Layout 1 ARTICLE Introduction Crocodiles and other reptiles serve various purposes in trop- ical regions, where they are utilized for food, traditional medicine, and ornamental items (Alves et al., 2006, 2008, 2009, 2013). In areas where crocodilians coexist with human populations, local attitudes towards them can range from tolerance to hostility. De- spite this, crocodiles can offer economic value through regulated harvesting, particularly in the skin trade (Caldwell, 2014). Beyond material benefits, communities may be encouraged to conserve crocodiles due to their ecological significance in sustaining wet- land ecosystems (Van der Ploeg et al., 2011). In parts of West Africa, rural populations show reverence for crocodiles, often protecting them due to traditional beliefs. In cer- tain cultures, crocodiles are symbolically linked to water avail- ability, where their disappearance is feared to result in water scarcity (Kpéra, 2003; Kpéra et al., 2004). The dwarf crocodile (Osteolaemus tetraspis), endemic to Africa, is the smallest extant crocodile species, easily recognized by its compact size, flattened skull, narrow snout, and characteristic body armor. It inhabits swamps, marshes, and dense forests, preying on aquatic organisms like fish, snails, and crabs. Adults typically grow up to 1.9 meters and weigh between 18 to 32 kilograms (Pauwels et al., 2007; Dou- glas, 2010; Sprawl et al., 2012; Shirley et al., 2016). Anatomical adaptations, such as thick body armor and osteoderms, enhance Species diversity and distribution of dwarf crocodile (Osteolaemus tetraspis) in bushmeat markets in Benin metropolis, Edo State, Nigeria Israel O. Ogorode,1 Aroloye O. Numbere,2 E. Abiodun Adams,3 Ijeoma F. Vincent-Akpu,2 Bolaji B. Babatunde,2 Francis Sikoki1 1Department of Biological Sciences, University of Africa, Toru-Orua, Bayelsa State; 2Department of Animal and Environmental Biol- ogy, Faculty of Science, University of Port Harcourt, Choba, Rivers State; 3Department of Biological Sciences, Faculty of Science, National Open University of Nigeria, Headquarters, Jabi, Abuja, Nigeria ABSTRACT This study investigated the diversity and distribution of dwarf crocodiles in eight bushmeat markets in Benin City, Edo State, Nigeria. The markets surveyed were New Benin, Oliha, Uselu, Oba Market, Ekiuwa, Ugbighokogho, Evbotubu, and Third Market between Oc- tober 2022 and May 2023. A multi-method approach was employed, combining observation and measurement techniques. The Shan- non-Wiener Index (H'), Simpson's Index (D), and Species Evenness (E) were calculated to determine the diversity and distribution of dwarf crocodiles. The results showed that Ekiuwa and Ugbighokogho markets had the highest Shannon-Wiener Index (H') values of 0.84 and 0.86, respectively, indicating higher species diversity. These markets also had the lowest Simpson's Index (D) values of 0.41 and 0.39, respectively, indicating lower species domi- nance. Additionally, Ekiuwa and Ugbighokogho markets had the highest Species Evenness (E) values of 0.92 and 0.94, respec- tively, indicating a more even distribution of individuals among species. However, when considering the monthly variation, Oliha, Ugbighokogho, and 3rd Market had the highest Shannon- Wiener Index (H') values of 1.05, 1.06, and 1.05, respectively, indicating higher species diversity. The results suggest that some markets have higher species diversity and evenness, indicating a more sustainable trade. The study recommends that conserva- tion efforts focus on regulating the trade in dwarf crocodiles and promoting sustainable harvesting practices. Corresponding author: Israel Omoghene Ogorode, Department of Biological Sciences, University of Africa, Toru-Orua, Bayelsa State, Nigeria. E-mail: israel.ogorode@uat.edu.ng Key words: Osteolaemus tetraspis, bushmeat markets, species di- versity, Shannon-Wiener Index, Benin metropolis. Authors’ contributions: all the authors made a substantive intellec- tual contribution. All the authors have read and approved the final version of the manuscript and agreed to be held accountable for all aspects of the work. Conflict of interest: the authors declare no potential conflict of in- terest. Funding: none. Availability of data and materials: all data generated or analyzed during this study are included in this published article. Acknowledgments: we are grateful to the Departments of Animal and Environmental Biology of the University of Port Harcourt, Choba Rivers State, Nigeria, and Delta State University, Abraka, Delta State, Nigeria for providing the field and laboratory materials for this research work. More so, we appreciate the Area chiefs and market women for granting us express permission to carry out this survey. Received: 12 May 2025. Accepted: 17 October 2025. Publisher’s note: all claims expressed in this article are solely those of the authors and do not necessarily represent those of their affil- iated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article or claim that may be made by its manufacturer is not guaranteed or endorsed by the publisher. ©Copyright: the Author(s), 2025 Licensee PAGEPress, Italy Advances in Oceanography and Limnology, 2025; 16:13976 DOI: 10.4081/aiol.2025.13976 This work is licensed under a Creative Commons Attribution- NonCommercial 4.0 International License (CC BY-NC 4.0). I.O. Ogorode et al.46 its survival. The species’ cranial structure is similar to the Cuvier dwarf caiman, suggesting ecological convergence. It also