ASPECTS OF BIONOMICS AND LIFE TABLE STUDIES OF FOUR BULINIID SNAILS FROM PARTS OF SOUTH EASTERN NIGERIA

*1NWAUGO V.O.; 2NDUKA, F.O and 2KALU, M.K

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Department of Microbiology Abia State University, Uturu, Nigeria

Department of Animal and Environment Biology Abia State University, Uturu Nigeria

*Corresponding Author Email: vonwaugo @yahoo.com Phone: 2348063494654.

ASPECTS OF THE BIONOMICS AND LIFE TABLE STUDIES OF FOUR BULINIID SNAILS FROM PARTS OF SOUTH EASTERN NIGERIA *1NWAUGO V.O.; 2NDUKA, F.O and 2KALU, M.K
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Department of Microbiology Abia State University, Uturu, Nigeria Department of Animal and Environment Biology Abia State University, Uturu Nigeria Buliniid snails are important intermediate hosts of Schistosoma heamatobium,

ABSTRACT
the causative agents of urinary Schistosomiasis. A study of their bionomics will help to provide an understanding of appropriate control measures for the infection. In this study some aspects of the bionomics and life table pattern of four buliniid snails from South Eastern Nigeria were carried out by observing their egg laying capacity and hatching in the laboratory. B. globosus and B. truncatus were observed to lay fewer egg masses but with more eggs/mass than B. forskalli and B. senegalensis. The Net Reproductive rate (Ro) varied according to species with the highest being 995.20 (B. globosus), 762.84 (B truncatus) 582.56 (B forskalli) and the least 541.0 (B. seneglansis). Mean Generation time (Tc) gave values of 132.01 (B. globosus), 122.67 (B truncatus), 11.27 (B. forskalli) and the least 97.88 (B. senegalensis). The cumulative effects of all these parameters gave similar values of (capacity to increase), as 0.0519 (B globosus) 0.0541 (B truncates), 0.0572 (B. forskalli) and 0.0643 (B senegalensis). These values were not statistically different (P < 0.05) an indication a natural population balancing effect. While B. globosus and B. truncatus eggs hatched within 7-14 days, those of B. forskalli and B senegalensis hatched within 6-12 days. Oviposition varied according to species though each had 2-3 peaks and falls. B globosus and B. truncatus began oviposition at 7+1 weeks with 41 and 33 eggs/snail and peaked at week 20 with 267.26 and 231.51 egg/snail respectively. B forskalli and B. senegalensis oviposited at 6+1 weeks with 27 and 26 eggs/snail and peaked at week 18 with 224.81 and 217.51 eggs/snail respectively. Ovipositon decreased extensively but did not cease till death. Growth showed similar pattern. Hatchlings ranged from 0.50 0.65mm which increased greatly till oviposition when it decreased and virtually ceased at the peak of oviposition indicating diversion of

growth energy to oviposition. The implication of these findings to the epidemiology of urinary schistosomiasis is then discussed.

Keywords; Bulinus species, bionomics, oviposition, life table, growth INTRODUCTION Schistosomiasis is a chronic blood disease caused by species of the digenetic trematode of the genus Schistosoma. The presence of the disease is determined by the presence of the susceptible snail intermediate hosts (Bulinus species) in addition to the socio-economic and cultural practices of the people. In view of this, schistosomiasis remains endemic where there is favourable disposition to its occurrence with regard to the factors above (WHO, 1991, Okafor, 1984, Savioli & Mott, 1989). In Nigeria, the presence of the main Bulinus species known to harbour S haematobium has been reported in various parts. These include B. globosus, B trunactus and B. senegalensis and B. forskalli. In northern parts of Nigeria, Ukoli & Asumu (1980) and Betterton & Fryer (1982) have reported the presence of B. globosus, B. truncatus, and B. forskalli. Cowper (1963) and Ndifon (1983) observed B. globosus in the west. The work of Okafor (1984), Emejulu et al (1994) and Nduka et al, (1995) have confirmed the presence of the Bulinus species in the East. B. senegalensis and B. forskalli have also been variously reported in Nigeria (Ukoli & Asumu, 1980, Nduka et al, 1995, Agi 1995b), Among these snails, there is little or no interchange of parasites which indicate species, strain and even locality specificity. (Chu et al, 1978; Woolhouse & Chardiwana 1990). Though the presence of the various Buinus species in Nigeria have been reported severally, attention has been in identifying their habitats and their roles as host to various Schistosoma species. This work therefore provides information concerning their life table pattern, and aspects of the reproductive bionomics.

