LIFE HISTORY OF THREE FAIRY SHRIMPS (BRANCHIOPODA: ANOSTRACA) FROM THAILAND. Wipavee Dararat, Peter L. Starkweather, and La-orsri Sanoamuang

JOURNAL OF CRUSTACEAN BIOLOGY, 31(4): 623-629, 2011 LIFE HISTORY OF THREE FAIRY SHRIMPS (BRANCHIOPODA: ANOSTRACA) FROM THAILAND Wipavee Dararat, Peter L. Starkweather, and La-orsri Sanoamuang (WD, LS, correspondence, la_orsri@kku.ac.th) Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand; (PLS) School of Life Sciences, University of Nevada, Las Vegas, U.S.A. ABSTRACT Life table characteristics of three species of fairy shrimps from Thailand, Streptocephalus sirindhornae, S. siamensis, and Branchinella thailandensis are reported, based on laboratory cultures at 24-26uC. Comparative data on their growth and fecundity revealed that B. thailandensis had the largest size (male 26.2, female 27.8 mm) and the highest hatchability 87.67%. The shortest hatching time was found in B. thailandensis, it hatched within 3 days, while the other two species hatched within four days. Branchinella thailandensis showed more rapid growth than the other two species, and it reached maturity within 6.5 days. In contrast, S. sirindhornae had the highest fecundity, producing a mean of 35.3 broods and 17,865 eggs per individual female over its life time, whereas the other two species produced less than 8000 eggs. Life span was equal for both males and females in all three fairy shrimp species (P. 0.05). In this study, the fecundity data indicated that the number of eggs per brood recorded in the three species showed a positive correlation with the body length and age of females. The result indicated that the Thai fairy shrimps show several biological attributes making them suitable for aquaculture and other commercial purposes; B. thailandensis shows rapid growth and high hatching percentage which is suitable for mass production, while S. sirindhornae has high fecundity which is appropriate for the commercial production of eggs. KEY WORDS: Branchinella thailandensis, fecundity, life span, Streptocephalus siamensis, Streptocephalus sirindhornae DOI: 10.1651/10-3410.1 INTRODUCTION Fairy shrimp inhabit ephemeral waters such as roadside ponds, rice field ditches, and grassy vernal pools (Gilchrist, 1978; Brendonck et al., 1993; Sanoamuang et al., 2000b). Degrees of rainfall and the depression are important factors for determining the water level in these temporary habitats. Ephemeral waters can be divided into different types of habitats, including predictable and unpredictable ponds (Al-Sayed and Zainal, 2005; Vanschoenwinkel et al., 2010). Under brief water inundation of temporary habitats, most aquatic organisms must adapt their life history traits to survive under this environmental stressful condition by having rapid growth, early egg hatching, and shorter life spans than those in predictable ponds (Hildrew, 1985; Starkweather, et al., 2001; Mura et al., 2003; Vanschoenwinkel et al., 2010). Fairy shrimp are one of the most unique crustacean groups that have adopted their life history, including early hatching, rapid maturation, and early start of egg production, to effectively inhabit temporary ponds. In Thailand, three species of fairy shrimps, Streptocephalus sirindhornae Sanoamuang, Murugan, Weekers, and Dumont, 2000a; S. siamensis Sanoamuang and Saengphan, 2006; and Branchinella thailandensis Sanoamuang, Saengphan, and Murugan 2002, have been discovered. Study of Thai fairy shrimps has mainly focused on their taxonomic status and environmental context. Information about the life history of fairy shrimp has been reported in many species (Hildrew, 1985; Mura, 1991; Simovich and Hathaway, 1997; Beladjal et al., 2003; Mura et al., 2003), but comparable information for the Thai fairy shrimps is lacking. Since they were discovered, efforts have been focused on the cultivation these new species in order to use them as a new live food for freshwater aquatic animals such as prawns, shrimp, and ornamental fish (Saengphan et al., 2005, 2006; Sanoamuang et al., 2006; Boonmak et al., 2007; Plodsomboon and Sanoamuang, 2007; Saengphan and Sanoamuang, 2009; Sriputhorn and Sanoamuang, 2011). Therefore, in order to better understand the life history and make use of these Thai fairy shrimps in aquaculture, the biological characteristics should be investigated. In view of growing needs, the objective of the present study was to define the biological characteristics of the three species of fairy shrimp mentioned above, focusing on the life history characters such as fecundity (number of broods per female, number of eggs per brood, and number of eggs per female), growth, life span, and maximum size. MATERIALS AND METHODS Life History Study Eggs of S. sirindhornae, S. siamensis, and B. thailandensis were obtained from the Applied Taxonomic Research Center, Khon Kaen University, Thailand. Nauplii of each species were hatched from eggs coming from the same batch, and they were fed approximately 1 3 10 6 cells ml 21 Chlorella sp. twice per day. Chlorella sp. was grown according to the 623