displays a unique dental formula, including 4 premaxillary and 12-13 max- illary teeth (Walmsley et al., 2013). Two subspecies exist: the West African dwarf crocodile (O. tetraspis) and the Congo dwarf crocodile (O. osborni), each adapted to distinct environments within tropical West and Central Africa. These habitats, rich in aquatic and terrestrial biodiversity, support the survival and reproduction of O. tetraspis, which prefers forested and shaded areas and avoids large water distur- bances (Sprawl et al., 2012; Amoah et al., 2021). Cave use, ob- served in western Gabon, may serve as a mechanism to maintain genetic diversity (Gabbatiss, 2016). Primarily nocturnal, O. tetraspis forages at night and remains concealed during daylight in burrows or beneath tree roots. This behavior minimizes expo- sure to predators and environmental stressors (Numbere & Chimezie, 2020). Its diet is flexible and varies regionally—pri- marily fish in the Democratic Republic of Congo, and more com- monly snails and crabs in Nigeria. Plant matter occasionally appears in their diet, likely ingested incidentally (Pauwels et al., 2007; Sprawl et al., 2012). Breeding occurs during the rainy sea- son (May-June), with females constructing nests of decaying veg- etation near water. Eggs hatch after 85-105 days, and hatchlings measure around 28 cm. Clutches typically contain up to 10 eggs, although some can hold as many as 20 (Somaweera et al., 2013). Both O. tetraspis and O. osborni are classified as vulnerable by the IUCN. However, population data remain limited due to insuf- ficient research. Habitat degradation and hunting for bushmeat are believed to be the primary threats. Despite being found in sev- eral protected areas, the species faces local extinction risks in countries like Gambia and Liberia. Although the demand for their skins is relatively low, they are still hunted for subsistence pur- poses (Smolensky, 2015). These crocodiles are kept in captivity in North America and Europe, where they are commonly bred in zoos. However, taxo- nomic uncertainties persist, particularly in regions like Nigeria, where many captive individuals may be hybrids (Shirley et al., 2015). Although there is documented folklore about crocodiles in West Africa (Kpéra et al., 2014), detailed knowledge of human- crocodile relationships remains sparse (Akani et al., 1999; Aust, 2009; Pooley, 2016; Rice, 2018; Brackhane et al., 2018; 2019; Somaweera et al., 2019). While studies of crocodile ecology are growing in West Africa (Shirley et al., 2009, 2018), particularly in Nigeria (Luiselli et al., 1999a, 1999b, 2012), there is a signifi- cant gap in research on local perceptions regarding crocodile abundance and their utility for food or income. Therefore, this research paper aims to investigate the species diversity and distribution of Osteolaemus tetraspis within the bushmeat markets in Benin Metropolis. Materials and Methods Study area Benin City serves as the capital and largest metropolitan cen- ter of Edo State, situated in southern Nigeria. Notably, it ranks as the fourth-most populous city in Nigeria, according to the 2006 national census, preceded only by Lagos, Kano, and Ibadan. Benin City is located near the Benin River, situated approximately 40 kilometres (25 miles) to the north, whilst its eastern perimeter lies 320 kilometres (200 miles) from Lagos via the arterial road net- work. The city’s municipal boundaries converge with those of sev- eral prominent neighboring towns in southern Nigeria (Fabolude & Aighewi, 2022), notably Agbor, Oghara, and Ekpoma. Benin City boasts an exceptionally fertile agricultural landscape and serves as the epicentre of Nigeria’s thriving rubber industry (Ojo, 2024). Additionally, the production of palm oil constitutes a sub- stantial sector, further underscoring the city’s prominence in Nige- ria’s agricultural economy. Benin has a population of over 10 million. Around 70% of the working population depends on agriculture. Benin has pro- duced yams, cassava, corn (maize), millet, beans, and rice since the mid-1980s to achieve self-sufficiency in these staple foods. The vegetation regions can be divided into five major types: for- est, grassland, tundra, desert, and ice sheet. Climate, soil, the abil- ity of soil to hold water, and the slope, or angle, of the land all determine what types of plants will grow in a particular region. Benin City exhibits a borderline tropical savanna climate (Köppen: Aw), closely approaching a tropical monsoon climate (Am). The city’s climatic conditions are characterized by persist- ently high temperatures and humidity levels throughout the year (Aruya-Emmanuel, 2020). Edo state where Benin City is situated has a tropical wet and dry or savanna climate. The city’s yearly temperature is 28.78°C (83.8°F) and it is -0.68% lower than Nigeria’s averages (Aruya- Emmanuel, 2020) The weather in Benin City is typically hot and humid, with average temperatures ranging from 25 to 30°C (77 to 86°F) throughout the year. The relative humidity is consistently high, often exceeding 60% (Balogun et al., 2023). While the city experiences a relatively uniform climate throughout the year, there is discernible seasonal variation. The period between July and September is particularly notable for its dull and overcast weather conditions, with reduced sunshine and increased precipitation (Odiana & Idahosa-Ohio, 2023; Floyd et al., 2016) (Figure 1). Study site/site selection This study was carried out across