JOURNAL OF CRUSTACEAN BIOLOGY, 12(1): 41-50, 1992

ASPECTS OF THE REPRODUCTION OF PANULIR US INFLATUS

(BOUVIER) AND P. GRACILIS STREETS
(DECAPODA: PALINURIDAE) FROM THE
PACIFIC COAST OF MEXICO
Patricia Briones-Fourzdn and Enrique Lozano-Alvarez
ABSTRACT
Reproductionof the tropicalspiny lobstersPanulirusinflatusand P. graciliswas studied in
Guerrero,MexicanPacificcoast, from February1979 to May 1980, from sampleswith bottomset nets and from commercialdivers'catches.Lobsterscaughtwith nets weretagged,and some
femaleswere recapturedup to 4 times duringthe studyperiod.The smallestovigerousfemales
measured49.4 mm (P. gracilis)and 54.7 mm (P. inflatus)carapacelength(CL).Reproductive
females of both species were found throughoutthe year, but analysis of the percentageof
reproductivefemales by size class showed that small lobsters (<60 mm CL) had a shorter
spawningseason and startedreproducinglater than largerlobsters(>70 mm CL). Females of
both species > 70 mm CL werecapableof producingat least 3 or 4 broodsper year,those from
60-70 mm CL up to 3, and smaller maturelobsters 1 or 2. In addition, females between 65
and 80 mm CL had the highest index of reproductivepotential,due to their highestnumbers
in the populationand their numberof broods per year. Fisheryimplicationsof these findings
are discussed.

The spiny lobsters Panulirus inflatus

(Bouvier) and P. gracilis Streets coexist
throughoutmost of the Pacificcoast of Mexico, and were confusedin the literatureuntil
Holthuis and Villalobos (1961) established
the taxonomic differences between them.
Although their distribution is similar in a
long stretch of coast, they have different

born, 1977; Briones et al., 1981), as well as

the fecundity of P. inflatus(Gracia, 1985).
In 1979-80, we conducted a taggingprogram to study the population dynamics of
both species in Guerrero,Mexico. Results
on population density and ecological notes
werepublished(Lozanoet al., 1982), as well
as some information on their fisheries (I1habitat preferences. Panulirus inflatus is re- lescas, 1981) and their stomach contents
stricted to areas of clear water and rocky (Aramoni, 1982). In this paper,we describe
bottoms, whereasP. gracilis has a tolerance reproductionin P. inflatus and P. gracilis
for a wider range of turbidity and inhabits during the taggingperiod.
both rocky and gravel-sandbottoms. Both
species exist in depths of < 1 to approxiMATERIALSAND METHODS
mately 40 m (Briones et al., 1981; Lozano
etal., 1982).
Panulirus inflatus and P. gracilis are lo-

cally importantas fisheryresources(Gracia

and Kensler, 1980); from 1981-1988 their
combined catch comprisedbetween 9.7 and
17.3%of the total spiny lobster production
in Mexico (Secretariade Pesca, 1987, 1989).
Panulirusinflatusis endemic to Mexico, but
P. gracilis,whose distributionextends south
to Peru,is also fishedin Panama(Butlerand
Pease, 1965), on the continental coast of
Ecuador (Loesch and Lopez, 1966; Baez,
1983), and in the Galapagos Islands (Holthuis and Loesch, 1967).
Population structure,sex ratio, and fisheries aspects of both species on the Mexican
Pacific coast have been described (Wein-

The study area is located in the state of Guerrero,

on the Mexican Pacific coast (Fig. 1). This coast is
characterizedby extensive cliffs with some sand and
gravelbeacheslocatedmainlyin baysandcoves (Weinborn, 1977). The bottom is rocky, with gravel-sand
patches,and muddy areasnearthe dischargeof rivers.
Surface-watertemperatureranges from 24-31?C and
salinity rangesfrom 33-35.5 %o(Brioneset al., 1981).
Specimens were obtained from February 1979 to
May 1980, with bottom-set nets installed in two experimentalzones (Fig. 1). Zone I was locatedbetween
IxtapaIslandandthe continentalcoast,and hada maximum depth of 10 m. The bottom consisted of rubble
(coarse gravel with rock heaps), rock outcrops, and
largerockformationsemergingfromthe watersurface.
In this zone, both P. inflatusand P. gracilisdwell, with
a predominanceof the former.Zone II was located 2
km northof zone I, in frontof the mouth of the Ixtapa
river. The bottom was mostly gravel-sandwith some
rubble,surroundedby sand-mudareas,and hada maximum depth of 8 m. The turbidityof the waterin zone

41

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42

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 12, NO. 1, 1992

Fig. 1. Zones I and II of study area.