624 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 31, NO. 4, 2011 Table 1. Life history characteristics (mean 6 standard deviation and range) of Streptocephalus sirindhornae, S. siamensis and Branchinella thailandensis, n 5 10. Values followed by the same letter(s) are not significantly different (P. 0.05). Life history characteristics S. sirindhornae S. siamensis B. thailandensis Life span (days) Male 71.9 6 14.49 a (59-99) 42.5 6 6.80 b (30-50) 23.9 6 1.11 c (23-27) Female 67.6 6 17.13 a (44-94) 37.7 6 7.80 b (27-51) 22.5 6 2.17 c (20-25) Body length (mm) Male 24.9 6 2.6 a (18.5-28.1) 17.1 6 1.1 b (16.0-19.3) 26.2 6 2.6 d (21.5-28.5) Female 24.3 6 1.8 a (22.3-27.5) 21.3 6 1.9 c (18.4-22.5) 27.8 6 2.2 d (24.0-30.5) Maturation time (days) 13.5 6 1.27 a (12-15) 8.5 6 0.97 b (7-10) 6.5 6 0.53 c (6-7) Number of broods per female 35.3 6 4.92 a (29-42) 18.1 6 4.89 b (13-30) 13.5 6 1.72 c (11-16) Number of eggs per brood 506 6 11 a (242-723) 419 6 41 b (78-682) 470 6 35 c (135-709) Total number of eggs per female for whole life span 17,865 6 2446 a (14,872-21,442) 7634 6 2507 b (5179-13,792) 6365 6 1025 c (5022-8006) Brood frequency (days) 1.26 6 0.13 a (1.07-1.49) 1.45 6 0.34 b (1-2.21) 1.14 6 0.06 c (1.08-1.21) Brood-pouch length (mm) 6.1 6 0.7 a (4.2-6.3) 4.6 6 0.7 a (3.4-5.6) 5.2 6 0.6 a (4.5-6.3) Hatching percentage (%) 64 6 9.17 a (53.33-76.67) 50.67 6 4.66 b (46.67-56.67) 87.67 6 4.98 c (80-90) method described by Saengphan et al. (2005). All experiments were conducted under static room conditions at water temperatures between 24-26uC. These are common temperatures when fairy shrimps usually occur during rainy season in Thailand. After maturation (when eggs are first present in the brood pouch of females and the full-grown antennae are formed on the males), male-female pairs of each species were cultured separately in 2 L capacity plastic jars containing 1 L dechlorinated tap water. Observations continued until the animals died. The growth was observed by measuring the body length of the shrimp along with their development every 5 days. The body length of each individual was measured from the tip of the head to the posterior margin of the caudal furca using a Vernier caliper. All deposited eggs from each individual female were collected and counted under a stereo microscope every day to determine the egg production. Eggs from the first brood were removed within 24 hours after laying and subsequently used for hatching tests. Eggs were first passed through a 1 mm sieve to remove debris and then collected on a 100 mm sieve. For the hatching experiment, 30 eggs were introduced into a 1.5 ml microtube and immersed in dechlorinated tap water for 4 weeks before the experiment according to the egg preparing methods described by Saengphan et al. (2005). Then eggs were incubated in plastic transparent cups containing 50 ml of dechlorinated tap water and then placed in an incubator at 27uC to determine hatchability. The numbers of nauplii were observed daily for 7 days. Data on egg hatching are expressed as 7 days cumulative hatching totals. Hatching percentages of eggs were subsequently calculated. Data Analysis The variance of the data is given as standard deviation (SD) of the mean of ten replicates. The data were analyzed by analysis of variance (one-way and two-way ANOVA). Significant differences (P, 0.05) were discovered by Duncan s multiple range tests. An independent t test was conducted for comparing life span and body length between male and female. Correlation analysis was tested for the relationships between two sets of data (P, 0.01). existed in S. siamensis. The body length of the females of S. siamensis (21.3 mm) was significantly longer than males (17.1 mm). The body lengths of male B. thailandensis and S. sirindhornae were 26.2 and 24.9 mm whereas the body lengths of females were 27.8 and 24.3 mm, respectively. Among the three species, B. thailandensis had the largest size (male 26.2 and female 27.8 mm). Maturation time (which is the time of first spawning) of the three species of fairy shrimps is presented in Table 1; B. thailandensis, S. siamensis, and