eight bush-meat markets (New Benin, Oliha, Uselu, Oba Market, Ekiuwa, Ugbighokogho, Evbotubu, and Third Market) within the Benin Metropolis, Edo State. Benin City, situated in the south-south geo-political region of Nigeria, is a major urban hub with coordinates 6.3350° N, and 5.6037° E, as shown in Figure 1 above. It is about 64 kilometres from the Gulf of Guinea, and it serves as a significant industrial and cultural center. The city’s economy thrives on rubber and oil production (Ikponmwosa, 2020), with a population nearing 1.2 million (Balogun & Onokerhoraye, 2017). Benin City’s residents predominantly trace their heritage to the rich Edo culture. Visitors can explore various attractions, including historic palaces, muse- ums, lush parks, and vibrant cultural festivals that take place throughout the year (Ogoanah and Omejife, 2017). Data collection and sampling technique Eight to ten crocodile traders each were sampled from the eight bushmeat markets. The bushmeat markets include New Benin, Oliha, Uselu, Oba Market, Ekiuwa, Ugbighokogho, Evbo- tubu, and Third Market. This study employed a multi-method approach, combining observation and measurement techniques. At the bushmeat mar- ket, the number and species of crocodiles were counted and Dwarf crocodile diversity In Benin Metropolis Markets 47 recorded. To facilitate measurement, graduated sticks and tape rules were used, enabled by the fact that many crocodiles were restrained or weakened (Eniang et al., 2020). The measured crocodiles were categorized into three size groups: Small Sized Dwarf Crocodile (SSDC, 50-60 cm or approximately 2ft), Medium Sized Dwarf Crocodile (MSDC, 70-90 cm or approxi- mately 3ft), and Large Sized Dwarf Crocodile (LSDC, 100-120 cm or approximately 4ft) (San-Diego, 2015). Additionally, in- terviews and focused-group discussions involving recorded oral interviews were conducted to understand how marketers acquire, preserve, and trade these crocodile species, whether dead or alive, until they are sold (Ogorode et al., 2023; Eniang et al., 2020). The dwarf crocodiles shaded for sale are represented in Figure 2 and 3. Figure 1. Map of study area showing the metropolitan locations which includes: Oredo, Egor, Ikpoba-Okha Ekenwan, Etsako-Wedt Ovia North East with Ikpoba Slope,a long sloppy hill in the middle of the town, Benin City and a large River with a bridge at the extreme. Each of the locations has a market where bushmeats such as crocodiles, Pangolins, antelopes, land snails and so on are sold as represented on the map. Figure 2. Dwarf crocodile shaded for sale at the Oliha Market, Benin City. The species shaded on the table are tied with ropes on their legs to ensure that they cannot move from one position. The mouth is opened and castrated with a stick then tied with a rope to avoid dangerous bites on customers and the sellers. This captivity technique makes the animal weak and become nearly dead on the table. The species is black in color with armored skin for protection and defense. Figure 3. Freshly supplied dwarf crocodile shaded at Ekiuwa market for sale. The species are lined up according to their sizes Being tied with ropes on their legs to their body to avoid move- ment and mouth castrated with sticks to avoid biting. The tails of the species are well decorated with sharp blades which are flipped at random when still very active. They are shaded on the table for visibility. Other animals like land snails, pangolins and antelopes are shaded and hanged for visibility close to the crocodile. I.O. Ogorode et al.48 Data analysis/statistical analysis The Shannon-wiener index (H) representing diversity, Simp- son index (D) representing dominance of species, and Species evenness (E) representing distribution among species were em- ployed to analyse the data collected: Shannon-Wiener Index (H’) H’ = - ∑ (ni/N) * ln(ni/N) where ni is the abundance of species i, and N is the total abun- dance. Simpson’s Index (D) D = ∑ (ni/N)² Species Evenness (E) E = H’/ln(S). where S is the number of species. Results From Table 1, a total of 8 traders were interviewed at inter- vals in each market, providing valuable insights into the bush- meat trade. According to the traders, the bushmeat markets have been in existence for over a century, with 2 traders confirming this. The traders also reported that they bring bushmeats and crocodiles to the markets frequently, depending on sales and re- quests by traders. This was confirmed by 4 traders. In terms of preservation methods, 3 traders reported that dead crocodiles are preserved through roasting or smoking. The traders also pro- vided information on the sales volume of crocodiles. According to 5 traders, the weekly average sales range from 5 to 10 croco- diles. The study also investigated the uses of crocodile species. According to 5 traders, the crocodile species is used largely for medicinal purposes, with a few used for food and very few resold for money. The traders reported that they obtain croco- diles from hunters and fishermen and sometimes travel to the riverside. This was confirmed by 5 traders. From Table 2, the Calculated H’ values for the diversity of species in different markets include: New Benin (0.69), Oliha (0.73), Uselu (0.63), Ekiuwa (0.84), Ugbighokogho (0.86), Evbo- tubu (0.65), 3rd Market (0.71) while the calculated D values in the market read: New Benin (0.53), Oliha (0.51), Uselu (0.58), Ekiuwa (0.41), Ugbighokogho (0.39), Evbotubu (0.56), 3rd Mar- ket (0.52). The Calculated