Ecological and conservation considerations on the reptile fauna of ...

HERPETOZOA 11 (3/4): 141-153Wien, 26. Februar 1999

Ecological and conservation considerationson the reptile fauna of the eastern Niger Delta (Nigeria)

Betrachtungen zu Ökologie und Schutz der Reptilienfauna des östlichen Niger Deltas (Nigeria)

GODFREY C. AKANI & LUCA LUISELLI & EDOARDO POLITANO *]

KURZFASSUNG

Qualitative und quantitative Daten zur Reptilienfauna des östlichen Niger-Deltas (Port Harcourt, Rivers State)werden präsentiert. 60 Reptilienarten wurden im Untersuchungsgebiet festgestellt: 3 Krokodile, 7 Schildkröten, 13Echsen und 37 Schlangen. Die quantitativen Angaben zur Diversität der Reptilien beruhen auf Fang-Wiederfang-Daten entlang von zehn Transekten, die jeweils verschiedene Spektren von Biotoptypen umfaßten und sich hinsichtlichder Diversität, Dominanz und Abundanz der vorkommenden Reptilienarten signifikant unterschieden. Grundsätzlichbeherbergten trockene und sumpfige primäre und sekundäre Regenwälder eine weit größere Artenzahl als die Man-groven. Echsen waren die häufigsten Reptilien. Dabei waren Individuen der Gattung Mabuya zahlenmäßig vor-herrschend, außer auf landwirtschaftlichen Flächen und im Siedlungsbereich, wo Agama agama die dominierendeEchse war. An den meisten Standorten war die Vielzahl der Schlangenarten bemerkenswert, doch nur sechs (Gastro-pyxis smaragdina, Psammophis phillipsi, Naja nigricollis, Python regiüs in terrestrischen Biotopen und Afronatrixanoscopus, Grayia smythii in aquatischen Biotopen) waren regelmäßig anzutreffen.

Die Anzahlen der je Transekt festgestellten Biotoptypen und Reptilienarten waren positiv korreliert (p < 0.05),

der je Transekt festgestellten Reptilienarten waren nicht signifikant korreliert. Dieser Befund ist nicht einleuchtend.Man würde erwarten, daß mit steigender Anzahl verfügbarer Biotoptypen die Dominanz einer einzelnen Art abnehmensollte. Bei Ausschluß von Mabuya aus dieser Berechnung war die zu erwartende negative Beziehung zwischen denVariablen gegeben (p < 0.05).

Einige negative Auswirkungen der industriellen Förder- und Transportaktivitäten fur Gas und Öl auf die Di-versität und Häufigkeit von Reptilien im Niger-Delta werden genannt. Crocodylus niloticus und C. cataphractusscheinen außerordentlich selten zu sein. Um eine Verarmung der Reptiliendiversität zu vermeiden, sollten die Förder-industrien ihre Öl- und Gasleitunsanlagen nicht mehr in primären oder sekundären Waldflächen verlegen, da diese diearten- und individuenreichsten Reptiliengemeinschaften beherbergen.

ABSTRACT

Qualitative and quantitative data on the reptile fauna of the eastern Niger Delta (Port Harcourt, Rivers State) ispresented. 60 reptile species were found in the study areas, 3 crocodiles, 7 turtles, 13 lizards, and 37 snakes. Quantifi-cation of reptilian diversity was based on capture-mark-recapture procedures along ten transects which covered differ-ent spectra of habitat types each and varied significantly in terms of reptile species diversity, dominance, and abun-dance. As a rule, both dry and swampy primary and secondary rain forests housed a significant excess of species ascompared with the mangroves. Lizards were the most abundant reptiles. Species of the genus Mabuya were numeri-cally dominant except in cultivated lands and suburbia where Agama agama was the dominant lizard. In most locali-ties there was a remarkable variety of snake species, but only six (Gastropyxis smaragdina, Psammophis phillipsi,Naja nigricollis, Python regius in terrestrial habitats and Afronatrix anoscopus. Grayia smythii in aquatic habitats)were regularly found.

The number of habitat types and the number of reptile species registered per transect were positively correlated(p < 0.05), but this general pattern was mainly due to the snakes. Dominance index of reptile species and number ofhabitat types registered per transect were not significantly correlated. This is a counter-intuitive result as one wouldexpect: the higher the number of macrohabitats available, the lower the dominance of a single species. When Mabuyawas excluded from this analysis, the expected negative relationship between these variables was obtained (p < 0.05).