II was relativelyhigher than in zone I because of the
river discharge.These characteristicsallow for the existence of P. gracilis,but not of P. inflatus.
The bottom-set nets measuredbetween 80 and 150
m long and 2 m in height.Mesh size rangedfrom 11.515 cm. Nets were installedon gravel-sandbottoms;in
zone I, they were set close to the rock heaps and formations,but not directlyon these. Lobstersweretagged
with "spaghetti"-typetags (Floy-TagFD-68B), inserted into the dorsolateralextensor muscle between the
cephalothoraxand the first abdominal segment.After
being tagged, the lobsters were immediately released
wherethey had been caught.Tag number,date, release
location, sex, carapacelength (CL, measuredfrom between the rostralhorns to the posteriordorsal edge of
the carapace, +0.1 mm), and reproductive stage of
females, accordingto the following scale (Briones et
al., 1981), were recordedfor each specimen.
Stage 1.-No extruded eggs nor traces of spermatophore.
Stage 2.-New and intact spermatophoreattachedto
the sternum.
Stage 3.-Newly extrudedeggs (brightorange).

Stage4.-Dark orangeeggs, with eye spots visible.

Stage5.--Brown eggs;eggcapsulestranslucent,embryo
and eye spots clearlyvisible.
Stage6.-Remnants of empty egg capsulesor remains
of eroded spermatophore.
The experimentalzones were fishedregularlyduring
the whole study period;this regularfishingallowedfor
multiple recaptures.In addition to specimens caught
in the nets, samples were taken from the commercial
catch obtained in nearbyareas. Fishermencatch lobsters by SCUBA and "hookah"diving, using a gaffto
extractlobstersfrom the crevicesin rocks.Fishingregulations for the two species include a closed season
from 1 June-15 September,a minimum size limit of
82.5 mm CL, and a prohibitionon the catchingof eggbearing females. However, these regulationsare not
enforced,and those regardingminimum size and eggbearingfemales are not observed by fishermen.Samples from the commercialcatch were taken in March,
April, May, October,and November 1979.
Size distributionand sex ratio, as well as monthly
percentageof reproductivefemales(reproductivestages
2-5) and of females in each reproductivestage were
obtainedfrom the pooled sample(catchfrom nets plus
commercialsamples).Durationof egg-carryingperiods
were calculatedfrom multiple-recapturedfemales.An
index of reproductivepotential (IRP; Kancirukand
Herrnkind,1976) was estimated for P. inflatus.
RESULTS

Species and Sex Ratio

The tagging program yielded a catch of
2,748 specimens, 745 (27.1%) P. inflatus,
and 2,003 (72.9%) P. gracilis. Conversely,
of the 1,335 lobsters sampled from the commercial divers' catch, 1,110 (83.1%) were
P. inflatus and 225 (16.9%) were P. gracilis
(Table 1). Sex ratio for each species of the
total tagging program and the total sample
from the commercial divers' catch showed
a smaller percentage of females in every case
(Table 1).
Size Distribution
Size distribution of the total net catch and
from samples of the commercial catch of P.
gracilis is depicted in Fig. 2. The smallest

Table 1. Speciesproportionand sex ratioof spinylobsterssampledfromcommercialdivers'catchand obtained

with nets.
males

Commercialcatch
(diving)
N= 1,335
Taggingprogram
(bottom-set nets)
N= 2,748

% of total

P. inflatus
P. gracilis

1,110
225

83.1
16.9

P. inflatus
P. gracilis

745
2,003

27.1
72.9

Females
%

Sex ratio
M:F

850
136

76.6
60.4

260
89

23.4
39.6

3.3:1
1.5:1

597
1,638

80.1
81.8

148
365

19.9
18.2

4.0:1
4.5:1

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BRIONES-FOURZAN

43

AND LOZANO-ALVAREZ: REPRODUCTION OF PANULIRUS

a,
oo

E
z

40

50

60

70
Carapace

80

90
length

100

110

120

vv D

OU

_v

.v.

130

Cmm)

F I
I Females
= IMales
IMemales
Males

Fig. 2. Size distributionof Panulirusgracilis,A) obtained with bottom-set nets, and B) caughtby divers.

specimens obtained were two females 37.1

mm CL caught in a net, while the largest
one, a male 127.0 mm CL was obtained
from the commercial catch.
Figure 3 shows the size distribution of the
total net catch and samples from the commercial catch of P. inflatus. The smallest
specimen, a male 36.2 mm CL, was obtained in a net and the largest, a male measuring 129.0 mm CL, came from the commercial catch.
Data for females from both samples were
pooled to examine the reproductive aspects
dealt with in this paper. Size distribution
for the pooled 454 females ofP. gracilis and
408 females of P. inflatus appears in Figs.
4, 5. Reproductive females (stages 2-5) accounted, respectively, for 49.1% and 51.5%
of the total.
The smallest ovigerous females obtained
in this study were 49.4 mm CL and 54.7
mm CL for P. gracilis and P. inflatus, respectively. The smallest ovigerous females
reported for both species are 45.6 mm CL
for P. inflatus (see Gracia, 1985) and 47.5
mm CL for P. gracilis (see Weinbom, 1977)
in the same area. However, the size class
where more than 30% of females were reproductive (Juinio, 1987) was 60-65 mm
CL in both species (Figs. 4, 5).

Carapace
Females

length Cmm)
= Males

Fig. 3. Size distribution of Panulirus iinflatus,A) obtained with bottom-set nets, and B) ca ughtby divers.