S. sirindhornae mature to adulthood within 6.5, 8.5, and 13.5 days, respectively (P, 0.05). The growth curves of the three fairy shrimp species are represented in Fig. 1 and Fig. 2. The growth data among the three species indicated that both males and females of B. thailandensis grew more rapidly than the other two species. The number of eggs per brood of S. sirindhornae, B. thailandensis, and S. siamensis were 506, 470, and 418 eggs, respectively. Comparative fecundity data showed that S. sirindhornae produced more broods and eggs than the other two species (P, 0.05), producing a total of 17,865 eggs per female and 35.3 broods per individual female in the whole life span. The frequencies of brood production in B. thailandensis, S. sirindhornae, and S. siamensis were 1.14, 1.26, and 1.45 days, respectively. RESULTS Fecundity and growth data of S. sirindhornae, S. siamensis, and B. thailandensis are summarized in Table 1. Life span was not significantly different between males and females in all three species (P. 0.05). Both males and females of B. thailandensis had shorter life spans (male 23.9, female 22.5 days) than S. siamensis (male 42.5, female 37.7 days) and S. sirindhornae (male 71.9, female 67.6 days) (P, 0.05). There were no significant differences in body length between males and females of B. thailandensis and S. sirindhornae (P. 0.05), but significant gender difference Fig. 1. Growth of male fairy shrimps (Branchinella thailandensis, Streptocephalus sirindhornae, and S. siamensis) estimated from their body lengths for 90 days.

DARARAT ET AL.: FAIRY SHRIMP LIFE HISTORIES 625 Fig. 2. Growth of female fairy shrimps (Branchinella thailandensis, Streptocephalus sirindhornae, and S. siamensis) estimated from their body lengths for 90 days. The hatching rate of the three fairy shrimps (Fig. 3) indicated that the highest hatching percentage was found in B. thailandensis (87.67%), followed by S. sirindhornae (64.0%), and S. siamensis (50.67%). Daily observation on egg hatching for 7 days after incubation showed that eggs of B. thailandensis hatched mostly within the first 3 days, which represented hatchability of 75.66% on the first day, followed by 8.66% on the second day, and 3.33% on the third day. The eggs of S. sirindhornae and S. siamensis hatched within 4 days, which showed a lower hatching rate than B. thailandensis. Eggs of S. sirindhornae and S. siamensis hatched 45.0 and 18.0% on the first day, followed by 11.33 and 21.66% on the second day, 7.0 and 7.66% on the third day, and 0.66 and 3.33% on the fourth day, respectively (Fig. 3). The longer hatching assay was found in S. sirindhornae and S. siamensis. Brood-pouch length of the three species (Fig. 4) positively correlated with the age of female fairy shrimp (P, 0.01; B. thailandensis, r 5 0.81; S. siamensis, r 5 0.71 and S. sirindhornae, r 5 0.69). These data indicate that the length of brood pouch depends on the body length of the female. The fecundity graph representing the number of eggs per body length of females (Fig. 5) shows that the number of eggs per Fig. 4. The relationship between length of brood pouch and age of female fairy shrimps, Branchinella thailandensis, Streptocephalus sirindhornae, and S. siamensis. brood of the three species had a positive correlation with the body length of females (P, 0.01; B. thailandensis, r 5 0.82; S. siamensis, r 5 0.84 and S. sirindhornae, r 5 0.91). Similarly, the number of eggs per brood correlated with the age of the female fairy shrimps (P, 0.01; B. thailandensis, r 5 0.83; S. siamensis, r 5 0.82 and S. sirindhornae, r 5 0.89) (Fig. 6). The pattern of the number of eggs per body length and age demonstrated that the number of eggs per brood increased when female fairy shrimps increased in size. There were no significant differences in the length of the brood pouch among the three species (P. 0.05). However, at the same age of the three species, the lengths of brood pouch and body of B. thailandensis were greater than those of the other two species (P, 0.05) (Fig. 2 and Fig. 4). Consequently, at the same age, among the three species, B. thailandensis produced more eggs than S. sirindhornae and S. siamensis (P, 0.05) (Fig. 6). DISCUSSION Among the three species studied, B. thailandensis had the shortest life span (male 23.9 6 1.11, female 22.5 Fig. 3. Daily hatching percentage of Branchinella thailandensis, Streptocephalus sirindhornae, and S. siamensis. Fig. 5. The relationship between number of eggs per brood and body length of fairy shrimps, Branchinella thailandensis, Streptocephalus sirindhornae, and S. siamensis.

