Marine Biology 114, 109-117 (1992) Marine ,===,Biology @ Springer-Verlag 1992
Population characteristics of the deep-sea lobsters Polycheles typhlops and Stereomastis sculpta (Decapoda: Polychelidae) in a bathyal mud community of the Mediterranean Sea
P. Abell6 and J. E. Cartes Institut de Ci6ncies del Mar, Passeig Nacional s/n, E-08039 Barcelona, Spain Date of final manuscript acceptance: April 24, 1992. Communicatedby J. M. Pdr6s, Marseille
Abstract. Distribution and abundance as a function of Introduction depth, and population parameters such as sex and size structure of the population and aspects of reproductive Polychelid lobsters are truly deep-water decapod crus- biology have been studied in the deep-sea polychelid lob- taceans whose life habits and ecology are still far from sters Polycheles typhlops and Stereomastis sculpta. Sam- being well understood. Little information is available on ples were taken by otter trawl in the Balearic Sea, a deep- their biology and ecology (Lagard6re 1977, Wenner 1979, sea basin in the northwestern Mediterranean Sea from Wenner and Boesch 1979, Wicksten 1980, Gore 1984). 1985 to 1989. In both species the largest size classes were Many difficulties are inherent to deep-sea sampling, and dominated by females. In S. sculpta, male and female the rarity of scientific cruises studying the bathyal and sizes were very similar. In P. typhlops, ovigerous females abyssal megafauna in many areas results in a lack of and those with external spermatophores were >23 and knowledge of the population biology and ecology of most 25 mm in carapace length (CL), respectively; males with deep-sea species. The limited number of captures of spec- external spermatophores were >17 mm in CL. In S. imens of some species of the Polychelidae family also sculpta, ovigerous females and those with external sper- accounts for the lack of knowledge about their general matophores were > 24 and > 19 mm CL, respectively, biology. and males with external spermatophores > 19 mm CL. Four different types of polychelid larvae are known to Highest densities of P. typhlops occurred along the mid- occur in the Mediterranean (Fredj and Laubier 1985); dle slope at depths between 500 and 1000 m. Only small- however, only two species of adult polychelids are known sized individuals were found at the deepest depths sam- from this sea: Polycheles typhlops Heller, 1862 and pled; some recruitment must therefore occur in waters Stereomastis sculpta (S.I. Smith, 1880). P. typhlops has much deeper than those usually inhabited by the adult been previously reported in the study area from the upper population. The sex-ratio was 1:1 in most samples, but in slope at ~ 300 m down to depths of ~ 2000 m (Zariquiey- some of the shallowest samples females predominated. Alvarez 1968, Abell6 and Valladares 1988), and S. sculpta The depth distribution range of S. sculpta was 981 to has been recorded at depths between 1196 and 2011 m 2253 m; densities clearly increased with increasing depth. (Abell6 and Valladares 1988). P. typhlops is a characteris- There were no apparent variations in size distribution as tic species of the middle subzone of the Mediterranean a function of depth. Since very few adult males and fe- bathyal mud assemblage, and S. sculpta is characteristic males were captured, the population in the survey area of the lower subzone (P6r6s 1985, Abell6 et al. 1988). would seem to be comprised mainly of juveniles. As a Some aspects of the biology and ecology of Polycheles function of depth, females were numerically dominant typhlops were firstly studied in the Ligurian Sea by San- only in some of the shallowest samples taken in the distri- tucci (1933). Additional data on its distribution and com- bution range of this species. There is strict habitat parti- munity ecology are rather scattered in the literature (e.g. tioning between the two species. In both species, the vari- Carpine 1970, Relini-Orsi and Relini 1972, Arena and Li ation in the sex-ratio as a function of depth suggests Greci 1973, Relini et al. 1986, Mura 1987, Abell6 et al. differential migration between the sexes, probably related 1988). No previous information on the biology and pop- to egg incubation and hatching. ulation ecology of Mediterranean Stereomastis seulpta is available. Some preliminary results on feeding ecology and size-distribution patterns of P. typhlops and S. sculp- ta were presented by Cartes and Abell6 (1990). Despite belonging to different genera, both species are very similar in morphology and size. Both probably occupy 110 P. Abell6 and J.E. Cartes: Ecology of polychelid lobsters