E (Species evenness) values read: New Benin (0.83), Oliha (0.85), Uselu, (0.79), Ekiuwa (0.92), Ug- bighokogho (0.94), Evbotubu (0.81), Third Market (0.84). The re- sults suggest that: Ekiuwa and Ugbighokogho markets have the highest Shannon-Wiener Index (H’) values, indicating higher species diversity, Secondly, Ekiuwa and Ugbighokogho markets also have the lowest Simpson’s Index (D) values, indicating lower species dominance. Ekiuwa and Ugbighokogho markets have the highest Species Evenness (E) values, indicating a more even dis- tribution of individuals among species. From the results obtained, Ekiuwa and Ugbighokogho stand out. From Table 3, the Calculated H’ values for diversity of the dwarf crocodile species read: New Benin (1.03), Uselu (1.01), Oba (1.02), Ekiuwa (1.04), Oliha (1.05), Ugbighokogho (1.06), Evbotubu (1.04), Third Market (1.05). The Calculated D values for the Simpson index for dominance of dwarf crocodiles read: New Benin (0.55), Uselu (0.57), Oba (0.56), Ekiuwa (0.54), Oliha (0.53), Ugbighokogho (0.52), Evbotubu (0.54), Third Market (0.53). The Calculated E values for species evenness read: New Benin (0.83), Uselu (0.81), Oba (0.82), Ekiuwa (0.84), Oliha (0.85), Ugbighokogho (0.86), Evbotubu (0.84), Third Market (0.85). The results suggest that Oliha, Ugbighokogho, and Third Market have the highest Shannon-Wiener Index (H’) values, in- dicating higher species diversity. Oliha, Ugbighokogho, and Third Table 1. Oral questions in bushmeat markets in Benin metropolis, Benin City, Edo State, Nigeria. S/N Oral questions General response presented Numbers of traders’ by traders interview at intervals in each market (sampling number of 8) 1 How long has these Bushmeat Markets been in existence? Over a century 2 2 How often do they bring bushmeats and crocodile to Frequently depending on sales/request by traders 4 these markets? 3 How do you preserve the crocodile species particularly Dead ones are roasted/smoked 3 the near dead ones? 4 How many crocodiles do you sell daily or weekly? 5-10 (weekly average) 5 5 What do people use the crocodile species for? Largely medicine 5 Few for food Very few resell for money 6 Where do you get the crocodile species from? Hunters and fishermen/transport to riverside 5 Table 2. Abundance of dwarf crocodiles in Bush meat markets in Benin City Edo State, Nigeria (2016-2017). Year New Benin Oliha Uselu Ekiuwa Ugbighokogho Evbotubu Third market 2016 20 20 10 100 120 10 20 2017 14 18 12 126 116 15 12 Source: Ogoanah & Omijie, 2017. Dwarf crocodile diversity In Benin Metropolis Markets 49 Market also have the lowest Simpson’s Index (D) values, indicat- ing lower species dominance the dwarf crocodiles. Oliha, Ugbighokogho, and Third Market have the highest Species Evenness (E) values, indicating a more even distribution of individuals among the dwarf crocodile species (Figure 2). Discussion The oral interview on the bushmeat markets in the metropolis yielded insightful results regarding the traders’ perceptions and operational practices. The first question queried the longevity of these bushmeat markets. The responses indicated that these mar- kets have been in existence for over a century, reflecting their deep-rooted presence in the community. This longstanding tradi- tion suggests that the trade of bushmeat, particularly crocodile species, is not only a means of livelihood but also part of the cul- tural fabric of the region, which is evident in recent literature (Petrozzi et al., 2016; Trefon, 2023). When asked how frequently traders supply bushmeats and crocodiles to the market, the con- sensus was that this process occurs frequently, often depending on sales and specific requests from traders. This variability in fre- quency indicates a responsive supply chain that adapts to market demands, as implied by McNamara et al. (2016) and Walenlign et al. (2019). Preservation methods for crocodile species, espe- cially those in a near-death state, were discussed, revealing that traders typically roast or smoke the meat. The smoking process not only aids in preservation but also enhances the flavor profile, making it more appealing to customers. The preservation tech- nique showcases the traders’ adaptability and their efforts to min- imize waste in a market where the freshness of the product is vital (Cowlishaw et al., 2005; Ahmadi et al., 2018), In terms of sales, traders reported an average sale of five to ten crocodiles weekly. This figure underscores the interest and de- mand for crocodile meat in local markets, indicating a sustainable level of consumption where both traders and consumers find value (Thorbjarnarson, 1999) The most significant use of crocodile species, as articulated by the traders, is in medicine, with a smaller fraction being utilized for food and even fewer being resold for profit (Moshoeu, 2017) This finding highlights the cultural and medicinal significance attributed to crocodiles, which may drive ongoing demand despite potential ethical or conservation con- cerns surrounding the trade. The source of crocodile species pri- marily stems from hunters and fishermen, with transportation occurring to riverside locations before reaching the markets. This sourcing method illustrates the interconnectedness of local com- munities and natural resources, emphasizing the need for sustain- able practices to protect both wildlife and the livelihoods that depend on them (Burgos-Ayala et al., 2020) The findings of this study indicate that Ekiuwa and