Spawning Season and Multiple

Spawnings
Reproductive females of P. gracilis appeared every month in a variable proportion, with a minimum of 22% in June (Fig.
6A). Females of P. inflatus were obtained
only from March-December 1979, because
tagging operations were discontinued in zone
I at the end of 1979. Reproductive females
occurred throughout this period, with the
lowest percentage in March (22%) (Fig. 6B).
Butler and Pease (1965), Loesch and Lopez
(1966), and Briones et al. (1981) reported
egg-bearing females of both species
throughout the year. The extended reproductive period indicates that multiple
spawnings occur in both species. Since
spawning and subsequent incubation of eggs
require a great expenditure of energy, this
could explain the small size of females compared to the large sizes attained by males
(Figs. 2, 3).
Briones et al. (1981) found evidence of
multiple spawnings when analyzing the gonad condition of egg-bearing females. Females in reproductive stages 3-5 had fully
mature gonads at different times of the year.
In this study, further evidence of multiple
spawnings was obtained through recaptured
females. Tables 2 and 3 show data of females of P. gracilis and P. inflatus, respec-

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44

?,

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 12, NO. 1, 1992

8-

60-

40z

E
z

20-

2040

50

60

Ovigerous
ez

80

70

Carapace

length
I

90

100

0-

Cmm)
Non ovigerous

50

60

With new spermatophore

70

Carapace

Fig. 4. Size distribution of pooled (net plus divers'

samples) of female Panulirus gracilis.

tively, that were recaptured once and from

2-4 times. In the case of P. gracilis, females
number 2 and 6 showed evidences of having
at least 3 or 4 spawnings in 265 and 153
days, while going through three and two
molts, respectively (Table 2). Similarly, female number 3 had at least 2 or 3 spawnings
and two molts in 98 days. Data from females recaptured more than twice allowed
for the estimation of the number of spawnings in those females recaptured only once.
Thus, for instance, females 16 and 17 might
have had 2 or 3 spawnings and 2 or 3 molts
in 121 and 131 days, respectively. Many of
these females molted and spawned regardless of the time of the year.
Multiple recaptures were less abundant
for P. inflatus, but a similar situation appears from Table 3. Female number 2
showed evidences of having four spawnings
and at least two molts in 164 days, while
females 1, 3, 4, and 5 all had at least two
spawnings in 61, 103, 110, and 141 days,
respectively. Few females of P. inflatus with
multiple recaptures showed growth, in contrast to females of P. gracilis.
These results indicate that females of both
species can have at least 3 or 4 spawnings
in one year, and possibly more, since multiple recaptures covered a maximum of only
265 and 164 days for P. gracilis and P. inflatus, respectively. Because several females
mated and spawned without having molted,
it is obvious that there is no need to molt
after each spawning before mating again. In
addition, 17 females of P. gracilis (65.890.4 mm CL) caught between April and

80

90

100

(mm)

length

I Non Ovigerous

Ovigerous

new spermatophore

EZWith

Fig. 5. Size distribution of pooled (net plus divers'

samples) of female Panulirus inflatus.

February, and two of P. inflatus (68.1 and

79.3 mm CL) obtained in July and October,
were found with a new spermatophore while
still carrying eggs.

A) P gracilis
24 55 33

100-

12

25

19

45

60

45

20

35

18

A S
O
MONTH

23

80-

60-

40-

20-

B) P. inflatus
39
75
100-

--

124

17

--

14

24

34

38

--

80-

600

40-

20-

Reproductive

J
A
MONTH
ZI

Non-reproductive

Fig. 6. Monthly percentage of reproductive and nonreproductive females of A) Panulirus gracilis and B)
P. inflatus. Figures above bars represent number of
females sampled each month.

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45

BRIONES-FOURZAN AND LOZANO-ALVAREZ: REPRODUCTION OF PANULIRUS

Table2. FemalesofPanulirusgracilisrecaptured(a) twice or moretimes and (b) those recapturedonce. Asterisk

indicates at least one molt occurred.

Number

Size at
first capture
(CL, mm)

Size at
recapture
(CL, mm)

Days
at large

Time of the year

Change in
reproductive
stage between
recaptures

21
4
126
85
54
48
50
70
9
10
112
5
10
17
65
61
43
38
15
9

Sep-Oct
Oct-Oct
Mar-Aug
Aug-Oct
Oct-Dec
Oct-Dec
Dec-Jan
Aug-Oct
Oct-Oct
Oct-Nov
Jul-Oct
Oct-Oct
Jun-Jun
Jun-Jul
Jul-Sep
Sep-Nov
May-Jun
Jun-Jul
Mar-Apr
Apr-Apr

1-1
1-1
1-3
3-2
2-3
5-2
2-2
3-3
3-5
5-6
6-3
3-3
6-1
1-3
3-4
4-3
3-6
6-2
3-5
5-3

Number of spawnings/
total days at large

(a)

