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УДК 595.384.12
SOME OF REPRODUCTIVE TRAITS OF SNAPPING SHRIMP ALPHEUS LOBIDENS DE HAAN 1850 (ALPHEIDAE) FROM THE NORTHERN COAST OF THE OMAN SEA
Z. Ansari, A. Maghsoodlou
НЕКОТОРЫЕ ОСОБЕННОСТИ РЕПРОДУКТИВНОЙ БИОЛОГИИ КРЕВЕТКИ-ЩЕЛКУНА ALPHEUS LOBIDENS DE HAAN 1850 (ALPHEIDAE) СЕВЕРНОГО ПОБЕРЕЖЬЯ ОМАНСКОГО МОРЯ
З. Ансари, А Махсудлю
Some biological traits of shrimp Alpheus lobidens reproduction have been studied for the first time. Shrimps were collected under rocks/boulders, in an intertidal area of the Oman Sea (Chabahar, Iran) in August 2014. 58 females have been studied of carapace length 6-15 mm including 48 ovigerous females of 7-14 mm. The mean realized fecundity (the number of embryo per pleopods) at stage 1 of embryonic development was 203±181. The realized fecundity values were significantly related to shrimp size. The volume of recently-produced embryos was on average 0.076 mm3, and embryo volume increased by 107.9 % during the incubation period. The eggs of A. lobidens showed changes in form and a constant increase in volume during the incubation period. The eggs of the first two stages were spherical, and then found ellipsoidal form by the end of development. The total percentage increase (107.9% from the first to the final stage of development) was high compared with other alpheids such as A. normanni, A. nuttingi and A. cf. burukovskyi. There are differences in embryo volume in these species of different areas that can be affected by different environmental strategies. The average of embryo loss during embryogenesis was 6.4%. Females carrying embryos close to hatching showed fully developed ovaries, suggesting consecutive spawning.
shrimps, Alpheus lobidens, reproductive traits, realized fecundity, the Oman Sea, continuous cyclicity of reproduction
Впервые исследованы некоторые особенности репродуктивной биологии креветки Alpheus lobidens De Haan 1850 (Alpheidae). Креветки собраны в августе 2014 г. на литорали Оманского моря (вблизи порта Чахбехар, Иран), на илисто-песчаном грунте с гравием и мелкими валунами, под которыми они скрывались во время отлива. Исследованы 58 самок с длиной карапакса 6-15 мм, из них 48 -яйценосные особи с длиной карапакса 7-14 мм. Средняя реализованная плодовитость (количество эмбрионов, вынашиваемых на плеоподах) в 1-й стадии эмбрионального развития была равна 203±181. Эта величина зависит от длины тела креветок. Объем только что отложенных яиц в среднем составляет 0,076 куб. мм, в процессе эмбрионального развития он возрастает на 107,9%. Яйца A. lobidens в данном процессе демонстрируют не только увеличение размеров, но и изменение формы. На первых двух стадиях эмбрионального развития они имеют почти сферическую форму, к концу процесса инкубации приобретая форму элипсоида. Общее увеличение с первой до последней стадии развития хорошо сравнимо с этим явлением у других видов рода, по которым имеется информация
(A. normanni, A. nuttingi и A. cf. burukovskyi ). Различия в объеме яиц у этих видов могут быть объяснены разными условиями их обитания. У самок с эмбрионами, близкими к вылуплению, гонады демонстрируют ооциты, готовые к нересту. Это говорит о том, что у них наблюдается непрерывная цикличность размножения.
креветки, Alpheus lobidens, репродуктивные особенности, реализованная плодовитость, Оманское море, непрерывная цикличность размножения
INTRODUCTION
Caridean shrimps of the family Alpheidae Rafinesque, 1815 are one of the most important, numerous and ecologically diverse groups among the decapods [1]. Alpheus Fabricius, 1798 is the most numerous genus of this family, and probably of the order Decapoda Latreille, 1802; it currently contains more than 280 species [2]. Genus Alpheus is represented by ten species, which have been reported from Iranian coast of Persian Gulf (8 species) and Oman Sea (2 species) [3, 4].