Polycheles fyphlops
..Q n:241 Po/ycheles fyphlops. E Males "1 2O I ~ ~ -~ ~K~<~< Z ~?14~?14~'''14~-~ ~.~ 0 E 0.75
r 0 0 0 0 O0 O 0 0.5 ...... Q ...... Q ...... e ...... o 20 /3- 000 O0000 0 0.25 / 0 NS
40 p Sfereomaslis sculpfo Stereomasfis scu/pfa 100 t, n=845 t ..O Males E 2:50 Ovigerous ~% .14 00 [-~ Females ~0.75 o o~ 0.5 ...... o o la. o o 5O 0.25 o NS n=1053 ,1~ p very similar ecological niches. However, as yet, little is Table 1. Trawls at 200 m depth-intervals used for present study in known about their population ecology. bathyal basin of Catalan Sea (no trawls were made between 800 and 1 000 m). Mean between maximum and minimum depth of each trawl was used to assign each trawl to a depth interval Depth range (m) No. of trawls Materials and methods 200- 400 26 The study area comprised part of the bathyal valley of the Catalan 400- 600 22 Sea, between the continental margins of the coasts of Catalonia and 600- 800 8 the Balearic Islands, between Latitudes 39~ and 42~ and 1 000-1 200 13 Longitudes 1~ and 3~ This area belongs to the Western 1 200-1 400 7 Mediterranean bathyal basin. 1 400-1 600 7 Samples were obtained from several research cruises performed 1 600-1 800 12 in 1985-1989 and aimed at the faunistic study of the epibenthic 1 800-2 000 11 macrofauna inhabiting the deep Mediterranean bathyal basin. Sam: 2 000-2 200 7 ples were taken using deep-water otter-trawls equipped with a 6 mm mesh-size cod-end. All trawls were performed during the day; 113 113 trawls were examined in the present study (Table 1). Abundances P. Abell6 and J.E. Cartes: Ecology of polychelid lobsters 111 Polycheles typhlops 2O Males 15 [ ~ with spermatophores J JD F10 -j Z t J [ q i i i i k r i t i i ~ t i i i i E r i i i 5 10 15 20 25 30 35 40 45 50 14 12 Non-ovigerous females 10 [ m with IoermatoDhores Z 4 5 10 15 20 25 30 35 40 45 50 Ovigerous females I e~ with spermatophore~ -i n=57 Z Fig. 3. Polycheles typhlops. Size-frequency distributions of males and of non-ovigerous and ovigerous females with external spermatophores irJl,,,, 5 10 15 20 25 30 35 40 45 50 CL(mm) were standardized to number of individuals captured per hour of Results trawling (individuals/h). A total of 736 individuals of Potycheles typhlops and 1896 Stereomastis sculpta were examined. Sex, size (carapace length, CL, Size dimorphism as millimeters from the median posterior margin of the carapace to the end of the rostral spines), and presence of external eggs and The overall size-frequency distribution recorded for both spermatophores were noted. Polycheles typhlops and Stereomastis sculpta (Fig. 1) indi- The sex ratio (as proportion of females in each sample) was calculated in those trawls where ten or more individuals were cap- cates that very few males attain the sizes reached by fe- tured. A likelihood-ratio G-test using Williams' correction (Sokal males; the proportion of females is very high at sizes and Rohlf 1981) was performed to test the deviations of the sex ~> 30 mm CL. In P. typhlops, males ranged between 9 and ratios recorded for each sample using a theoretically expected ratio 46 mm CL in size, females between 9 and 49 mm CL. In of 1:1. Deviations in sex ratio as a function of size were tested using S. sculpta, female sizes were generally very similar to a heterogeneity G-test (Sokal and Rohlf 1981). those of males; males ranged between 8 and 35 mm in Size-frequency distributions are presented by 200 m depth inter- carapace length, and females between 5 and 46 ram. In vals, except in the deepest distribution range of Polycheles typhlops where (in view of the scarceness of this species) samples were both species, both sexes displayed a clearly marked poly- grouped into larger depth intervals of 1100 to 1600 m and 1600 to modal size-frequency distribution. In P. typhlops, the 2300 m. largest sizes (> 30 mm CL) were clearly dominated by 112 P. Abell6 and J.E. Cartes: Ecology of polychelid lobsters Stereomastis sculpta 120 Males 100 8O I ~ with