626 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 31, NO. 4, 2011 Fig. 6. The relationship between number of eggs and age of the three fairy shrimp species, Branchinella thailandensis, Streptocephalus sirindhornae, and S. siamensis. 62.17 days), biggest size (male 2.62 6 0.26, female 2.78 6 0.22 cm) and more rapid maturation (6.5 6 0.53 days) than the other species. Besides, this species had a higher hatchability, ranging from 80-90%. Although S. sirindhornae had the longest lifespan, its fecundity (number of broods per female, number of eggs per brood, and number of eggs per female) was higher than those of other species. Our observation corresponds well with the previous studies by Boonmak et al. (2007), which reported that B. thailandensis had a more rapid growth than S. sirindhornae, whereas S. sirindhornae produced much more eggs than B. thailandensis. According to our results, B. thailandensis matured to adulthood within a week (6.5 days) under culture conditions, while the duration from nauplii to adulthood of S. siamensis and S. sirindhornae were about 8.5 and 13.5 days, respectively. All of the Thai species presented grew more rapidly than Streptocephalus vitreus (Brauer, 1877) (17 days) from Kenya (Hildrew, 1985), S. sudanicus Daday, 1910 (16 days) from Sudan (Brendonck et al., 1993), Branchinecta schantzi Mackin, 1952 (15-21 days) from India (Raina and Vass, 2006), and Polyartemia forcipata Fischer, 1851 (3-4 months) from Europe (Brtek and Thiery, 1995). Thus, B. thailandensis is suitable for commercial culture as documented by Saengphan et al. (2006). One of the most interesting biological characteristics of these Thai fairy shrimps is their substantially high fecundity compared to other species previously studied. The number of egg productions by S. sirindhornae, S. siamensis, and B. thailandensis were 17,865, 7634, and 6365 eggs per individual female covering the entire life span, respectively, while S. proboscideus (Frauenfeld, 1873) and S. sudanicus produced 4890 and 3550 eggs (Brendonck et al., 1993). Two species recorded from the Mediterranean region, Branchipus schaefferi Fischer, 1834, and S. torvicornis (Waga, 1842) produced only 1734 and 2460 eggs (Beladjal et al., 2003). The fecundity data indicated that the size of the female showed a positive relation to the fecundity of the three fairy shrimp species, it was also seen that the length of brood pouch increased in size corresponding to the age of the females. Additionally, the number of eggs per brood recorded in the three species increased, as the length and the age of females also increased. For Thai fairy shrimps, of the same age, length of brood pouch and body length of B. thailandensis were longer than the other two species. Hence, at the same age, B. thailandensis produced more eggs than S. sirindhornae and S. siamensis. Consequently, the smaller size fairy shrimps appear to have fewer eggs while larger ones produce more eggs. Hence, the larger sized female fairy shrimps can bear more eggs than the smaller ones. The pattern of egg productions in all three Thai fairy shrimp species exhibited a positive relationship with the brood size and body length of the female as recorded in S. sudanicus and S. proboscideus (Brendonck et al., 1993). Several species of fairy shrimps exhibited different life histories (Simovich and Hathaway, 1997; Beladjal et al., 2003). The time to maturity varies among species even within the same habitat (Al-Sayed and Zainal, 2005). In terms of energy allocation for growth, some species adapt their life history to devote more energy to growth while being insensitive to temperature change (Hazelwood and Hazelwood, 1985). Besides, eggs of fairy shrimp hatch at variable times, thereby spreading the risk of mortality (Hildrew, 1985; Simovich and Hathaway, 1997). The differences of biological performance on life span, growth and fecundity among species might be due to their different natural habitats. Fairy shrimp occur in ephemeral waters such as ponds, roadside canals, rice field ditches and grassy vernal pools, the wet phase of which are subject to periodic drying up, flooding, and erratic rainfall (Gilchrist, 1978; Brendonck et al., 1993; Sanoamuang et al., 2000b). Ephemeral or temporary habitats are typically characterized for shortlasting and long-lasting ponds by variation in timing and duration of water inundations (Vanschoenwinkel et al., 2010). Previous research