Some negative effects of industrial gas and oil transmission and extraction activities on the biodiversity andabundance of reptiles in the Niger Delta area are indicated. Crocodylus niloticus and C. cataphractus appear to beextremely rare. To avoid impoverishment of reptilian diversity, industry companies should no longer lay their oil andgas transmission installations through primary and secondary rain forest patches, which house both the highest diver-sity of taxa and the highest densities of reptile specimens.

KEY WORDS

Niger Delta, Nigeria; reptile communities, species diversity, dominance, abundance, ecology, habitat, impact ofgas and oil industries on reptile populations, threat, conservation

Authors in alphabetical order

©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

142 GODFREY e. AKANI & LUCA LUISELU & EDOARDO POLITANO

INTRODUCTION

Nigerian oil and gas industry activi-ties concentrate mainly on the Niger Deltawhich, concurrently, represents one of theworldwide largest wetlands (more than20,000 km2) and harbours a remarkablenumber of reptile taxa in a variety of ecol-ogically different habitats.

In spite of that, the herpetofauna ofthe Niger Delta is poorly known althoughAfrican rain forests have frequently beensubject to careful research during the recentyears. However, data on reptile communi-ties are scarce, and quantitative studies onthe functioning of these communities re-mained at an embryonic stage (SCOTT1982; but see LUISELLI & al. 1998). Somecollective information on Amphibians andReptiles of the Niger Delta is presented inclassical monographs of West African her-petofaunas or in specific papers on Nige-rian species (e.g., ROMER 1953; GRANDI-SON 1956, 1968; BLACKWELL 1967; SCHI0TZ1967, 1969; ARNOULT & LAMOTTE 1968;EKUNDAYO & OTUSANYA 1969; DÜNGER1967a, 1967b, 1967c, 1968, 1971a, 1971b,1972a, 1972b, 1973; ViLLiERS 1975; RO-MAN 1976; VANEUSDEN 1978; JOGER 1979,1981; HUGHES 1983; LOUMONT 1984;GARTSHORE 1985; BUTLER & REID 1986;REID 1986; SODEINDE & KUKU 1989;SODEINDE & OGUNJOBI 1994). In all thisliterature, however, very little is reportedon ecological distribution and eco-ethology

of the various taxa. Moreover, in a recentsurvey in which an environmental devel-opment strategy for the Niger Delta is de-fined (SINGH & al. 1995), only five repre-sentatives of the reptiles (three crocodiliansand two pythons) are listed.

The present research constitutes thefirst attempt to describe the composition ofthe reptile fauna of several Niger Delta lo-calities both in terms of quality and quan-tity and in relation to the variation ofhabitats available. Considerations on thethreats to the reptile fauna and immediatemanagement needs are presented in thelight of potential adverse effects of the fastdeveloping gas and oil transmission andextraction activities.

Aim of this paper is neither to presentexhaustive faunistic records nor to reviewthe reptilian zoogeography of the region,but just to give a commented summary ofover two years of intensive field explora-tions in a country which, mainly because ofinternal problems limiting the foreigners'access (e.g. political misunderstandingbetween the Federal Government of Nige-ria and the western countries, and presenceof organized criminality), is still largelyunknown from the zoological point ofview. The quantitative data in this paperhave already been presented in an unpub-lished report by the authors to the company"T.S.K.J. Nigeria Ltd.".

STUDY AREA AND METHODS

Study area

The quantitative analyses in this pa-per are based on field data collected in theeastern Niger Delta (Port Harcourt region,Rivers State, southeastern Nigeria) duringthe wet seasons of 1996 through 1997, andthe dry season of 1997 (fig. 1). Additionalqualitative data (e.g. presence/absence oftaxa) were, however, collected also duringthe wet and the dry season of 1998.

The environment of the study regionis extremely heterogenous in that water andsoil patchily alternate. The variation of thesalinity level of waters (due to concomitantpresence of freshwater and tidal water) is

an important factor. Most parts of the areaare heavily populated, with hundreds oftowns and villages surrounding Port Har-court City.

Both dry and swamp forest patchesare interspersed in a mosaic of farmlandand cultivations (mainly cassava, oil palms,bananas and plantains). Dryland rainfor-ests can be assigned to one of three catego-ries: primary forest, secondar)' forest, andshrubland. Periodically and permanentlyflooded freshwater swamp forests may beprimary or secondary. Mangrove formations(dominated by Avi cenni a and Rhizophora)are present in an arboreal (high man-groves) and bushy (low mangroves) form.