Ipm'matophore8 t .0 E 60 =I n=843 Z 4O l 20 t ~ (i t i i i i i t i i i I=lll~LIJIIIk~lll 5 10 15 20 25 30 35 40 45 50 120 Non-ovigerous females 100 [ ~ with spermatophores 8o n=1o24 J~ E 60 Z 4O 20 5 10 15 20 25 30 35 40 45 50 Ovigerous females 5 ~ with $permatophore= I 4 J~ n=27 E -1 3 Z Fig. 4. Stereomastis sculpta. Size-frequency distributions of males and of non-ovigerous and ovigerous females with external spermatophores 5 10 15 20 25 30 35 40 45 50 CL (mm) ovigerous females. The size distribution observed in S. 200-400 however, suggests that very few of the females sculpta, E 400-500- captured were adults. In Polyeheles typhlops, the sex ratio plotted as a func- 600-800 tion of size (Fig. 2) did not differ significantly from 1:1 at 800-1000 sizes ~< 25 to 30 mm CL; it then increased steadily until, 1000-1200 at >40 mm CL, almost 100% of the population was 1200-1400 comprised of females. Stereomastis sculpta displayed ap- P. typhlops 1 I S. scu,#to proximately the same pattern as P. typhIops; the sex ratio 1400-1600 of this species did not differ significantly from 1:1 at 1600-1800 %25 mm CL, and then steadily increased up to 100% at 1800-200@ > 35 mm CL. 2000-2200- i _ '0 20 40 Mean abundance (ind/h) Reproductive biology Fig. 5. Polycheles typhlops and Stereornastis seulpta. Bathymetric distribution and mean abundance (no. of individuals per hour of The size range of ovigerous female Polycheles typhlops trawling) in bathyal valley of Catalan Sea. Samples are grouped into lay between 23 and 49 mm CL (Fig. 1). Sizes of females 200 m depth-intervals. (No trawsl were made between 800 and with spermatophores ranged between 25 and 44 mm CL 1 000 m) P. Abell6 and J.E. Cartes: Ecology of polychelid lobsters 113 Polyche/es Iyphlops n=16 10 300-500 m 10~ n=65 ~ 1900-1t00 rn I k~ ..(3 .13 E LJ M J u E z z 10 10] u u U~JLY l 20 n=40 20 ~109 ..... I ...... I ...... I ...... 1,, 5 15 25 35 45 5 15 25 55 45 CL (mr'n) CL (mrn) 10 t n=102 1.500-700 m I 10 n=16 [1100--1600 m l .Q Z3 _ k~ ~r~ r~k~ #~ E E I~ ~Llj~u='~ HJ~UH~ z Z 10 20 In=239 20 n=29 I,~lE,~r,ll,4L,lCE ~I,,E,I,,I,t~I,~, +Ill III III ~tll I~ I] ,~ ]1 i kl t $1 i i I I I I I I ~ I I I I I 5 15 25 35 45 5 15 25 35 45 CL (mr'n) C/ (turn) n=37 n=9 10 700-900 m o; 1600-2300 m E LJU z Z ~ Males 10. 10 Ovigerous 20. n=56 20 n=18 [--[ Females ..... ,,,I ...... I ...... I ...... I .... , t 5 25 35 45 5 15 25 35 45 CL (~m) CL (mr-n) Fig. 6. Polycheles typhlops. Size-frequency distribution as a function of depth intervals in bathyal valley of Catalan Sea. CL: carapace length (Fig. 3). Males carried external spermatophores from an Distribution and abundance earlier size than females (between 17 and 32 mm CL). Most ovigerous females bore external spermatophores. Polycheles typhlops was found between ,-,300 and Ovigerous female Stereomastis sculpta were scarce in ,-,1900 m depth throughout the survey area and was the samples, and ranged in size between 24 and 44 mm present in both the continental slope and the bathyal CL (Fig. 1). Females with spermatophores ranged be- basin. However, the highest densities were recorded tween 19 and 46 mm CL in size; males with external sper- along the middle slope between 600 and 800 m depth matophores between 19 and 35 mm CL (Fig. 4). As in (Fig. 5); no trawls were made at a mean depth between Polycheles typhlops, most ovigerous females also bore 800 and 1000 m (Table 1); below 1000 m, in the bathyal external spermatophores. basin, densities were fairly constant but rather low. 114 P. Abell6 and J.E. Cartes: Ecology of polychelid lobsters Stereomaslis scu/pta 40 n=6 ]1200--1400 m n:294 m ~ 1[]_800-2000 rnll 2O 2O k- '~l J3 E ~E '7 Males 2O 20 Ovig, females 40 n=5 Females 40 t n=332 ...... I ...... I ...... ,I ...... I,,, 5 15 25 .55 45 5 15 25 55 45 CL (ram) CL (mm) 40 40 n=22 [ 1400--1600 m n=95 [2000-2200 m] 20 20 _(3 c~ E E z z 20 20 40 n=76 40 n=99 ...... I ...... i ...... I ...... I , ,,~- 5 15 25 35 45 5 15 25 35 45 C/ (turn) CL (mm) 40- n=269 ~ N [1 600-1'800 m-7 40 n=160 22~0_0--2300m] 20. 2O ..