50.9

59.6

71.6

76.1

80.3

80.6

86.2

90.1

50.9
50.9
71.7*
75.0*
78.5*
74.8*
78.0*
79.1*
79.5
79.5
83.5*
83.7
84.3*
84.5
84.3
88.0*
86.2
88.9*
90.4
90.4

0/25
3/265
2 or 3/98
2/89
1 or 2/117

3/153
2/81
2/24

Estimated
spawnings

(b)
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30

51.5
53.0
64.4
66.6
66.6
66.9
69.7
71.4
73.1
73.6
74.4
76.0
76.5
77.0
78.0
78.4
78.7
80.8
82.0
83.6
83.8
100.4

67.0*
56.3*
63.0
75.2*
71.5*
67.0
72.9*
82.7*
76.2*
73.3
80.0*
75.9
76.4
81.9*
81.5*
80.0*
78.7
81.2
82.0
88.3*
100.6

167
55
44
127
74
8
49
121
131
5
98
10
24
39
59
75
28
33
27
45
11
23

There was no significant difference (x2, P

= 0.05) in the percentage of reproductive
females among seasons in either species
(Table 4). However, differences in the percentage of reproductive females by size class
appeared among seasons (Table 5). For P.
gracilis, higher percentages of reproductive
small females (<60 mm CL) appeared in
winter (December through February), while
reproductive females in the 60-70 mm CL

Jul-Dec
Nov-Jan
Aug-Sep
Oct-Mar
Nov-Jan
Aug-Aug
Nov-Dec
Apr-Aug
Jul-Nov
Nov-Dec
Feb-May
Jun-Jun
May-Jun
Oct-Nov
Mar-May
Oct-Jan
Oct-Nov
Feb-Mar
Feb-Mar
Oct-Nov
Jul-Aug
Oct-Nov

1-3
1-3
1-3
4-5
5-6
6-6
6-3
5-6
3-2
6-6
1-3
3-4
3-6
6-4

3-3
5-5
3-6
2-1
2-6
2-1
3-5
6-2

1 or 2
1

1
2
2

1
2 or 3
2 or 3

1 or 2
1
2
2
I

1
1

class were more abundant from autumn to

spring, and females >70 mm CL reproduced during the four seasons. Data for P.
inflatus did not cover the four seasons, but
a similar scheme appears to be evident from
the three seasons represented, with large
(>80 mm CL) reproductive females peaking in autumn.
In our analysis of reproductive stage by
season (Table 6), females in stages 2, 4, and

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46

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 12, NO. 1, 1992

Table 3. Females of Panulirus inflatus recaptured (a) twice or more times and (b) those recaptured once.
Asterisk indicates at least one molt occurred.
Size at
first capture
(CL, mm)

Size at
recapture
(CL, mm)

Days at large

66.8

68.7

71.9

77.1

77.8

66.6
66.7
70.8*
72.6*
72.5
72.4
73.0*
73.0
77.5
77.5
78.0
78.7
79.0

45
16
71
65
8
20
63
40
55
55
24
89
28

Number

Time of the year

Change in
reproductive stage
between recaptures

Number of
spawnings/total
days at large

(a)
6-3
3-3
3-5
5-3
3-5
5-3
6-3
3-3
4-6
6-1
6-6
6-3
3-6

Aug-Sep
Sep-Oct
Apr-Jun
Jun-Aug
Aug-Sep
Sep-Sep
Aug-Oct
Oct-Dec
Oct-Dec
Dec-Feb
Jul-Jul
Jul-Oct
Oct-Nov

2/61

4/164
2/103
2/110

2/141
Estimated
spawnings

(b)

6
7
8
9
10
11
12
13
14
15
16
17
18
19

59.8
62.6
64.5
67.0
72.3
73.7
74.0
74.4
74.9
75.0
77.9
79.5
79.8
83.2

60.9*
62.5
65.9*
68.0
73.6*
75.0*
74.0
76.4*
77.7*
74.7
78.4
82.5*
79.8
83.4

78
9
7
54
88
80
27
73
27
28
30
47
12
12

5 were scarcer than females in other reproductive stages, indicating that the time between mating and spawning (stage 2) and
the incubation period (stages 3-5) were short.
Data from recaptured females (Tables 2, 3)
allowed for the estimation of 12 days as the
time between mating and spawning, an incubation time (stage 3 to stage 6) of 27-30
and 24-28 days, respectively, and a complete reproductive cycle (any stage to the
same stage) of 30-73 days in P. inflatus and
50-75 days in P. gracilis.
Index of Reproductive Potential
The estimation of the index of reproductive potential (IRP) was possible only for P.
inflatus, because it is the only one of the two
species for which fecundity information exists (Gracia, 1985). The index is useful to
determine which female size classes have
the highest potential larval production
(Kanciruk and Herrkind, 1976). Table 7
shows the percentage of ovigerous females,
egg-carrying capacity (Gracia, 1985), size

6-6
3-4
3-4
3-4
6-1
3-1
2-1
3-3
6-2
3-6
3-3
6-6
2-3
2-3

Jun-Aug
Sep-Sep
Sep-Sep
Aug-Oct
Sep-Dec
Feb-May
Nov-Nov
Mar-Jun
Nov-Nov
Oct-Nov
Jul-Aug
Apr-May
May-Jun
Oct-Oct

1
2
1
2
1
2
1
1
2
1
-

frequency, IRP, and contribution to total

egg production by size class.
Only 11.1% and 16.7% of females in size
classes 50-55 and 55-60 mm CL, respectively, were ovigerous (Table 7, column B).
Percentage of ovigerous females increased
until 59.6% in size class 70-75 mm CL, and
Table 4. Seasonal percentage of reproductive females
ofPanulirusgracilis(March 1979-May 1980) and Panulirus inflatus (March 1979-February 1980).