Our studied shrimps are the new record of Alpheus lobidens De Haan 1850, for Oman Sea. Sheibani (2007) [4] and Naderloo et al. (2013) [3] summarized the available information on marine decapod diversity in Iranian waters of Persian Gulf and Oman Sea, but did not mention A. lobidens,. The shrimp A. lobidens has so far only been reported from two locations in the intertidal zones of Oman Sea (Iran).
Considering the great diversity and ample geographic distribution of the members of the family Alpheidae, there are still relatively few studies on this group. In Iranian waters of Persian Gulf and Oman Sea, investigations on alpheids are totally lacking, with no previous studies on the reproduction of this group. Therefore, the purpose of this study is descriptions on some reproductive characteristics of A. lobidens
MATERIALS & METHODS
All studied shrimps were collected in August 2014 during field trip in the intertidal zone of Chabahar beach, on the coast of the Sea of Oman, Iran in Djod station (25°26'58'' N 59°30'28'' E) (fig. 1). 58 females studied of carapace length CL (mm) included 48 ovigerous females of CL (mm). The structure of substrate was composed of silty-sandy with pebbles and boulders. Samples were taken during low tide by hand under boulders, by four people and fixed in 4% neutralized formalin solution. The sea surface temperature and salinity in sampling time varied between 30°C and 38 ppt., respectively.
The carapace length (CL, distance between distal part of the eye socket to the posterior margin of the carapace) was measured using a stereoscopic microscope MBS-10. The weighing of shrimps was made on electronic balance "Professional digital table top scale" up to 0.1 g. The weighing of gonad prespawning females and the egg production was determined on balance TU 64-1-990-72 up to 0.005 g.
In order to identify an absolute individual fecundity (number of oocytes in the gonad of prespawning females - ABF) and realized fecundity (number of embryos in females- RBF) is completely counted the number of oocytes in the gonad and the number of embryos.
The length and width of 10 embryos in each clutch and 10 oocytes from each prespawning gonads were measured with a eyepiece micrometer. The volume of the embryo was calculated using the formula V = 1/ 6 x ir x d Xx ( d2 )2, where d 1 greater diameter and d 2 lesser diameter [5]. To determine the relationship between the length and number of embryos was used nonlinear regression analysis.
The developmental stage of embryo and stage of ovarian development was assigned following the methodology proposed by Burukovsky [6]. To avoid possible egg loss and change in egg volume during incubation and embryonic development, only females with recently produced eggs (Stage 1) were included in the fecundity analysis.
Fig. 1. Sampling locality along the Iranian coast of the Oman Sea Рис. 1. Место сбора проб на иранском побережье Оманского моря
To evaluate changes in the number and volume of the embryo during the incubation was used one-sided analysis of variance.
RESULTS
A total of 58 shrimp females were studied with CL 6-15 mm (fig. 2). Of these, 10 were non-ovigerous females (17.2%) and 48 were ovigerous females (82.7%). The ovigerous females with eggs were at different developmental stages: 43.7% females carried embryos in stage 1, 16.7% in Stage 2, 12.5% in stage 3, 14.6% in stage 4, and 12.5% in stage 5.
25
20
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□ Non-ovigerous
□ Ovigerous
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6 7 8 9 10 11 12 13 14 15 Carapac length (mm)
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Fig. 2. Frequency distribution of carapace length of females of A. lobidens (N=58); Oman Sea, Iran Рис. 2. Размерный состав самок A. lobidens (N = 58); Оманское море, Иран
Early ovarian stages (Stage II-III) predominated in females carrying recently-extruded embryos (Stage 1-2), while ovaries filled with vitellogenic oocytes (Stage IV) reached its highest occurrence in females with embryos close to hatching (Stage 3) (fig. 3).
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Fig. 3. Stage of ovarian development in relation to the stage of embryo development from females of A. lobidens (N=58), Oman Gulf, Iran Рис. 3. Стадия зрелости яичников по отношению к стадии развития эмбрионов самок A. lobidens (N = 58) берегов Оманского моря Ирана
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Fig. 4. Relationship between number of embryos in stage 1and carapace length of A. lobidens females (N=58), Oman Gulf, Iran Рис. 4. Соотношение между количеством эмбрионов на стадии 1 и длиной карапакса A. lobidens (N=58) самки берегов Оманского моря Ирана
The average number of stage 1 embryos was 203 ± 181 embryos per female, with a minimum and maximum of 22 and 785, respectively. Relative fecundity ranged from 3 to 56 embryos/mm carapace length (mean ± SD: 18±13). In this study,
3 ovigerous females were in V stage of ovarian development with a mean number of oocytes 217. The maximum and minimum of oocyte in these females were 267 and 190, respectively. The average of oocyte diameter for these females was 0.771 mm with a minimum and maximum of 0.765 and 0.785 mm.