Ug- bighokogho bushmeat markets exhibit higher species diversity, lower species dominance, and a more even distribution of indi- viduals among species. These results are consistent with recent studies on dwarf crocodiles, which have highlighted the impor- tance of habitat diversity and evenness in maintaining a healthy population (Riley & Huchzermeyer, 2009). It was also found that dwarf crocodiles in habitats with higher vegetation diversity and evenness had higher population densities and greater genetic di- Table 3. Species abundance of dwarf crocodile in Bushmeat Markets in Benin Metropolis Edo State Nigeria (2022-2023). Markets Sizes of Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Total dwarf 2022 2022 2022 2023 2023 2023 2023 2023 crocodile New Benin Market LSDC 3 0 3 0 0 3 0 0 6 MSDC 2 5 2 0 0 1 3 2 15 SSDC 1 2 0 1 1 0 1 4 10 Uselu Market LSDC 0 1 4 2 1 0 0 1 9 MSDC 2 3 0 1 0 0 3 1 10 SSDC 1 0 0 0 3 0 1 1 6 Oba MarketMarkets LSDC 0 1 2 0 5 0 0 1 9 MSDC 0 2 0 0 2 4 2 1 11 SSDC 1 3 0 5 1 3 4 1 15 Ekiuwa Market LSDC 12 3 5 5 7 4 7 7 50 MSDC 8 5 7 7 6 5 5 14 57 SSDC 5 2 4 3 2 6 3 10 35 Oliha Market LSDC 3 3 1 1 0 3 2 2 15 MSDC 4 4 0 4 6 1 1 1 20 SSDC 2 1 1 2 2 3 1 2 14 Ugbighokogho Market LSDC 3 1 1 3 1 3 3 1 16 MSDC 3 4 3 0 2 2 3 2 19 SSDC 2 2 4 1 3 2 1 3 17 Evbotubu Market LSDC 3 2 3 1 2 1 2 1 14 MSDC 3 4 0 3 5 2 0 2 19 SSDC 5 3 1 2 1 1 2 3 18 Third Market LSDC 4 1 0 1 2 0 2 3 13 MSDC 4 1 1 1 1 3 2 3 16 SSDC 1 1 3 0 1 1 2 0 9 LSDC, large sized dwarf crocodile; MSDC, medium sized dwarf crocodile; SSDC, small sized dwarf crocodile. I.O. Ogorode et al.50 versity (Van Asch et al., 2019; Cao et al., 2020). Similarly, a study by Eaton et al., (2009) reported that dwarf crocodiles in areas with more even distributions of resources had lower levels of compe- tition and aggression. The high species diversity and evenness values observed in Ekiuwa and Ugbighokogho markets may be attributed to the prox- imity to Okomu National Park where the presence of suitable habitats and resources, such as vegetation, water, and prey are ob- tainable (Thorbjarnarson and Eaton 2004; Eaton and Barr 2005; Eaton 2006; Shirley 2007; Shirley et al. 2009). These markets may serve as important hubs for dwarf crocodile populations, pro- viding connectivity and facilitating gene flow between different habitats. The research results indicate relatively low diversity, domi- nance, and evenness values for dwarf crocodile populations across eight different locations. While some locations show slightly higher values than others, the overall range is narrow, suggesting a fairly homogeneous distribution of species within these populations (Eaton et al., 2009; Bouquin, 2008; Van Asch et al., 2019) Shannon-Wiener Index (H’): The H’ values (ranging from 1.01 to 1.06) are quite low. Most ecological studies reporting di- versity indices for reptile populations, especially those with lim- ited species richness, will often show higher values (Ficetola et al., 2013; Pincheira-Donoso et al., 2013). Studies on other croc- odilian species, or even broader herpeto-faunal surveys, frequently report H’ values exceeding 2 or even higher depending on habitat complexity and species richness (Kafash et al., 2020). This low H’ suggests that the dwarf crocodile populations studied might be dominated by a single or very few species, even if they appear di- verse at a coarse level. Recent literature emphasizes the impor- tance of functional diversity within ecosystems by these megafauna species (Malhi et al., 2016; Pimiento et al., 2020). Crocodiles and alligators play a vital role in maintaining healthy ecosystems, and their decline could have far-reaching conse- quences, according to a recent study (Eversole et al., 2015). These large reptiles contribute to ecosystem balance by creating complex burrow systems that provide shelter for other animals and by prey- ing on invasive pests that damage crops (Somaweera et al., 2020). However, up to 38% of the ecological functions performed by crocodilian species are under threat, highlighting the need for en- hanced conservation efforts. Other researchers have consistently shown that large predators particularly, crocodiles play a vital role in maintaining the balance and health of ecosystems on land and in the ocean (Heithaus et al., 2008a; Ritchie et al., 2012; Ripple et al., 2014). Simpson’s Index (D): The D values (0.52 to 0.57) are indica- tive of relatively low dominance. A value closer to 1 indicates dominance by a single species. The values obtained suggest a rea- sonably even distribution, albeit still within a narrow range. This aligns with the low H’ values, implying a potentially limited num- ber of species contributing to the community structure. According to Evans (2016), who studied the impact of habitat fragmentation on crocodilian populations was of the view that fragmentation could lead to reduced diversity and increased local dominance, even if overall regional diversity is higher. Habitat fragmentation poses a significant threat to biodiversity worldwide (Fahrig 2003; Fletcher et al. 2018; Ewers & Didham 2006). When habitats are destroyed or degraded, they often break into smaller, isolated frag- ments, disrupting delicate ecosystem balances and imperiling species survival (Raghubanshi & Tripathi 2009). The widespread clearance of natural forests and subsequent fragmentation of re- maining areas has become a pressing global concern (Echeverría et al. 2006, 2008). Species Evenness (E): The E values (0.81 