Panulirus gracilis
Spring
Summer
Autumn
Winter
Spring
Panulirus inflatus
Spring
Summer
Autumn
Winter

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% reproductive

112
56
128
71
65

50
45
51
51
55

238
40
96
11

51
48
62
55

47

REPRODUCTIONOF PANULIRUS
AND LOZANO-ALVAREZ:
BRIONES-FOURZAN

Table 5. Seasonal incidence (in percentage) of reproductive females by size class for Panulirus gracilis (March

1979-February1980) and Panulirusinflatus(March-November1979).

Size class
(CL,mm)

Spring%
reproductive

Summer%
reproductive

Autumn%
reproductive

Winter%
reproductive

0
17
46
74
62

8
5
12
20
15

0
20
25
55
60

9
11
31
57
23

0
18
45
58
70

3
13
14
32
10

33
31
43
59
60

0
38
55
56
55

1
5
13
16
6

0
20
38
63
50

1
7
27
53
10

0
29
67
58
90

Panulirusgracilis
4
<50
50-60
23
60-70
35
27
70-80
24
>80
Panulirusinflatus
1
<50
72
50-60
60-70
118
70-80
43
11
>80

remainedwithin a rangeof 36.4 and 50.0%

in the rest of the size classes. This indicates
that, although ovigerous females were relatively more abundantin the size class 7075 mm CL, the rest of the size classes had
also a largepercentageof ovigerous females
and their proportion did not increase with
size.

The IRP developed by Kanciruk and

Herrnkind(1976) assumesone spawningper
year, which is not the case for P. inflatus.

7, column D). However, by dividing the

percentageof estimatedtotal eggproduction
of size class/size class percentageof all females (F = E/A in Table 7), an index of
productivityis obtained. This indicatesthat
the highest productivity(6.3) lies in the 9095 mm CL class, which makes this size class
2.5 times more productive than the 70-75
mm CL class, and 12.6 times more productive than the newly mature 50-55 mm
CL class, emphasizing the importance of
large females for the yearly production of
eggs.

Therefore,we introducedin the calculations

the number of broods per year for each size
class considered. The value of the constant
D (1136.5) in the IRP equation was chosen
DISCUSSION
to set the index of the 70-75 mm CL size
The inverse proportion of species in
class to 100 as the standard.Resultsindicate
that females between 65 and 80 mm CL had catches made with nets and by divers is a
highervalues of IRP than females from 50- consequence of the different fishing meth65 and those largerthan 80 mm CL (Table ods employed as related to the habitats of
each species. Fishermen prefer to dive for
Table 6. Percentageof females of Panulirusgracilis lobsters in rocky bottoms where visibility
and P. inflatusfound seasonallyin each of the six re- is better, thus obtaining largerproportions
productivestagesduringthe study period.
of P. inflatus. On the other hand, the nets
aremore selective forP. gracilisbecausethis
Percentagein each
species is more abundanton the type of botreproductivestage
tom where the nets were installed (Lozano
N
1
4
2
3
5
6
etal., 1982).
Panulirusgracilis
The sex ratio in both samples favored
4
113 36
5 32
8 13
Spring
males
over females in both species. Since
7
4 16
Summer 56 39
5 29
divers searchactively for lobsters,the ratios
Autumn 128 34 16 20
7 16
8
71 35 14 24
Winter
7 14
6
from their catch are less biased than those
65 32 15 12 15 12 12
Spring
obtained with the nets. Weinbor (1977)
Panulirusinflatus
Spring 238
Summer 40
Autumn 96
Winter
11

and Briones et al. (1981) obtained a sex ratio

20
18
14
36

9
0
11
9

30
28
30
27

7
8
8
0

5
13
13
18

29
25
24
9

closer to the unity in samples taken from

divers' catches. In addition, egg-bearingfemales display a more seclusive behavior
(Kanciruk and Herrnkind, 1976; Morgan,

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48

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 12, NO. 1, 1992

Table 7. Estimation of the index of reproductivepotential (IRP) for females of Panulirus inflatus. (A =
percentageof ovigerousfemales with respectto total numberof females;B = percentageof ovigerousfemales
within a particularsize class;C = mean numberof eggsper size class;C' = C x numberbroods/year;D = IRP;
E = percentageof total eggs produced;F = E/A, i.e., an index of productivityof size class).