Recently-produced embryos (Stage 1) had an average volume of 0.076 mm , and those closed to hatching (Stage 5) 0.158 mm3 (table 1), representing a volume increase of 107.9 % during the incubation period.
DISSCUSSION
In studied sample 60.4% females carried embryos in stage 1 and 2, 12.5% in stage 3, and 27.1% in stage 4 and 5. In the last three stages of embryonic development, the ovaries filled with vitellogenic oocytes. For ovigerous females in recent stages (IV and V), there are embryos near to hatching stage. Therefore, it is evident that the females of A. lobidens have continuous cyclicity of reproduction.
Table 1. Data on the number of studied specimens, mean ± standard deviation of carapace length (CL), embryo number, and embryo volume according to the stage of embryo development in females of A. lobidens from Oman Gulf, Iran
Таблица 1. Связь между изменением размеров карапакса, стадией эмбрионального развития, реализованной плодовитостью и объемом эмбриона_
Stage of embryo development n CL (mm) Embryo number Embryo volume (mm3)
1 21 10 ± 2.4 203 ± 181 0.076 ±0.007
2 8 9.8 ± 1.8 152± 117 0.076 ±0.009
3 6 11.2 ± 1.7 265 ± 205 0.085 ± 0.007
4 7 10.6 ± 2.1 248 ± 303 0.112 ± 0.019
5 6 11.7 ± 2.5 190± 154 0.158 ± 0.015
There was not significant egg loss during embryo development process. The egg losses of up to 23% for Alpheus nuttingi [7] and 12% for A. armillatus [8] have been reported. In contrast, we observed much lower level of egg loss for A. lobidens but this finding was similar to level of egg loss estimated for A. cf. burukovskyi [9]. Therefore, our results indicate the presence of efficient parental care from the embryos in A. lobidens during the developmental process. It may be related to the presence of sufficient space under the female abdomen to care the number of eggs produced until they hatch.
The eggs of A. lobidens showed changes in form and a constant increase in volume during the incubation period. The eggs of the first two stages were spherical, and then found ellipsoidal form by the end of development. The total percentage increase (107.9% from the first to the final stage of development) was high compared with other alpheids such as A. normanni [10], A. nuttingi [7] and A. cf. burukovskyi [9]. There are differences in embryo volume in these species of different areas that can be affected by different environmental strategies.
The maximum relative fecundity to carapace length for A. lobidens (56 eggs/mm CL) is much lower than given for some other species of the Alpheus, e.g., 78 eggs/mm CL for A. normanni, 149 eggs/mm CL for A. armillatus, 237 eggs/mm CL for A. nuttingi, 131 eggs/mm CL for A. cf. burukovskyi. However, the maximum relative
fecundity to CL for A. lobidens is about 2.5 times higher than the value reported for A. heterochaelis (Table 2)
The differences in fecundity found among species related to the genus Alpheus may reflect the differing environmental conditions of their habitat and different reproductive strategies.
Table 2. Data on the number of studied shrimps, carapace length, embryo number, embryo volume, and fecundity (max)/CL of females of species of Alpheidae family, from literature sources and the present study.
Таблица 2. Сравнительная характеристика основных репродуктивных параметров некоторых видов креветок из рода Alpheus____
Species N Carapace length (mm) Embryo number (Max) Embryo volume (mm3) Fecundit y (Max) / mm CL Locality Referen ce
A. normanni 7 4.1-7.4 584 0.09 78 Florida, USA [10]
A. armillatus 31 7.7-14.1 1979 0.1 149 Florida, USA [8]
A. lobidens 48 14 785 0.076 56 Oman Sea Present study
A. cf. burukovskyi 75 14 1843 0.11 131 Oman Sea [9]
A. nuttingi 27 11.2-18.6 4222 0.09 237 Sao Paulo, Brazil [7]
A. saxidomus 5 10.1-17.4 - 0.25 The east of Pacific [11]
A. heterochaelis 5 10.2-13.4 336 0,91 25 Florida, [10]
THE LIST OF THE USED REFERENCES
1. Anker A., Dworschak P.C. Jengalpheops rufus gen. nov., sp. nov., a new commensal alpheid shrimp from the Philippines (Crustacea: Decapoda). Zoology Study, 2007, no. 46, pp. 290-302.
