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Some Biological Characteristics of Tadpole Shrimp, Triops Cancriformis, from Seasonal Pools of West Azarbaijan (Iran)

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Some Biological Characteristics of Tadpole Shrimp, Triops Cancriformis, from Seasonal Pools of West Azarbaijan (Iran) J. Agric. Sci. Technol. (2009) Vol. 11: 81-90 Some Biological Characteristics of Tadpole Shrimp, Triops cancriformis, from Seasonal Pools of West Azarbaijan (Iran) A. Golzari1, S. Khodabandeh1*, and J. Seyfabadi1 ABSTRACT The tadpole shrimp of genus Triops is a well-known living fossil whose fundamental morphology has been unchanged for 220 million years. We collected specimens of Triops cancriformis in temporary water bodies near the southern part of Urmia Lake (in the Fall of 2005). Some biological characteristics of this Triops were investigated. The feeding re- gime of T. cancriformis was found to be related to the fauna and flora of the temporary pools. Invertebrates and animal detritus were found to constitute major part of the feed- ing regime. The existence of Triops cysts and particles in the gut also showed certain de- gree of cannibalism. Morphological and histological investigations showed that the popu- lation of T. cancriformis was female and there was only one male among 400 samples col- lected. Observation of sperm among follicle ducts of a few samples indicated some degree of hermaphrodity, but the animal seemed to reproduce mainly through parthenogenesis. Fecundity, varying from 100 to 2500 cysts, was with a few exceptions related to the body size. The average cyst diameter was 400±±±85 µµµm. Keywords: Crustacean, Feeding Regime, Notostraca, Reproduction, Shrimp, Tadpole Triops cancriformis. INTRODUCTION Petrov, 1999). Tadpole shrimps are charac- terized by their wide fluctuations in popula- Notostracans are freshwater crustaceans tion density. When their density becomes adapted to temporary water bodies (Su and very high, they can barely find enough food, Mulla, 2001). They are commonly called and they become very active in searching for tadpole shrimps because of their superficial food. Food shortage inhibits their develop- resemblance to frog larvae. They are recog- ment and reduces fecundity, while mortality nizable by their large, horseshoe–shaped increases as a result of cannibalistic behav- dorsal carapace (Martin and Boyce, 2005). ior (Scholnick and Snyder, 1996). They are benthic branchiopods which swim The tadpole shrimp genus Triops is a well- and feed ventral surface down, burrowing known living fossil whose fundamental into the bottom sediments in search of detri- morphology has remained unchanged for tus and a varietyArchive of small organisms (Schol- of220 million SID years. While the order has wide nick and Snyder, 1996). Their feeding de- geographical distribution, many species have pends on their habitats. Being facultative a restricted local distribution (Suno–Uchi et detritus feeders or scavengers or predators, al., 1997). they eat algae, bacteria, protozoa, rotifers, The ability of Triops to develop and grow earthworms, insects, fairy shrimps, frog eggs quickly is crucial to its success (Davis and and also tadpoles (Cvetkovic-Milicic and Madison, 1999). Fecundity is related to oxy- _____________________________________________________________________________ 1 Department of Marine Biology, Faculty of Marine Sciences, Tarbiat Modares University, Noor, P.O. Box: 46414-356, Islamic Republic of Iran. * Corresponding author, [email protected] 81 www.SID.ir _______________________________________________________________________ Golzari et al. gen tension and temperature and so small tini, 1980; Scanabissi et al., 2005). changes in temperature or oxygen dramati- Although some scattered networks on this cally change the total number of eggs laid in organism- under its general name “Apus”- one season (Scholnick, 1995). have been traced to fish culture ponds in The females possess two ovisacs (brood Iran (Kheirandish, 1975; Kohnehshahri and pouches) on the 11th appendages. The two Takami, 1974), no specific work on its iden- ovisacs hold an almost equal number of tification and biological characteristics, par- eggs. Eggs are laid randomly and not in ticularly feeding and reproduction, have clusters, although all the eggs in both brood been conducted. In this work, therefore, ef- pouches are released simultaneously (Sea- forts are concentrated towards precise identi- man et al., 1991). The fecundity of T. can- fication, and study of the feeding and repro- criformis is high; individuals may lay more ductive biology of this organisms in sea- than 1000 eggs (Fry and Mulla, 1996). Their sonal ponds of West Azarbaijan Province. eggs lie dormant in the soil and can survive as such for many years. They are resistant to MATERIALS AND METHODS severe drought and extreme temperatures (Takahashi, 1997). It has been suggested that the distribution of many branchiopods Animals results from the transfer of resting eggs, ei- ther by birds, people or cattle and sheep. The first adult specimens of T. cancri- Most eggs hatch within a few days of being formis were collected in the Fall of 2003 submerged, but some remain unhatched in from vernal pools of West Azarbaijan Prov- the soil. This is an effective way for the ince in Iran (Figure 1A) (Khodabandeh et shrimps to maintain their populations under al., 2008). For biological studies 400 speci- unpredictable fluctuations in environmental mens (length = 2-5 cm) were collected dur- conditions (Takahashi, 1997). ing the Spring and Fall of 2005. 