MATERIALS AND METHOD (i) Snail collection and maintenance; Buliniid snails used in this study were obtained from water-filled abandoned Quarry pits in parts of Eastern Nigeria. The specific areas were Lokpaukwu, Ishiagu and Agwu, All the communities lie between Okigwe and Enugu along the Port Harcout Enugu Expree highway. Twenty (20) snails of each Bulinus species were used for the studies. The reproductive bionomics of these snails were studied using snails maintained in the laboratory in small plastic containers harbouring five (5) each and water from the Quarry pits (where they were collected). Fresh lettuce leaves were supplied to the snails ad libitum for their feeding. The water was changed every two days to get rid of the snail wastes while glass slides were provided as oviposition materials. LIFE TABLE STUDIES Freshly laid eggs of each Bulinus species were transferred to filtered Quarry pit water for incubation. Eggs which failed to hatch within 3 days after the last hatch was regarded as not viable. Changing of water in the containers harbouring juvenile snails was done using pipettes while 3 drops of 10% solution of ground chalk was added into the Quarry pit water to supply calcium ions. Following regular observation of these laboratory- bred snails, life tables were constructed for each Bulinus species using the method of Schiff (1964) as described by Okafor (1984) and Okoronkwo (1986). The table which consisted of pivotal age of snail (x), time of egg-laying, (zero=0), survival rate (Lx) showed records of death and the number of living snails left, expressed as fraction of 1 (initial number = 1 = 100%). Fecundity rate (Mx) was calculated as the number of birth or fertile eggs laid per unit time by snail of pivoted age (x).

Multiplication of survival rate (Lx) by fecundity rate (Mx) gives LxMx and the summation of all LxmX gives the net reproductive rate (Ro). The capacity for increase was derived from the reproductive rate as follows, R c = Loge Ro Tc Where: rc = capacity for increase, Tc = Generation time Ro = Net reproductive rate Tc is the mean time from birth of parents to birth of offspring and calculated as follows; Tc = LxMxX = LxMxX LxMx GROWTH The increase in shell length of the laboratory bred Bulinus species was used as an index of growth. The snail shell length (SSL) was measured from day of hatching to the 20-30th week bi-weekly to the nearest 0.01mm using the vernier caliper. (ii) VIABILITY Six hundred eggs of each Bulinus species were incubated till all viable eggs hatched. Any egg which failed to hatch within 3 days after the last hatch was regarded as not viable. The percentage hatch was then calculated. RESULTS FECUNDITY RATES The four Bulinus species had different egg production rates at different periods (Table 1). Fecundity rates increased gradually from the point of oviposition (6-7 weeks) having several peaks and falls and decreased gradually till death. B globosus began oviposition by week 7 with 41 egg/snails and rose to 221.34 eggs/snail at week 12 while the highest rate was 267.76 eggs/snail (week 20). For B trancatus, oviposition also began at week 7 with the peak at Ro