2. De Grave S., Fransen C.H.J.M. Carideorum catalogus the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea, Decapoda) // Zool. Meded. Leiden V, 2011, no 9, pp. 195-588.
3. Naderloo R., Turkay M., Sari A. Intertidal habitats and decapod (Crustacea) diversity Of Qeshm Island, a biodiversity hot spot within the Persian Gulf. Marine biodiversity, 2003, ISSN 1867-1616. DOI 10. 1007/s 12526-013-0174-3.
4. Sheibani R. Zoogeography and taxonomy of littoral Caridean shrimps of the Persian Gulf and Gulf of Oman. PHD Thesis. Shahid Beheshti University, Faculty of Biological Sciences. Tehran, 2007.144 p.
5. Turner R.L., Lawrence J.M. Volume and composition of echinoderm eggs: implications for the use of egg size in life-history models. In: Stancyk SE (Ed.) Reproductive Ecology of Marine Invertebrates. University of South Carolina Press, Columbia, USA, 1979. pp. 25-40.
6. Буруковский Р.Н. Методика биологического анализа некоторых тропических и субтропических креветок. Промыслово-биологические исследования морских беспозвоночных [Biological analysis procedure of some tropical and subtropical shimps. Field- biological research of marine invertebrates] . ВНИРО, М. -1992.-С. 74-84.
7. Pavanelli C.A.M., Mossolin E.C., Mantelatto F.L. Maternal investment in egg production: environmental and population-specific effects on offspring performance in the snapping shrimp Alpheus nuttingi (Schmitt, 1924) (Decapoda, Alpheidae). Animal Biology, 2010, no. 60, pp. 237-247.
8. Pavanelli C.A.M., Emerson C. Mossolin., Fernando L. Mantelatto. Reproductive strategy of the snapping shrimp A. armillatus H. Milne-Edwards, 1837 in the South Atlantic:fecundity, egg features, and reproductive output. Invertebrate Reproduction and Development, 2008, no. 52 (3), pp. 123-130.
9. Ансари З, Махсюдлю A. Репродуктивная биология креветки Alpheus cf. burukovskyi (Crustacea, Alpheidae) с Иранского побережъя Оманского моря, научный журнал «Известия» КГТУ 2016 [Reproductive biology of a shrimp Alpheus cf. burukovskyi (Crustacea, Alpheidae) from the Iranian coast of the Oman sea, scientific journal "Izvestiya" KSTU 2016], no. 42, pp. 21-29.
10. Corey S., Reid D.M. Comparative fecundity of decapod crustaceans. I. The fecundity of thirty-three species of nine families of caridean shrimp. Crustaceana, 1991, no. 60, pp. 270-294.
11. Wehrtmann I.S., Graeve M. Lipid composition and utilization in developing eggs of two tropical marine caridean shrimps (Decapoda: Caridea: Alpheidae, Palaemonidae). Comp. Biochem. Physiol, 1998, no. 121, pp. 457-463.
ИНФОРМАЦИЯ ОБ АВТОРАХ
Ансари Зейнаб - Калининградский государственный технический университет; аспирантка кафедры «Ихтиопатология и гидробиология»;
E-mail: zeinab6228@gmail.com
Ansari Zeinab - Kaliningrad State Technical University; Postgraduate student of the Department «Ichthyopathology and Hydrobiology»;
E-mail: zeinab6228@gmail.com
Махсюдлю Аболвахаб - Иранский Национальный институт океанографии и атмосферных наук; руководитель океанических наук Научно-исследовательского
центра; E-mail: Wahab@inio.ac.ir
Maghsoodlou Abdolvahab - Iranian National Institute for Oceanography and Atmospheric Sciences; Head of Ocean Science Research Center;
E-mail: Wahab@inio.ac.ir