Size class
(CL,mm)

50-55
55-60
60-65
65-70
70-75
75-80
80-85
85-90
90-95
>95

Number
of
Total
number of ovigerous
females
females

18
24
68
89
99
64
28
11
2
2

2
4
23
39
59
30
13
4
1
0

0.5
1.0
5.6
9.6
14.5
7.4
3.2
1.0
0.2
0.0

11.1
16.7
33.8
43.8
59.6
46.9
46.4
36.4
50.0
0.0

C(x103)

Number
broods/
year

129.4
168.2
214.1
267.4
328.8
398.6
477.6
566.1
664.7
774.0

1980; Lozano et al., 1982). Egg-bearingfemales ofP. inflatustend to occupy the deepest partsof crevices, which makes them less
accessibleto divers (E. Lozano,personalobservation).The sex ratio of P. gracilistaken
by divers is less biased, because the habitat
of this species is less intricate than that of
P. inflatus,and females are more easily detected by divers. In contrast, when using
nets, the lobstersplay the active role. Males
are more active than females and thus are
more abundant in catches made with nets.
This type of behaviorhas also been reported
in catches from traps (Newman and Pollock, 1971; Davis, 1977). Kanciruk(1980)
and Morgan (1980) concluded that disproportionate sex ratios reportedin spiny lobsters are probably due to differencesin female and male behavior.
Protracted spawning seasons, implying
multiple spawnings, are a current issue in
tropicalpalinurids(Berry,1970, 1971;Kanciruk, 1980; MacDonald, 1982; Juinio,
1987). Females ofPanulirus inflatusand P.
gracilis have multiple broods per year. In
addition, females >70 mm CL spawn
throughout the year, while those between
60 and 70 mm CL have a shorterspawning
season, with a minimum in summer, and
females < 60 mm CL have only one spawning peak in winter.
In the study area, water temperatureis
29-31?C in summer, 28-29?C in autumn,
26-27?C in winter,and 24.4-26?C in spring
(Briones et al., 1981). Organicmatter, wet
weight of molluscs (the main diet component) (Aramoni, 1982), and density of lobsters (Lozano et al., 1982) are higher in au-

2
2
3
3
4
4
4
4
4
4

C'(xl03)

258.8
336.4
642.3
802.2
1,315.2
1,594.4
1,910.4
2,264.4
2,658.8
3,096.0

0.1
0.5
10.7
29.7
100.0
48.7
25.0
7.3
2.3
0.0

Number
eggs
produced/
year(xl03)

517.6
1,345.6
14,772.9
31,258.8
77,596.8
47,832.0
24,835.2
9,057.6
2,658.8
0.0

0.2
0.6
7.0
14.9
37.0
22.8
11.8
4.3
1.3
0.0

0.5
0.6
1.3
1.6
2.5
3.1
3.7
4.3
6.3
-

tumn in zone I, which indicates that

conditions are optimal in this season for
reproductive success. First-stage phyllosomata densities are also higher in September-October (Lozano and Briones, unpublished data). Gracia (1985) found that
fecundityof females ofP. inflatuswas higher and the mean size of the eggs was significantlylargerin autumnthanin spring.Thus,
the autumn spawning seems to be the first
of the yearfor most of the medium and large
females, at least for P. inflatus, with subsequent spawningsbeing less abundant, as
has been reportedfor P. argus (see Creaser,
1950), P. japonicus (see Ino, 1950), and P.
ornatus (see MacFarlane and Moore, 1986).

Berry(1971) found that large females of P.

homarus rubellus in South Africa spawn
from winter to early autumn, and small females from late springto late summer. Lipcius (1985) found a similar situation for P.
argus in Florida, with largefemales spawning earlier(in early spring)than smaller females (in summer), which grew in early
springratherthan reproducing.Since molting and reproductionare coordinated(Lipcius and Herrnkind,1987), a similar explanation can be offered for P. gracilis, with

smaller lobsters probably growing in autumn and spawning in winter, as opposed

to largerlobsters who start spawningin autumn.
When the time to complete a reproductive cycle (Tables 2, 3), and the evidence
obtainedthroughrecapturesareconsidered,
females >70 mm CL of both species are
probably capable of producing up to five
broods in one year; females 60-70 mm CL

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BRIONES-FOURZAN AND LOZANO-ALVAREZ: REPRODUCTION OF PANULIRUS

up to four, and those smaller than 60 mm

CL from one to three in one year. This emphasizes the fact that females >70 mm CL,
in addition to having the highestIRP values
(Table 7), are more relevant for egg-production than their numbers would suggest.
The only other palinurid species for which
evidence of as many as four broods per year
has been found are P. homarus rubellusin
South Africa (Berry, 1971) and P. penicillatus in the Philippines (Juinio, 1987).
Female P. argusin captivity that had successfully copulated avoided the approaches
of courting males and even responded aggressively (Lipcius and Herrnkind, 1985).
However, the finding of several females of
P. gracilisand a few P. inflatusshowingboth
extrudedeggs and a new spermatophoreindicates that a second mating took place before the release of larvae from the previous
brood. This also has been recorded for P.
argusin South Florida(Gregoryet al., 1982)
and for P. ornatus in Papua New Guinea
(MacFarlaneand Moore, 1986), suggesting
that it is not an uncommon event in tropical
palinuridswith repetitive breeding.
The proportionof lobsterslargerthan the
currentminimum legal size (82.5 mm CL)
is small (Figs. 2, 3). This minimum size was