to 0.86) show rel- atively high evenness. This suggests that the few species present are fairly evenly distributed in terms of abundance within each location. However, the high evenness coupled with low diversity suggests a community that is likely not highly complex. This was supported in Pyron, (2010) that ecological communities comprise various species coexisting in the same space and time. Re- searchers have identified three diversity levels within these com- munities: Local diversity which refers to the variety of species within a specific habitat, habitat transition diversity which: meas- ures changes in species diversity across different habitats or en- vironmental gradients and regional diversity which encompasses both local and transition diversity, providing a broader landscape perspective (Bestelmeyer et al., 2003; Pavoine & Bonsall, 2011). Enhanced conservation efforts are necessary to protect dwarf croc- odile populations and their habitats. Habitat preservation and restoration, education and outreach programs, and sustainable management of bushmeat trade are crucial for maintaining ecosys- tem balance and biodiversity. Conclusions This study emphasizes the importance of habitat diversity and evenness in maintaining healthy dwarf crocodile populations and highlights the need for enhanced conservation efforts. Recommendations The following recommendations were made: i) conduct eco- logical assessments of wild Osteolaemus tetraspis populations; ii) implement sustainable harvesting guidelines, particularly in biodiversity hotspot markets; iii) promote alternative livelihoods and conservation education to reduce dependence on crocodile trade; iv) engage local communities and traders in co-managed conservation initiatives; v) establish monitoring programs to track trade volumes and species diversity across markets. References Ahmadi S, Maman S, Zoumenou R, et al. Hunting, sale, and con- sumption of bushmeat killed by lead-based ammunition in Benin. International Journal of Environmental Research and Public Health 2018;15:1140. Akani GC, Luiselli L, Politano E. Ecological and conservation considerations on the reptile fauna of the Eastern Niger Delta (Nigeria). Herpetozoa 1999;11:141-53. Alves RRN, Filho GAP, Lima YCC. Snakes used in ethnomedi- cine in North-east Brazil. Environmental Development and Sustainability 2006;9:455-64. Alves RRN, Léo Neto NA, Santana GG, et al. Reptiles used for medicinal and magic religious purposes in Brazil. Applied Herpetology 2009;6:257-74. Alves RRN, Vieira WLS, Santana GG. Reptiles used in traditional folk medicine: conservation implications. Biodiversity and Conservation 2008;17:2037-49. Dwarf crocodile diversity In Benin Metropolis Markets 51 Alves RRN, Vieira WLS, Santana GG, et al. Herpetofauna used in traditional folk medicine: conservation implications. In: RRN. Alves, IL Rosa (eds.), Animals in Traditional Folk Med- icine. Springer; Berlin, Germany; 2013. Amoah E, Danquah E, Ross JP. Nesting ecology of West African dwarf crocodiles in a heavily disturbed landscape in Chire- hin, Ghana. International Journal of Ecology 2021. Available from: https://onlinelibrary.wiley.com/doi/10.1155/2021/ 8871631 Aruya-Emmanuel I, Ariko DJ. Assessment of spatial variation of urban canopy heat island in the wet season in Benin City. Gombe Journal of Geography and Environmental Studies 2020;1. Aust PW. The ecology, conservation and management of Nile crocodiles Crocodylus niloticus in a human dominated land- scape. Division of Biology, Imperial College London, UK, 2009. Balogun TF, Onokerhoraye AG. Spatio-temporal growth of Benin City, Nigeria, and its implications for access to infrastructure. Journal of Geography and Geology 2017;9:11-23. Balogun VS, Ekpenkhio E, Ebena B. Spatiotemporal trends and variability analysis of rainfall and temperature over Benin Metropolitan Region, Edo State, Nigeria. Geography, Envi- ronment, Sustainability 2023;16:6-15. Bestelmeyer BT, Miller JR, Wiens JA. Applying species diversity theory to land management. Ecological Applications 2003;13: 1750-61. Bourquin SL. The population ecology of the Nile crocodile (Croc- odylus niloticus) in the panhandle region of the Okavango Delta, Botswana. Stellenbosch University; Stellenbosch, South Africa; 2008. Brackhane S, Webb G, Xavier FME, et al. When conservation becomes dangerous: human-crocodile conflict in Timor- Leste. The Journal of Wildlife Management 2018;82: 1332-44. Brackhane S, Webb G, Xavier FME, et al. Crocodile management in Timor-Leste: drawing upon traditional ecological knowl- edge and cultural beliefs. Human Dimensions of Wildlife 2019;24:314-31. Britton A. Crocodilians: Natural History & Conservation. 2017. Available from: https://crocodilian.com/cnhc/cbd-con-1.htm Burgos-Ayala A, Jiménez-Aceituno A, Torres-Torres AM, et al. Indigenous and local knowledge in environmental manage- ment for human-nature connectedness: a leverage points perspective. Ecosystems and People 2020;16:290-303. Caldwell J. World trade in crocodilian skins 2010-2012. United Nations Environment Programme- World Conservation Monitoring Centre UNEP-WCMC, Cambridge, UK, 2014. Cao R, Somaweera R, Brittain K, et al. Genetic structure and di- versity of Australian freshwater crocodiles (Crocodylus john- stoni) from the Kimberley, Western Australia. Conservation Genetics 2020;21:421-9. Coe M, Webb GJW, Manolis SC, Whitehead PJ. Wildlife man- agement: crocodiles and alligators. Journal of Applied Ecol- ogy 1988;25. Cowlishaw G, Mendelson S, Rowcliffe MJ. Structure and opera- tion of a bushmeat commodity chain in southwestern Ghana. Conservation Biology 2005;19:139-49. Eaton MJ. Ecology, conservation and management of the Central African dwarf crocodile (Osteolaemus tetraspis), a progress report. In: Proceedings of the 18th working meeting of the IUCNSSC Crocodile Specialist Group. 