on life history adaptations of freshwater invertebrates, along with the different ranges of a gradient hydroperiod, was reported (Wissinger et al., 2003; De Roeck et al., 2010). Fairy shrimp have also developed their biological characters to help them persist under unpredictable habitats, including the hatching pattern (Vanschoenwinkel et al., 2010), maturation time (Hathaway and Simovich, 1996), and life span (Beladjal et al., 2003). Here, we investigated differences life history of the three Thai fairy shrimps, S. sirindhornae, S. siamensis, and B. thailandensis under laboratory condition. Based on these results, it would be possible to evaluate these biological data in relation to their natural habitats. Generally, short-lasting ponds have extreme fluctuations in temperature and dissolved oxygen concentration (Gilchrist, 1978; Brendonck et al., 1993). Typically temporary habitats are characterized by water level, short period of inundation, and desiccation phase in the pool bodies (Hathaway and Simovich, 1996; Vanschoenwinkel et al., 2010). Besides, the water level of the ponds depends on erratic rainfall and these ponds would dry up completely within a few weeks. In the short-lived ponds, the time stress is expected to select for early hatching or a short period of hatching. It seems that it is advantageous for fairy shrimp to

DARARAT ET AL.: FAIRY SHRIMP LIFE HISTORIES 627 hatch as soon as possible before the pond dries up (Vanschoenwinkel et al., 2010). Based on our data, we found some biological characters supporting for this hypothesis: B. thailandensis inhabits short-lasting ponds (water inundation approximately 1-2 months) (personal observation), it expressed early hatching up to 87.67% within 3 days after inundation and presented higher hatching rate for the first day (75.67%) than the other species occurring in more stable ponds. Additionally, selection in the short-lasting pond is believed to promote rapid maturation, early start of reproduction and shorter life span (Hathaway and Simovich, 1996; Beladjal et al., 2003; Mura et al., 2003; Vanschoenwinkel et al., 2010). Based on our observations, B. thailandensis shows the biological characteristics that might have evolved as life mechanisms to survive under these environmental conditions by having characteristics of rapid growth, early maturation and short life span. This observation also corresponds to the work of Mura et al. (2003) who suggested that the population of fairy shrimp, Chirocephalus diaphanus Prévost, 1803, occurring in a shallow habitat presented rapid growth, precocious reproduction and a short life cycle. Therefore, it implies that the life history traits of B. thailandensis may relate with its unpredictable habitats as previously recorded in Tanymastix stagnalis (Linnaeus, 1758) (cf. Mura, 1991), Branchinecta sandiegonensis Fugate, 1993 (Hathaway and Simovich, 1996), and Branchipus schaefferi (Beladjal et al., 2003). On the other hand, slower maturation, late reproduction and long life span were recorded from animals inhabiting long-lasting ponds or predictable habitats. The long-lasting ponds are considered as a predictable and relatively constant environment which seems to match with the adaptive ability and life history pattern of fairy shrimps including long period of hatching, late reproduction and long life span (Hathaway and Simovich, 1996; Beladjal et al., 2003; Mura et al., 2003; Vanschoenwinkel et al., 2010). The long-lasting pond might stimulate animals to allocate more energy to growth and postpone their reproduction (Johansson and Suhling, 2004). Consequently, slow maturation can be beneficial by making animals produce a greater total of egg and higher quality offspring (Vanschoenwinkel et al., 2010). Our findings correspond with this hypothesis. S. sirindhornae inhabiting long-lasting ponds, the life history may match with their predictable habitats. S. sirindhornae was normally found in long-lasting ponds (water inundation approximately for 3-6 months) (personal observation). The longer hatching assays were found in S. sirindhornae, hatching within 4 days and showing a lower hatching rate compared with B. thailandensis. The lower hatchability 45.0% for the first day and 64.0% for cumulative hatching were recorded in this species. The life history of S. sirindhornae is characterized by slow growth, late reproduction, and a long life span. Based on biological data, S. sirindhornae