week 20 (231.0 eggs/snail). In both snails, oviposition showed other lower peaks which decreased gradually till death (Table 1). B forskali and B senegalensis showed similar oviposition rate which were slightly different from B globosus and B trancatus. B forskalli and B. senegalensis began oviposition by week 6 and had the peak fecundity at week 18 with 224.81 and 217.91 eggs/snail respectively. There was less fluctuation in fecunditry rates after the peak which was followed by a gradual fall (Table 1). B. globosus had the highest net reproductive rate (Ro) of 965.6, followed by B truncatus (762.84) and .B senegalensis had the least (541.0). The observed Tc varied from 132.09 in B. globosus to 97.88 in B. senegalensis (Table 2),while the rc was between 0.0539 (B. globosus) to 0.0643 (B. senegalensis). The Ro and Tc showed significant variation based on snails species (P = 0.05) while the rc did not vary significantly (P = 0.05). SNAIL GROWTH The growth rate of the four Bulinus species showed; a similar pattern: Hatclings ranged from 0.65mm (B. globosus) to 0.50 (B. senegalensis). There was gradual increase in growth rate from week 1 to week 10 in each snail (Table 3), This was followed by a decline from week 12. B globosus and B truncatus reached 13.7mm and 12.5mm at week 20, with each reaching the greatest SSL at week 36 with 14.3 mm and 13.5mm respectively (Table 3). For B. senegalensis and B. foskalli, steady increase in growth rate was also observed from weeks 1 to 7, followed by a decline from week 10. Using, Walford plot Maximum SSL was reached by week 26 with 11.4mm and 10.7mm for B. forskalli and B. senegalensis respectively (Table 3). formula of Lt = 1 on Lt, the line describing this relationship intercepted at 45 0 diagonal at Lt values of 10.3mm (B. globosus), 10.05 (B. truncatus), 9.6mm (B. forskalli) and 9.4mm (B. senegalensis).

(iii)

MORTALITY

The Bulinus species showed the highest mortality rate at weeks 1-2 (Table 4). From week 4 to week 18 death rate of snails remained similar and gradual for B globosus and B trancatus till death.Similarly the death rate of B forskalli and B senegalensis was also high between weeks 1 to 2, followed by a sharp and steep rate of death from weeks 18-20 respectively (Table 4) (vii) INCUBATION PERIOD B. globosus and B. truncatus periods of incubation were between 7 to 14 days while those of B. forskalli and B. senegalensis were 6 to13 days. Hatching began on the 6th day for B. forskalli and B. senegalensis and on the 7th day for B. globosus and B. trunacatus. Any egg which failed to hatch after 3 days at the end of the longest incubation periodwas regarded as not viable. The difference was not statistically significant (Table 5 & 6). DISCUSSION The life table studies of the four Buliniid snails were based on the net reproductive rate (Ro), generation time (Tc) and capacity for increase (Rc). This work shows that the rc of the four Buliniid snails ranged from 0.054 to 0.064. These values are slightly lower than the reports of Okafor (1984) and Anya & Okoronkwo (1991b) who reported 0.65 and 0.06 respectively for B. globosus which are not much different. From the summation of the Ro, and Tc which culminated in rc, B. senegalensis had the highest potential to increase, followed by B. forskalli, B. trunactus and B. globosus. The observations in this work indicated the interaction of all the life table factors to produce a similar rc. This shows a natural balancing effect of the Tc, Ro and rc, meaning that each species has similar potential with any other one to be found in the Quarry pit water barring other selective effects. The hatchability studies showed that B. senegalensis laid the most viable eggs as only 10% were not viable, compared to B. forskalli, B. truncatus and B. globosus. Okoronkwo (1986) reported that 18.7% and 16.18% of B.
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truncatus and B. globosus were not viable as some were empty egg-masses without embryoes. This observation further confirms that some eggs of Bulinus species are not viable and can not contribute to snail population. Similarly, incubation periods of the buliniid snails did not vary much but agreed with the observations of Freyer et al (1990), Okafor (1984) and Anya & OKoronkwo (1991b) who reported 6 to 12 days and 6-14 days. This observation implies that eggs laid within the same period will hatch within the same time and juveniles found are of similar age .The non-viable eggs therefore reduce the potential capacity to increase by the laying snail. Hatchlings of the Buliniid snails ranged from 0.50 0.65mm. These values are similar to the 0.6mm and 0.65mm reported by Okafor (1984) and Anya & Okoronkwo (1991b). The growth pattern observed in the snails studied was similar as growth rate was geometrical in the first 5-8 weeks of life before the rate decreased drastically. Similarly Woolhouse and Taylor (1990) and Fryer et al (1990) had reported similar growth pattern. This implies that from week 7, the snails were beginning to lay eggs.Growth materials therefore were diverted to egg production. On the other hand, Doumbo et al (1992) and Raymond & Probert (1993) had reported decreased growth just before oviposition. This could be attributed to the conservation of food materials for the would-be-hatchlings in the eggs. The gradual or slow growth rate observed during oviposition, showed that small amount of materials were still reserved for growth, an indication that a change of physiological and biochemical activities from growth to oviposition occured Results of this study on the mortality rate indicated that most of the four Bulinus species died within the first 2 weeks of life. Similarly another high death rate periods were recorded between weeks 18-24 for B forskalli and B senegalensus and between weeks 20 & 26 for B globosus and B truncatus. This observation suggests that hatchlings are faced with difficult environmental