(3 t , E -~ ~,~ z z 20t 2O 40- n=381 40 n=159 I, .... ,,,,I,,,,, .... I,,,,,,,,,I,,,,,,,, 5 15 25 35 45 5 15 25 .55 45 CL (mm) CL (rnm) Fig. 7. Stereomastis sculpta. Size-frequency distribution as a function of depth intervals in bathyal valley of Catalan Sea. CL: carapace length Stereomastis sculpta inhabits the bottom of the West- Size distribution as a function of depth ern Mediterranean bathyal basin. The deepest bathymet- tic distribution limit of this species has, therefore, not Fig. 6 shows size-frequency distributions as a function of been discovered. First occurrences were recorded at sex and depth for Polycheles typhlops. Only small-sized 1000 m. The depth range for this species in the surveys individuals were found at the deepest depths sampled. was 981 to 2253 m. Densities clearly increased with The highest proportion of ovigerous females was record- depth, attaining maximum values in the bathyal basin at ed for the shallowest levels of the bathymetric distribu- around 2000 m depth (Fig. 5). tion range of the species (300 to 700 m). P. Abell6 and J. E. Cartes: Ecology of polychelid lobsters 115 ,Pol/cheles typhlops portion of all females (Fig. 6), this suggests reproductive migrations related to egg laying and/or spawning. 20O In Stereomastis sculpta, sex ratio as a function of E O depth revealed that females tended to comprise most of 5O0 .X2 the population in the shallowest depths of the species' Q- distribution range (Fig. 8). Nevertheless, the sex ratio did C3 800 0 not differ significantly from 1:1 in most samples; howev- q~ er, the few ovigerous females in the samples were scat- 1100 D U [] tered among all the depth strata surveyed (Fig. 7). 1400 I [] NS Discussion 1700 Little information is available on the biology and ecology l D of polychelid lobsters. The difficulties of adequately sam- 2000 i : i 0 0.2 0.4 0.6 0.8 pling their bathyal and abyssal habitat account for the Sex ratio scarcity of knowledge of the deep-sea fauna. The present survey provided the opportunity of studying a few as- pects of the biology and ecology of both Polycheles ty- Slereomosti7 sculpto phlops and Stereomastis sculpta. Despite a few early stud- ies on P. typhlops (e.g. Santucci 1933), little is known of 1400 the biology of this deep-water lobster, while previous E ra [] data on the life history of S. sculpta in the Mediterranean "'1600 was completely lacking. The patterns of size dimorphism of the two species ODD D [] differ markedly from those observed for many benthic C3 1800 D D decapods, in which males usually reach larger sizes than females (Wenner 1972). In both Polycheles typhlops and 2000 [] Stereomastis sculpta, females attained larger sizes than D males. The size ranges reported by Mura (1987) for P. o NS 2200 [] [] typhlops and by Abell6 and Valladares (1988) for both p In Stereomastis sculpta, ovigerous females and those area constitutes only a part of the extensive (and in some with attached spermatophores occurred from ~24 mm sections much deeper) bathyal basin of the western Med- and from ~ 19 mm CL, respectively. In males, external iterranean, the adult population may occupy mainly the spermatophores occurred from ~19 mm CL. All these deeper parts of the basin, and the juveniles the bottom of sizes are much smaller than those reported by Wenner the Catalan Sea. (1979) for the western North-Atlantic S. sculpta popula- As a function of depth, the sex ratio displayed a simi- tion. In comparison with Polycheles typhlops, a Very lar pattern in both species. The greater part of the females small number of ovigerous female S. seulpta was cap- occurred in the shallowest depths of the species' distribu- tured. tion range, suggesting differential population movements Sex ratio as a function of size suggested differential between the sexes probably related to egg incubation and growth between the sexes: from ~25 to 30 mm CL in hatching, as suggested by Santucci (1933). For Polycheles typhlops, and from ~ 25 mm CL in Stereomas- Stereomastis sculpta, Wenner (1979) reported that most tis sculpta. The size at which differences in the sex ratio ovigerous females occurred in the shallowest depths of become significant has been suggested to be an indirect the species' distribution range; this pattern has also been indicator of size at the puberty moult (Abell6 et al. 1990). documented for other deep-water decapods such as litho- In both P. typhlops and S. sculpta this size closely match- did crabs (Abell6 and Macpherson 1991). es the size at which external eggs and spermatophores appear. The size-related sex