may have developed a life history to optimize its fitness, longevity, and reproductive ability in its predicable habitat as previously reported in C. diaphanus (Mura, 1991), S. woottoni Eng, Belk and Eriksen, 1990 (Hathaway and Simovich, 1996) and S. torvicornis (Beladjal et al., 2003). Fairy shrimp have evolved biological characteristics to live under ephemeral conditions. Life history characters such as rapid growth to maturity, short life span, fecundity and ability to form resistant eggs have been suggested to be important and essential characters for survival of the group (Simovich and Hathaway, 1997; Starkweather et al., 2001; Mura et al., 2003). In term of species distribution, the distribution of fairy shrimps relates to the movement of eggs by wind (Graham and Wirth, 2008). Among the three species, S. sirindhornae is the most widely distributed species in Thailand (Sanoamuang et al., 2000a), which is in agreement with the findings of Brtek and Mura (2000) who reported that Streptocephalus is the most widespread and tolerant genus of fairy shrimps in the world. The widespread distribution of S. sirindhornae in Thailand may depend on their high sexual ability, i.e., high fecundity. Our results revealed that S. sirindhornae can produce a lot of eggs, up to 17,865 eggs per individual female. Therefore, its eggs would have more chance to be dispersed by wind than the other two species. Another explanation could be that S. sirinhornae seems to be a tolerant species in Thailand. It occurs in a wide range of habitats with many environmental variables when compared with B. thailandensis (Sanoamuang et al., 2000b). The co-occurrence of fairy shrimps in Thailand has been reported. B. thailandensis co-occurs in several localities with the common species S. sirindhornae. However, B. thailandensis always emerged prior to S. sirindhornae and usually lived for 4-5 weeks (Sanoamuang et al., 2002). The two coexisting species generally inhabit a group of ephemeral pond. The egg hatching of these two species seem to respond to the same hatching stimuli (Saengphan et al., 2005, 2006), since their nauplii can coexist but they can be separated by morphological size difference along with their different maturation times. The significant differences recorded as to size, time in attaining sexual maturity and life span are in favor of a niche separation pattern as previously recorded in the cooccurring species C. diaphanus and T. stagnalis (Mura, 1991); Branchinecta sandiegonensis and S. woottoni (Hathaway and Simovich, 1996); Branchiopus schaefferi, and S. torvicornis (Beladjal et al., 2003). The life histories of S. sirindhornae and B. thailandensis, in particular the different maturation times are supposed to reflect the size difference, also in filtering apparatus level relating to the different size prey selections (Mura, 1991). Hence, the different life history traits of coexisting species might reduce their food competition enabling them live together. Therefore, the maturation time and life span might play a role in restricting the coexisting fairy shrimp species S. sirindhornae and B. thailandensis in their natural habitats. Regarding S. siamensis, it is a very rare species found only in five temporary pools out of 700 sampled localities from only two provinces of Thailand. They have not been found in the water bodies of natural habitats. Their samples were obtained only from dried mud collections (Sanoamuang and Saengphan, 2006). Since it was observed only

628 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 31, NO. 4, 2011 from the egg hatching experiment, the environmental variable data is limited. Hence, the life history pattern of S. siamensis from our experiment can be used as information for indicating its natural habitats. Life history information can also be useful for the cultivation system of Thai fairy shrimp species. The result indicated that the Thai fairy shrimps show several biological attributes making them suitable for aquaculture and other commercial purposes. B. thailandensis, for example, shows rapid growth and high hatching percentage which is suitable for mass production, while S. sirindhornae has high fecundity which is appropriate for the commercial production of eggs. Further research on mass production of fairy shrimp will be necessary to further promote these crustaceans for commercial purposes. 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