conditions in the 1-2 weeks of life after hatching, and if they survive, will face another tough time by weeks 18-22 of life. In conclusion therefore, B. globosus was found to be highly adapted to the Quarry pit water followed by B. truncatus, B. forskalli and then B. senegalensis. A comparison of values in this work to other researchers values, showed that no focus of snail population is ideal rather each focus varies in its own ways (Fryer et al, 1990, Klumpp and Chu, 1977, Agi 1995b).

REFERENCES Agi, P.I. (1995b) Distribution of fresh water snail vector of schistosomiasis and study of physico-chemical parameters in Ogoni Communities (Rivers State Nigeria. Acta Hydrobiologica, 37(2)69076. Anya, A.O. & Okoronkwo, F.O (1991b) Studies of infection with Schistosoma haematobium and its effects on the reproductive biology of Buliniid snail in South Eastern Nigera. Proceedings of Nigerian Academy of Science 3:1425. Betterton, C. & Fryer, S.E. (1982) S. haematobiun in Northern Nigeria; A complex of snail hosts and parasite strain. Parasitology 85(2):1-5 (suppl xxii). Chu, K.Y.Kpo H.K.& Klump. K.K (1978) Mixing of S. haemotobium strains in Ghana. Bulletin of World Health Organization 56:601-608. Cowper. S.G.(1963). Schistosomiasis in Nigeria. Annals of Tropical Medicine and parasitology 57:307-322. Doumbo, O. Dabo, A. Diallo, M. Doucaure, B. Akory A. Balique, H.& Quilici. M. (1992). Epidemiology of human urban schistosomiasis in Bamko, Mali Medicine in Tropics 52(4):427-434. Emejulu, A.C. Albaronye, F.F Ezenwaji H. M.G. & Okafor F.C. (1994). Investigations into the prevalence of urinary schistosomiasis in the Agulu lake are of Anambra State, Nigeria. Journal of Heliminhology 2:119-123. Fryer S.E.oswold, R.S. Probert, A.J & Runham, N.W. (1990). The effect of S. haematobium infections on the growth and fecundity of 3 symmetric species of Buliniid snail. Journal of parasitology 76(4) 557-563. klumpp, R.K. & Chu K.Y. (1997). Ecological studies of Bulinus truncatus, the intermediate host of S. haematobium in the Volta Lake. Bulletin of World Health Organization 55(6) 715-736.
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Nduka F.D.Ajaero C.M.U. Nwoke, B.E.B. (1995). Urinary Schistosomisis among School children in an endemic community in South Eastern Nigeria, Applied parasitology, 36:34-40. Ndifon, G.I.(1983) Human helminthiasis in the Taga Lake basin, Kano, Nigeria. The Nigeria Society for Parasitology, VIITH Annual Conference Book of Abstract (Ed. Iwuala, M.O.E) 14-15. Okafor, F.C. (1984). The Eco-physiology and biology of snail hosts of Schistosoma haematobium Biharziasis, in Anambra State, Nigeria Ph.D. Thesis University of Nigeria, Nsukka. Okoronkwo, F.C.(1986). Host parasite relationship of S. haematobiunm and its snail hosts in Anambra State, Nigeria, Ph.D. Thesis University of Nigeria Nsukka. Raymond, K. & Probert A.D. (1993).The effect of infections with S. margrebowei on the growth of Bulinus natalensis. Journal of Helminthology 67 (1): 10-16. Savioli, L. & Mott. K.E. (1989). Urinary Schistosomiasis on Pemba Island, Lowcost diagnosis for control in primary Health care setting. Parasitology today 5(10) 333-337. Ukoli, F.M.A. & Asumu, D.I. (1980) Ecological Studies on snail relation to the transmission of Schistosomiasis and other snail borne-diseases in Kainji Lake, Nigeria, Journal of platythelminths 12(1) 59-68. WHO (1991). Schistosomiasis; Tropical Disease Progress in Research 19891990. tenth Programme Report. UNDP World Bank WHO Special Progress Report for Research (TDR) General 39-54. WHO (1994) Schistosomiasis; 20 years of progress, Highlights, 1993-19994. TDR 12TH Programmes Report UNDP World Bank/WHO special Report for Research (TDR) 77-86. Woolhouse, M.E.J.& Chandiwana, S.K. (1990). Temporal patterns in the Epidemiology of Schistosome infections of snails, a model for field data Parasitology; 100:247-253. Woolhouse, M.E.J & Taylor, P. (1990) Survival rates of Bulinus globosus during Aestivation Annals of Tropical Medicine and Parasitology 84(3): 293-295.

