to grow into 0+ breed- superintendents of the Gyotoku Wild-bird are Sanctuary, ers themselves. This pattern is similar to the thankedfor providingthe opportunity to use the study reproduction slips between 1+ and 2+ gener- site. We also thankDr. K. Shortfor editingthe English. ations observed in L. oceanica (Willows, LITERATURE CITED 1984, 1987). The one-year life cycle for L. cinerascens Carefoot,T. H. 1973a. Studies on the growth,reproduction,andlife cycle of the supralittoral isopodLigia in this study is similar to that reported for L. pallasii.-Marine Biology 18: 302-311. oceanica (L.) in the western Baltic (Jons, . 1973b. Feeding,food preference,and the uptake of food energyby the supralittoral isopod Ligia 1965), L. pallasii Brandt in British Columpallasii.-Marine Biology 18: 228-236. bia (Carefoot, 1973a), and L. dilatata Brandt Biologie und Okologie von Ligia in South Africa (Koop and Field, 1980), but Jons, D. 1965. Zur oceanica (L.) in der westlichen Ostsee.-Kieler is shorter than that reported for L. oceanica 21: 203-207. Meeresforschungen in England (Nicholls, 1931; Willows, 1984). Koop, K., andJ. G. Field. 1980. The influenceof food on population availability dynamicsof a supralittoral The shortened life cycle observed in this of Experimenisopod,LigiadilatataBrandt.-Journal study, in which females breed and die durtal MarineBiology andEcology48: 61-72. ing a single breeding season, is the first re. 1981. Energy transformation , and by the ported for species of Ligia. The study site is of supralittoral isopodLigiaditatataBrandt.-Journal at a lower latitude than previous studies (exExperimental Marine Biology and Ecology 53: 221-233. cept for South Africa), and higher summer A. G. 1931. Studieson Ligia oceanica I. A. Nicholls, temperatures may account for the fast growth Habitat andeffect of changeof environment on respirequired for the shortened life cycle. ration.B. Observation on moultingand breeding.Female L. cinerascens at the study site Journalof the Biological Associationof the United Kingdom17: 655-673. showed semelparous (one brood production N. 1983. Studies on the terrestrial isopod through a life) reproduction, most likely ne- Nunomura, in JapanI. Taxonomy crustaceans of the FamilyLigicessitated by their short adult life. The rearand Olbrinidae.-Bulletinof the idae, Trichoniscidae ing experiment, however, showed that feScienceMuseum5: 23-68. Toyama males of L. cinerascens are capable of a Sutton,S. L., M. Hassall,R. Willows, R. C. Davis, A. andK. D. Sunderland. 1984. Life histories of Grundy, longer adult life and consequently able to proterrestrial isopods:a studyof intra-and inter-specific duce up to three or even more broods during of theZoological variation.-Symposia Societyof Lona breeding season. European L. oceanica are don 53: 269-294. also essentially semelparous, but some fe- Willows,R. 1984. Breeding phenologyof woodliceand males produce a second brood (Jons, 1965; oostegite development in Ligia oceanica (L.) (Crustacea).-Symposia of the ZoologicalSociety of LonWillows, 1984, 1987). In addition, reared L. don 53: 469-485. oceanica produce two, three, or more broods . 1987. Population dynamicsand life historyof (Nicholls, 1931). A similar increase in brood two contrasting of Ligia oceanica (Cruspopulations tacea:Oniscidea)in the rocky supralittoral.-Journal production has also been reported for semelof AnimalEcology56: 315-330. parous terrestrial isopods when they were reared (reviewed by Sutton et al., 1984). This RECEIVED: 5 June 1998. iteroparous reproduction seen in reared fe- ACCEPTED:29 April 1999. males of Ligia suggests that the semelparous Address:Department of Biology,Facultyof Science, reproduction pattern observed for most females of Ligia in the field may be necessi- Toho University, Miyama 2-2-1, Funabashi, Chiba tated by external biological and/or physio- 274-8510, Japan.(e-mail:furota@bio.sci.toho-u.ac.jp) chemical conditions, rather than genetic reproductive characteristics.