ratio did not follow any of Acknowledgements. We wish to thank our colleagues at the Institut the patterns indicated by Wenner (1972). de CiSncies del Mar de Barcelona, and especially to all members of the research projects BATIMAR and AQUDE, as well as Drs. E One of th~ most remarkable features concerning the Sardfi, E. Macpherson and M. Demestre for their helpful com- comparative distribution of Polycheles typhlops and ments. Stereomastis sculpta lies in the low degree of habitat over- lap between these two deep-water lobsters. P. typhlops was caught within a wide depth zone throughout the survey area. The highest densities of P. typhlops were on Literature cited the continental slope between ~ 400 and 1000 m, whereas S. sculpta was collected from the deep bathyal basin, with Abellt, P., Macpherson, E. (1991). Distribution patterns and mi- highest densities occurring deeper than ~1600 m. A gration of Lithodes ferox (Filhol) (Anomura: Lithodidae) off Namibia. J. Crustacean Biol. 11:261-268 somewhat similar distribution pattern, with congeneric Abel16, P., Pertierra, J. P., Reid, D. G. (1990). Sexual size dimor- species showing overlapping depth range but with differ- phism, relative growth and handedness in Liocarcinus depurator ent depths of maximum abundance, has been demon- and Macropipus tuberculatus (Brachyura: Portunidae). Scientia strated for S. sculpta and S. nana in the northwestern mar. 54:195-202 Atlantic by Wenner and Boesch (1979). Abell6, P., Valladares, K J. (1988). Bathyal decapod crustaceans of The patterns of size distribution as a function of depth the Catalan Sea (northwestern Mediterranean). Mtsogte 48: 97-102 revealed different characteristics for the two species in the Abell6, P., Valladares, E J., Castell6n, A. (1988). Analysis of the present study. In Polycheles typhlops, some recruitment structure of decapod crustacean assemblages off the Catalan takes place in waters much deeper than those usually coast (North-West Mediterranean). Mar. Biol. 98:39-49 inhabited by the adult population: the smallest individu- Arena, P., Li Greci, F. (1973). Indagine sulle condizioni faunistiche als were found in the deepest sampled levels, where very e sui rendimenti di pesca dei fondali batiali della Sicilia occiden- few adult lobsters were caught. Environmental stability is tale e della bordura settentrionale dei banchi della soglia Siculo- Tunisina. Quad. Lab. Tecnol. Pesca 1:157-201 usually considered to increase with depth; therefore, Attrill, M. J., Hartnoll, R. G., Rice, A. L., Thurston, M. H. (1990). adult P. typhIops would appear to inhabit more unstable A depth-related distribution of the red crab, Geryon trispinosus habitats than juveniles. Competition between adult P. (Herbst) [= G. tridens Kroyerl: indications of vertical migration. typhlops and Stereomastis sculpta may also account for Prog. Oceanogr. 24:197-206 this pattern. Beyers, C. J. de, Wilke, C. G. (1980). Quantitative stock survey and some biological and morphometric characteristics of the deep- The pattern in Polycheles typhlops is clearly different sea red crab Geryon quinquedens off south west Africa. Fishery from that of benthic littoral and shallow-water decapods Bull. S. Afr. 13:9-19 which inhabit the continental shelf. In these species, juve- Carpine, C. (1970). Ecologie de l'~tage bathyal dans la Mtditerra- niles tend to occur in shallow water or at lesser depths nte occidentale. Mtm. Inst. octanogr. Monaco 2:1-146 than those inhabited by the adults, and favour more en- Cartes, J., Abell6, P. (1990). Comparative size distribution and ergetic, biologically productive and unstable waters. feeding ecology of Polycheles typhlops and Stereomastis sculpta (Decapoda, Polychelidae) in the Mediterranean bathyal mud Some deep-water crustaceans do, however, sometimes assemblage. Rapp. P.-v. Commn int. Explor. Scient. Mer Mtd- display an opposite pattern, with juveniles occupying iterr. 32(1): 35 deeper waters than the adults or than the main part of the Fredj, G., Laubier, L. (1985). The deep Mediterranean benthos. In: adult population (Pereyra 1968, Wigley etal. 1975, Moraitou-Apostolopoulou, M., Kiortsis, V. (eds.) Mediter- Beyers and Wilke 1980, Somerton and Otto 1986, Attrill ranean marine ecosystems. 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