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TABLE 1: FECUNDITY RATES OF FOUR Bulinus-species STUDIED (Mx) Age in weeks (X) 1 2. 4 6 7 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 B. globosus B. truncatus B. forskalli B. senegalensis

41.0 + 0.02 72.56 + 0.03 136.12 + 0.03 221.34+ 0.04 132.57 + 0.06 224.48 + 0.06 202.42 + 0.06 267.26 + 0.11 137.92 + 0.03 114.79 + 0.04 92.20 + 0.06 162.20 + 0.07 86.34 + 0.07 51.62 + 0.08 35.31 + 0.08 20.14 + 0.06 10.10 + 0.07 6.10 + 0.08

35.1 + 0.04 61.23 + 0.06 127.47 + 0.05 201.06 + 0.05 112.81 + 0.06 207.56 + 0.07 187.57 + 0.08 231.51 + 0.07 152.12 + 0.06 120.72 + 0.06 106.23 + 0.08 118.10 + 0.07 24.20 + 0.03 16.31 + 0.03 10.21 + 0.04 8.12 + 0.05 6.51 + 0.05 -

32.1 + 0.10 38.44 + 0.10 44.23 + 0.09 92.31 + 0.08 141.57 + 0.08 142.31 + 0.06 212.43 + 0.05 224.81 + 0.05 103.71 + 0.05 60.81 + 0.06 72.19 + 0.04 21.91 + 0.04 8.43 + 0.05 6.52 + 0.06 -

30.13 + 0.04 45.20 + 0.05 62.13 + 0.05 80.43 + 0.05 131.14 + 0.04 146.23 + 0.04 187.47 + 0.04 217.91 + 0.05 163.91 + 0.06 100.27 +0.06 43.84 + 0.06 26.32 + 0.08 18.43 + 0.08 10.11+ 0.08 -

X = Age in weeks - = Age at which snail died. TABLE 2: SUMMARY OF THE REPRODUCTIVE BIONOMICS OF FOUR BULINIID SNAILS STUDIED Snail Bulinus globosis B truncatus B forskalli B senegalensis Key: Ro 965.6 762.84 582.51 541.0 To 132.09 122.68 11.27 97.88 Rc 0.0539 0.0541 0.0572 0.0643