2006. Eaton MJ, Barr B. LacTele, Republic of Congo. Crocodile Spe- cialist Group Newsletter 2005;24:18-20. Eaton MJ, Martin A, Thorbjarnarson J, Amato G. Species-level diversification of African dwarf crocodiles (Genus Osteolae- mus): a geographic and phylogenetic perspective. Molecular phylogenetics and evolution 2009;50:496-506. Echeverria C, Coomes DA, Hall M, Newton AC. Spatially explicit models to analyze forest loss and fragmentation between 1976 and 2020 in southern Chile. Ecological Modelling 2008;212: 439-49. Echeverría C, Coomes D, Salas J, et al. Rapid deforestation and fragmentation of Chilean temperate forests. Biological Con- servation 2006;130:481-94. Eniang EA, Godfrey C, Akani DD, et al. People’s perception of crocodiles in Nigeria. Herpetological Journal 2020;30: 112-6. Evans LJ. Assessing the impacts of habitat fragmentation and sub- sequent anthropogenic expansion on the behavioural, nesting and population ecology of the estuarine crocodile, Crocodylus porosus. 2016. Available from: https://orca.cardiff.ac.uk/id/ eprint/90075/ Eversole CB, Henke SE, Wester DB, et al. Responses of American alligators (Alligator mississippiensis) to environmental con- ditions: implications for population and ecosystem monitor- ing. Herpetologica 2015;71:37-45. Ewers RM, Didham RK. Confounding factors in the detection of species responses to habitat fragmentation. Biological Re- views 2006;81:117-42. Fabolude G, Aighewi IT. Evaluation of the extent of land use-land cover changes of benin city, edo state, nigeria from 1987- 2019. Journal of Applied Sciences and Environmental Man- agement 2022;26:1443-50. Fahrig L. Effects of habitat fragmentation on biodiversity. Annual Review of Ecology, Evolution, and Systematics 2003;34:487- 515. Ficetola GF, Bonardi A, Sindaco R, Padoa-Schioppa E. Estimating patterns of reptile biodiversity in remote regions. Journal of Biogeography 2013;40:1202-11. Fletcher Jr RJ, Didham RK, Banks-Leite C, et al. Is habitat frag- mentation good for biodiversity? Biological Conservation 2018;226:9-15. Floyd AC, Oikpor R, Ekene B. An assessment of climate change in Benin City, Edo State, Nigeria. FUTY Journal of the Envi- ronment 2016;10:87-94. Gabbatiss J. Weird Orange Crocodile found gorging on bats in Gabon’s caves. 2016. Available from: https://www.newscien- tist.com/article/2107837-weird-orange-crocodiles-found- gorging-on-bats-in-gabons-caves/ Heithaus MR, Wirsing AJ, Thomson JA, Burkholder DA. A re- view of lethal and non-lethal effects of predators on adult ma- rine turtles. Journal of Experimental Marine Biology and Ecology 2008;356:43-51. Ikponmwosa F. Colonialism and industrial development in Benin Province, Nigeria. Romanian Journal of Historical Studies 2020;3:20-9. Kafash A, Ashrafi S, Yousefi M, et al. Reptile species richness as- sociated to ecological and historical variables in Iran. Scien- tific Reports 2020;10:18167. I.O. Ogorode et al.52 Kpéra GN. Note on crocodiles in Bénin. Crocodiles Specialist Group/IUCN SSC Newsletter 2003;22:3-14. Kpéra GN, Aarts N, Tossou RC, et al. “A pond with crocodiles never dries up”: a frame analysis of human–crocodile rela- tionships in agro-pastoral dams in Northern Benin. Interna- tional Journal of Agricultural Sustainability 2014;12. Kpéra GN, Mensah GA, Sinsin B. Utilisation desproduits et sous- produits de crocodiles en médecine traditionnelle au Bénin. Bulletin de la Recherche Agricole du Bénin 2004;44:1-12. Luiselli L, Dendi D, Pacini N, et al. Interviews on the status of West African forest tortoises (genus Kinixys), including pre- liminary data on the effect of snail gatherers on their trade. Herpetological Journal 2018;28:171-7. Luiselli L, Petrozzi F, Akani GC, et al. Rehashing bushmeat –in- terview campaigns reveal some controversial issues about the bushmeat trade dynamics in Nigeria. Revue d’Ecologie (Terre Vie) 2017;72:3-18. Luiselli L, Politano E, Akani GC. Crocodile distribution in SE Nigeria. Part II. Crocodile Specialist Group Newsletter 1999;19:4-6. Malhi Y, Doughty CE, Galetti M, et al. Megafauna and ecosystem function from the Pleistocene to the Anthropocene. Proceed- ings of the National Academy of Sciences 2016;113:838-46. McNamara J, Rowcliffe M, Cowlishaw G, et al. Characterising wildlife trade market supply-demand dynamics. PloS One 2016;11:e0162972. Moshoeu TTJ. Overview of the trade of reptile taxa consumed for therapeutic purposes across Africa. 