95 speci- 15 species of Triops have been reported in mens were fixed for 24 hours in Bouin’s the world. Triops cancriformis inhabits tem- fixative and alcohol 70% separately for porary pools and rice fields in Eurasia and feeding regime and reproduction studies. Africa (Cesari et al., 2004; Cvetkovic- Milicic and Petrov, 1999). T. cancriformis distribution in Europe has been reported by Feeding Regime Zaffagnini and Trentini (1980). While the order has a wide geographical distribution, Study of the feeding regime was accom- many of the species have a restricted local plished by investigating the gut contents of distribution (Takahashi, 1997). 50 specimens under the microscope, using T. cancriformis populations reproduce Rose Bengal staining and histological meth- gonochorically, hermaphroditically or even ods. The Triops, gut was separated by scal- parthenogenetically (Zaffagnini and Tren- pel under a stereo microscope, and its con- tini, 1980; CesariArchive et al., 2004). They com- tentsof were SIDpreserved in alcohol 70% and prise bisexual populations, with an equal then studied under a light microscope. Rose male: female sex ratio or with a female bias, Bengal (0.5 g in 500 cc distilled water) was and unisexual populations, either hermaph- added to the contents of the gut to distin- roditic or parthenogenetic (Mantovani et al., guish the vegetal or animal origins of the 2004). The more northerly European popula- food. tions are parthenogenetic, while incidents of For classic histological studies, the animals gonochorism begin appearing towards the were fixed for 24 hours in Bouin’s fixative. southern latitudes, with the northern African The specimens were then fully dehydrated in populations presenting both sexes and am- a graded ethanol series and embedded in phigonic reproduction (Zaffagnini and Tren- paraffin. Sections (5µm) were cut on a mi- 82 www.SID.ir Biology of Triops cancriformis ________________________________________________ crotome, collected on albumine-glycerine 1D-F). The anterior dorsal portion of the slides and stained with Haematoxylin, Eosin carapace is equipped with a relatively and Methyl green (Martoja and Martoja- unique optical arrangement (Figure 1E). The Pierson, 1967; Khodabandeh et al., 2005a trunk consists of a thorax bearing append- and b). ages and an abdomen without them. The oral region of the cephalon possesses specific Reproduction appendages including antennules, antennae, maxillae and mandibles that are covered by a labrum and used for food handling. The Morphological observation and a histo- anterior portion of the thorax consists of 11 logical method were used for sex differentia- segments, each bearing a pair of appendages tion and identification of reproductive sys- (thoracopods) (Figure 1F). The 11th append- tem structure. Fecundity was determined by ages of females form ovisacs (brood counting the cysts in the ovisacs and gonads pouches) (Figure1F). The many appendages of 45 specimens. Biometry of the cysts was posterior to the 11th move the spent feeding accomplished by light microscope with a and respiratory current away from the body. micrometer. The diameters of 12 cysts from The proximal portion of the abdomen pos- each specimen were measured, and the aver- sesses 65 pairs of fine, hair-like appendages age, maximum and minimum diameter was that beat in a rhythmic fashion to assist in calculated. movement and food channeling. The distal portion of the abdomen terminates in a RESULTS prominent telson and subsequently branches into two large caudal furca (Figures 1D-F). Stereo microscopic studies revealed that Triops cancriformis is reported from the gut of T. cancriformis was filled with throughout the southern Palaearctic region, microscopic vegetal, animal, and unknown and from vernal pools of West Azarbaijan particles (Figures 2A-H). Invertebrates and Province in Iran (Khodabandeh et al., 2008). animal detritus were found to constitute a Morphologically the material is a mixture of major part of the T. cancriformis feeding specimens bearing the carapace carinal spine regime. The existence of Triops cysts and arrangements of both T. c. cancriformis and Triops particles in the guts also showed T. c. simplex subspecies, casting doubts as to some degree of cannibalism (Figures
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