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Ro = Net Reproductive rate. Tc = mean generation time Rc = capacity for increase

TABLE 3: GROWTH RATE OF THE FOUR Bulinus species MAINTAINED IN THE LABORATORY Age weeks (X) 0 2 4 6 8 10 12 14 16 18 in B globosus B truncatus B forskalli B senegalensis

0.65 + 0.618 1.8 + 0.014 3.2 + 0.07 8.8 + 0.15 9.0 + 0.03 9.6 + 0.08 10.4 + 0.17 11.3 + 0.06 12.1 +0.08 12.6 + 0.08

0.60 + 0.04 1.8 + 0.09 3.0 + 0.11 5.2 + 0.08 8.1+ 0.11 9.1 + 0.14 9.8 + 0.12 10.9 + 0.16 11.3 + 0.11 11.8 + 0.10
12

0.61+ 0.02 1.02 + 0.08 2.0 + 0.08 4.1 + 0.11 8.2 + 0.10 8.1 + 0.11 9.2 + 9.13 9.8 + 0.14 10.2 + 0.09 10.4 + o.19

0.50 + 6.02 0.80 + 0.11 1.65 + 0.14 3.0 + 0.15 6.1 + 0.11 6.4 + 0.26 8.0 + 0.19 9.0 + 0.21 9.4 + 0.20 9.6 + 0.21

20 22 24 26 28 30 32 34 36 38 40

13.0 + 0.05 13.2 + 0.08 13.4 + 0.04 13.7 + 0.06 13.8 + 0.06 13.7 + 0.116 13.8 + 0.12 13.8 + 0.09 13.8 + O.15 13.8 + 0.09 13.8 + 0.10

12.1 + 0.08 12.3 + 0.11 12.4 + 0.10 12.6 + 0.07 12.6 + 0.07 12.6 + 0.13 12.5 + 0.10 12.6 + 0.10 12 + 0.10 12 + 0.09

10.6 + 0.17 10.8 + 0.11 10.9 + 9.09 11.0 + 0.13 11.0 + 0.13 11.0 + 0.10

9.0 + 0.14 9.7 + 0.08 9.9 + 0.11 10.0 + 0.12 10.0 + 0.09 10.0 + 0.11

TABLE 4:

MORTALITY RATE ACCORDING THE MORTALITY FORMULA loge NT - loge NT + 1 OF THE VARIOUS Bulinus species

Age in weeks (x) 1 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40

B globosus 1 0.36 0.38 0.44 0.46 0.47 0.48 0.48 0.48 0.48 0.50 0.51 0.53 0.55 0.57 0.60 0.64 0.69 0.72 0.74 0.74

B truncatus 1 0.32 0.36 0.40 0.44 0.46 0.47 0.47 0.47 0.49 0.50 0.55 0.60 0.66 0.66 0.70 0.73 0.77 0.81 0.81

B forskalli 1 0.24 0.34 0.37 0.4 0.41 0.43 0.44 0.49 0.50 0.58 0.60 0.60 0.64 0.74 0.88

B senegalensis 1 0.20 0.26 0.32 0.38 0.40 0.42 0.45 0.58 0.59 0.59 0.64 0.89 0.90 0.94 0.94

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TABLE 5: VIABILITY OF THE LAID EGGS AS DETERMINED BY HATCHING OF THE EGGS SPECIES B globosus B fruncatus B forskalli B senegalensis NO OF EGGS 600 600 600 600 % HATCHED 82.8 (497) 82.8(485) 85.34(512) 89.8(539) % UNHATCHED 17.2(1030) 19.21(115) 14.66(94) 10.20(61)

TABLE 6: INCUBATION PERIODS OF THE FOUR Bulinus species Bulinus species B globosus B truncatus B forskalli B senegalensis Days of incubation 7-14 7-14 6-13 6-13

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