2017. Available from: https://wiredspace.wits.ac.za/items/26ad77c7-301d-4905- bfb8-c31e616ed414 Numbere AO, Chimieze BWI. Burrow distribution, size and chemical composition of the African dwarf crocodile (Osteo- laemus tetraspis) of the Niger Delta Nigeria. London Journal of Research in Science: Natural and Formal 2020;20. Odiana S, Idahosa-Ohio ME. Variations in rainfall in Benin City Nigeria and its implication in flood occurrences. Ethiopian Journal of Environmental Studies & Management 2023;16. Ogoanah OS, Omijife F. Animal species used for traditional med- icine in Benin City, Nigeria. Society for Experimental Biology of Nigeria 2017;17. Ogorode IO, Vincent-Akpu IF, Babatunde BB, et al. Illegal wildlife trade and Osteolaemus tetraspis. A case study of ac- tivities in bushmeat markets (Uwa and Edumanom) in Niger Delta. Sankalp Journal of Multidisciplinary Studies (SJMS) 2023;3. Ojo PE. A comprehensive evaluation of urbanization impact on land cover in Benin City. Journal of Energy Technology and Environment 2024;6:235-43. Pauwels B, Sanchez B. Diet records for the dwarf crocodile, Os- teolaemus tetraspis in Rabi oil fields and Loango National Park, southwestern Gabon. Hamadryad 2007;31:258-64. Pavoine S, Bonsall MB. Measuring biodiversity to explain com- munity assembly: a unified approach. Biological Reviews 2011;86:792-812. Petrozzi F, Amori G, Franco D, et al. Ecology of the bushmeat trade in West and Central Africa. Tropical Ecology 2016;57:545-57. Pimiento C, Leprieur F, Silvestro D, et al. Functional diversity of marine megafauna in the Anthropocene. Science Advances 2020;6:7650. Pincheira-Donoso D, Bauer AM, Meiri S, Uetz P. Global taxo- nomic diversity of living reptiles. PloS One 2013;8:e59741. Powell CB. Sites and species of conservation interest in the central axis of the Niger Delta (Yenagoa, Sagbama, Ekeremor, and Southern Ijaw Local Government Areas). A report of recom- mendations to the Natural Resources Conservation Council (NARESCON). NARESCON; Abuja, Nigeria; 1993. Powell CB. Final report submitted to the Environmental Affairs Department, Shell Petroleum Development Company (SPDC) of Nigeria, Ltd., Eastern Division. SPDC of Nigeria; Port Harcourt, Nigeria; 1995. Pyron M. Characterizing communities. Nature Education Knowl- edge 2010;3:39. Raghubanshi AS, Tripathi A. Effect of disturbance, habitat frag- mentation and alien invasive plants on floral diversity in dry tropical forests of Vindhyan highland: a review. Tropical Ecol- ogy 2009;50:57. Rice PM. Maya crocodilians: Intersections of myth and the natural world at early Nixtun-Ch’ich’, Petén, Guatemala. Journal of Archaeological Method and Theory 2018;25:705-38. Riley J, Huchzermeyer FW. African dwarf crocodiles in the Lik- ouala swamp forests of the Congo Basin: habitat, density, and nesting, Copeia 1999;2:313-20. Ripple WJ, Estes JA, Beschta RL, et al. Status and ecological ef- fects of the world’s largest carnivores. Science 2014;343: 1241484 Ritchie EG, Elmhagin B, Glen AS, et al. Ecosystem restoration with teeth: what role for predators? Trends in Ecology & Evo- lution 2012;27:265-71. San Diego Zoo Wildlife Alliance. Dwarf Crocodile (Osteolaemus tetraspis) Fact Sheet. 2015. Available from: http://ielc. libguides. com/ sdzg/factsheets/ dwarf_crocodile Shirley MH. Crocodile conservation in West Africa: planning for the future. Final report to the Wildlife Division of the Forestry Commission, Ghana and the Wildlife Directorate, Ministry of Water and Forests, Cote d’Ivoire. 2007. Shirley M, Burtner B, Oslisly R, et al. Diet and body condition of cave-dwelling dwarf crocodiles (Osteolaemus tetraspis, Cope 1861) in Gabon. African Journal of Ecology 2016;55:411-22. Shirley MH, Oduro WE, Yaokokore Beibro H. Conservation and status of crocodiles in Ghana and Cote d’Ivoire, West Africa. Oryx 2009;43:136-45. Shirley MH, Villanova VL, Vliet KA, Austin JD. Genetic barcod- ing facilitates captive and wild management of three cryptic African crocodile species complexes. Animal Conservation 2015;18:322-30. Smolensky NL. Co-occurring cryptic species pose challenges for conservation: a case study of the African dwarf crocodile (Os- teolaemus spp.) in Cameroon. Oryx 2015;49:584-90. Somaweera R, Brien ML, Platt SG, et al. Direct and indirect in- teractions with vegetation shape crocodylian ecology at mul- tiple scales. Freshwater Biology 2019;64:257-68. Somaweera R, Brien M, Shine R. The role of predation in shaping crocodilian natural history. Herpetological Monographs 2013;27:23-51. Somaweera R, Nifong J, Rosenblatt A, et al. The ecological im- portance of crocodylians: towards evidence-based justifica- tion for their conservation. Biological Reviews 2020;95: 936-59. Sprawls S, Ashe J, Howell K, Drewes R. A field guide to the rep- Dwarf crocodile diversity In Benin Metropolis Markets 53 tiles of East Africa: Kenya, Tanzania, Uganda, Rwanda and Burundi. Academic Press; Cambridge, USA; 2002. Thorbjarnarson J. Crocodile tears and skins: international trade, economic constraints, and limits to the sustainable use of croc- odilians. Conservation Biology 1999;13:465-70. Thorbjarnarson JB, Eaton MJ. Preliminary examination of croc- odile bushmeat issues in the Republic of Congo and Gabon. In: Proceedings of the 17th working meeting of the IUCNSSC Crocodile Specialist Group. 2004. Trefon T. Bushmeat: culture, economy and conservation in Cen- tral Africa. Oxford University Press; Oxford, UK; 2023. Van Asch B, Versfeld WF, Hull KL, et al. Phylogeography, genetic diversity, and population structure of Nile crocodile popula- tions at the fringes of the southern African distribution. PLoS One 2019;14:e0226505. Van der Ploeg J, Cauillan-Cureg M, van Weerd M, Persoon G. Why must we protect crocodiles?’ Explaining the value of the Philippine crocodile to rural communities. Journal of Integra- tive Environmental Sciences 2011;8:287-98. Walelign SZ, Nielsen RM, Jakobsen BJ. Price elasticity of bush- meat demand in the greater Serengeti ecosystem: insights for managing the bushmeat trade. Frontiers in Ecology and Evo- lution 2019;7:162. Walmsley CW, Smits PD, Quayle MR, et al. Why the long face? The mechanics of mandibular symphysis proportions in croc- odiles. PLOS One 2013;8:e53873.