49

help in field and laboratoryactivities, and Benjamin

Sotelo, Alfredo Vargas,and Simbad Vargasfor their
help in samplingactivities. Commercialsampleswere
obtained from catches of the Soc. Coop. "Vicente
Guerrero,"S.C.L. This projectwas supportedby the
Instituto Nacional de la Pesca (Secretariade Pesca,
Mexico) and UNAM.
LITERATURECITED

Aramoni,G. P. 1982. Alimentaci6nde las langostas

Panulirusinflatus(Bouvier)y P. gracilis Streetsen
Zihuatanejo,Guerrero,y su relaci6ncon el bentos.Tesis Profesional,Facultadde Ciencias,Universidad
Nacional Aut6noma de Mexico. Pp. 1-66.
Baez, P. 1983. Larvasphyllosoma y puerulusde la
langostaverde PanulirusgracilisStreets, 1871, procedentes de la expedici6n Costa Rica 1973 (Crustacea:Decapoda:Palinuridae).-Revista de Biologia
Marina,Valparaiso(Chile) 19: 79-111.
Berry,P. F. 1970. Matingbehaviour,ovipositionand
fertilizationin the spiny lobsterPanulirushomarus
(Linnaeus). -Oceanographic Research Institute
(Durban)InvestigationalReport 24: 1-16.
1971. The biology of the spiny lobsterPanulirus homarus (Linnaeus) off the east coast of
Southern Africa.-Oceanographic Research Institute (Durban)InvestigationalReport 28: 1-75.
Briones, P., E. Lozano, A. Martinez, and S. Cortes.
1981. Aspectos generalesde la biologia y pesca de
las langostas en Zihuatanejo,Gro., Mexico (Crustacea: Palinuridae).-Anales del Instituto de Ciencias del Mar y Limnologia Universidad Nacional
Aut6noma de Mexico 8: 79-102.
Butler,J. A., and N. L. Pease. 1965. Spiny lobster
explorationsin the Pacificand Caribbeanwatersof
the Republic of Panama.-United States Fish and
set for P. interruptus in Baja California, a
Wildlife Service, Special ScientificReport on Fishthat
reaches
subtropicalspecies
largersizes,
eries 505: 1-26.
and it seems to have been extended to P.
Chapa, H. 1964. Contribuci6nal conocimiento de
inflatus and P. gracilis before any studies on
las langostasdel Pacificomexicanoy su pesqueria.the size distributionof these two specieshad
InstitutoNacionalde InvestigacionesBiol6gico-Pesbeen undertaken(Chapa, 1964). However,
queras,Secretariade Industriay Comercio(Mexico),
Publicaci6n6: 1-68.
this regulation is not enforced. Fishermen
E. P. 1950. Repetition of egg-layingand
Creaser,
empirically assume that the minimum size
numberof eggsof the Bermudaspiny lobster.- Prolimit is incorrectfor these two species, and
ceedingsof the Gulf and CaribbeanFisheriesInstitute 2: 30-31.
do not observe it at all, with the conseEffectsof recreationalharveston
quence that fishing is exerted on the pop- Davis, G. E. 1977.Panulirus
a spiny lobster,
argus, population.-Bululation from very small sizes. This action
letin of MarineScience 27: 223-236.
could endangerthe fishery,and, in fact, be- Gracia, A. 1985. Variaci6nestacional en la fecuntween 1976 and 1980 the mean size of the
didadde la langostaPanulirusinflatus(Bouvier,1895)
lobster catch decreased significantly(Illes(Crustacea:Decapoda:Palinuridae).- CienciasMa11: 7-27.
et
Briones
al.
cas, 1981).
(1981) suggested rinas, (Mexico)
and C. B. Kensler. 1980. Las langostasde
a reduction in the minimum legal size to
Mexico: su biologia y pesqueria.-Anales del Insti75.0 mm CL because 50%of the population
tuto de Cienciasdel Mary Limnologia,Universidad
Nacional Aut6noma de Mexico 7: 111-128.
of both species was below that size and females startedreproductiveactivities at ap- Gregory,D. R., Jr., R. F. Labisky,and C. L. Combs.
1982. Reproductivedynamics of the spiny lobster
proximately 50 mm CL. The results of the
Panulirusargus in South Florida.-Transactions of
presentstudysupportthis suggestion,as long
the AmericanFisheriesSociety 111: 575-584.
as the new measurementis enforced.
Holthuis, L., and H. Loesch. 1967. The lobsters of
the Galapagos Islands (Decapoda, Palinuridea).ACKNOWLEDGEMENTS
Crustaceana12: 213-222.
The authors thank Carlos Illescas, Leonardo San, and A. Villalobos. 1961. Panulirusgracilis
Streets y Panulirusinflatus (Bouvier),dos especies
tarelli,Adolfo Gracia,and GabrielaAramonifor their

This content downloaded on Tue, 29 Jan 2013 20:47:36 PM

All use subject to JSTOR Terms and Conditions

50

JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 12, NO. 1, 1992

de langosta (Crustacea,Decapoda) de la costa del

Pacificode Am6rica.-Anales del Institutode Biologia, UniversidadNacionalAut6nomade Mexico 32:
251-276.

Illescas,C. M. 1981. Pescaexperimentaly aportaci6n

al conocimientobiol6gicode las langostasPanulirus
inflatusy Panulirusgracilisen Zihuatanejo,Gro.Tesis Profesional,Facultadde Ciencias,Universidad
Nacional Aut6noma de Mexico. Pp. 1-81.
Ino, T. 1950. Observationson the spawningcycle of
the Ise-ebi, Panulirusjaponicus(V. Siebold).-Bulletin of the JapaneseSociety of ScientificFisheries
15: 725-727.

Juinio, M. A. R. 1987. Some aspects of the reproductionof Panuliruspenicillatus(Decapoda:Palinuridae).-Bulletin of MarineScience 41: 242-252.
Kanciruk,P. 1980. Ecologyofjuvenile and adultPalinuridae.--In: J. S. Cobb and B. F. Phillips, eds.,
The biology and managementof lobsters. Vol. 2.
Ecology and management. Pp. 59-96. Academic
Press, New York, New York.
, and W. F. Herrnkind. 1976. Autumnal reproduction in Panulirus argus at Bimini, Bahamas.-Bulletin of MarineScience 26: 417-432.
Lipcius, R. N. 1985. Size-dependentreproduction
and molting in spiny lobsters and other long-lived
decapods.--n: A. Wenner, ed., Crustaceanissues.
Vol. 3, Crustaceangrowth:Factorsin adult growth.
Pp. 129-148. BalkemaPress,Rotterdam,The Netherlands.
,and W. F. Herrnkind. 1985. Photoperiodic
regulationand daily timing of spiny lobster mating
behavior.-Journal of ExperimentalMarineBiology
and Ecology 89: 191-204.

. 1987. Controland coordination

,and
of reproductionandmoltingin the spinylobsterPanulirusargus.-Marine Biology 96: 207-214.
Loesch,H., and E. Lopez. 1966. Observacionessobre
la langosta de la costa continental del Ecuador.-Boletin Cientificoy T6cnico, Instituto Nacional de
Pesca (Ecuador)1(5): 1-30.
Lozano, E., P. Briones, L. Santarelli,and A. Gracia.
1982. Densidad poblacional de Panulirusgracilis
Streets y Panulirus inflatus (Bouvier) (Crustacea:
Palinuridae)en dos areas cercanas a Zihuatanejo,

Gro., Mexico.-Ciencia Pesquera,Instituto Nacional de la Pesca Secretariade Pesca (M6xico)3: 6173.

MacDonald,C. D. 1982. Catchcompositionand reproductionof the spiny lobsterPanulirusversicolor

at Palau.-Transactions of the American Fisheries
Society 111: 694-699.
MacFarlane,J. W., and R. Moore. 1986. Reproduction of the ornate rock lobster, Panulirus ornatus
(Fabricius), in Papua New Guinea.-Australian
Journalof Marineand FreshwaterResearch37: 5565.
Morgan,G. R. 1980. Populationdynamics of spiny
lobsters.--In:J. S. Cobband B. F. Phillips,eds., The
biologyand managementof lobsters.Vol. 2. Ecology
and management.Pp. 59-96. AcademicPress,New
York, New York.
Newman, G. G., and D. E. Pollock. 1971. Biology
and migrationof rocklobsterJasus lalandiiandtheir
effecton availabilityat ElandsBay, South Africa.South African Division of Sea Fisheries Investigational Report 94: 1-24.
Secretariade Pesca. 1987. PesqueriasMexicanas:Estrategiasparasu administraci6n.-Direcci6n General de Administraci6nde Pesquerias,Secretariade
Pesca (Mexico). Pp. 1-1061.
SecretariadePesca. 1989. AnuarioEstadistico1988.Direcci6nGeneralde Informatica,Estadisticay Documentaci6n,Secretariade Pesca (Mexico). Pp. 1350.
Weinborn,J. A. 1977. Estudiopreliminardela biologia, ecologiay semicultivode los Palinuridosde Zihuatanejo,Gro., Mexico, Panulirusgracilis Streetsy
Panulirusinflatus(Bouvier).-Anales del Centrode
Ciencias del Mar y Limnologia, Universidad Nacional Aut6noma de Mexico 4: 27-78.
24 May 1991.
RECEIVED:

ACCEPTED:
4 September1991.
Address:UniversidadNacionalAut6nomade M6xico, Instituto de Ciencias del Mar y Limnologia,Estaci6n "Puerto Morelos," Ap. Postal 1152, Cancuin,
QuintanaRoo, 77500 Mexico.

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