Ecology and conservation of Nigerian reptiles 143

Cameroon

Fig. 1: The study area in the southeastern Niger Delta (Nigeria). P.H. - Port Harcourt City.Abb. 1: Das Untersuchungsgebiet im südöstlichen Niger Delta (Nigeria). P.H. - Port Harcourt City.

Methods

General survey procedures. Oursurvey is based on (i) original observationsin the field, (ii) examination of preservedspecimens in the collections of the Depart-ment of Biological Sciences of Port Har-court University of Science and Technol-ogy, and (iii) original interviews with se-lected persons from the main villages of thearea. With regard to point (i), most obser-vations were made in forested areas of thePort Harcourt region (see POLIT ANO 1997);others were done in villages, at the bordersof roads where local people were involvedin marketing animals (AKANI & al. 1998),and along the course of rivers (Bonny, NewCalabar, Sambreiro, Orashi). With regardto point (iii), we considered only interviewsin which the interviewed person was ableto doubtlessly describe the observed species(especially snakes), as well as a typical be-havioural display of that species (e.g. typi-cal defensive display of snakes, basking

postures of lizards, etc).Various observation techniques were

applied. Reptiles were searched for bywalking along random routes leadingthrough all habitat types available. Due tologistic problems, field work was predomi-nantly restricted to the daylight hours(08.00 to 18.00) which is of course not thebest time for finding Amphibians andReptiles in African rain forests, as severalof these species have their above-groundactivity peaks early in the morning and latein the afternoon (BRANCH 1988). Mostanimals encountered during our surveyswere captured by hand, drift fences or pit-fall traps and photographed.

To avoid unrecognized recaptures,lizards were marked by toe-clipping andsnakes by ventral scale clipping. Crocodileswere not captured and remained unmarked.Turtles were marked by unique sequencesof notches filed into the marginal scutes.

Quantitative survey procedures.Ten study sites were selected along the


144 GODFREY e. AKANI & LUCA LUISELU & EDOARDO POLITANO

Table 1: Transects studied in the region of Port Harcourt (Rivers State, Nigeria) including local names andgeographic coordinates of the study areas.

Tab. 1 : Die untersuchten Transekte im Gebiet Port Harcourt (Rivers State, Nigeria) sowie die Ortsnamen undgeographischen Koordinaten der Untersuchungsgebiete.

Transect

TIT2T3T4T5T6T7T8T9TIO

Name of Place / Ortsname

KreigeniAbarikpoOtariRumujiOrubiriSoku - Eiern SangamaTombia ForestTombia MangroveOrashi RiverPeterside (Bonny)

Latitude / Breite

N 05°1759.2N05°08'11.9N 04°53'22.3N04°5719.3N 04°42'25.6N 04°40'39.8N 04°46'34.9N 04°46'50.9N O4044.43 j

N 04°29'14.9

Longitude / Länge

E006°3741.3E 006°37*45.7E006°41'19.7E006°46'28.1E007°01'13.6E 006°40'54.2E 006°53'56.9E 006°51*53.9E006°38'10.1E007°10'04.8

"LNG Nigeria Gas Transmission SystemProject" pipe-line (AA.VV. 1997) in such away that the various natural habitats pres-ent along this pipe-line were equally repre-sented. Detailed methods describing theselection criteria of these ten study sites aregiven elsewhere (POLITANO 1997). In gen-eral, every possible effort was made toavoid unbalanced surveying in the varioushabitats.

Geographic coordinates and the nameof each study site are indicated in table 1,the main habitat types present in these lo-calités are given in table 2. In each studysite, a study transect was determined whichpassed through the various habitat typespresent in this location. The transects wereapproximately 7 km long each, and werewalked along only in one direction andonce per day to avoid multiple counts ofindividuals per observation unit. To avoidsampling differences between study sites,we walked along the transects always at thesame time of the day (from 09.00 to 13.00,Lagos time), maintaining constant velocity.Only those reptile specimens which wereclearly visible to us (i.e. not hidden undercovering objects, etc) were counted for thequantitative analyses. The specimens whichwere found hidden within the transects(this was the case especially in snakes)were recorded for describing the faunalcomposition of the site, but were not usedfor quantitative statistical inter site com-parisons. Exclusion of hidden specimenswas done under the assumption that theproportion of active (visible) specimensshould be the same in all study transects

(given that the time of surveying was thesame in all cases) and because the prob-ability to find covered animals might havebeen different in the various sites (e.g., be-cause of different frequencies of objects onthe ground, etc).

These records formed the basis forthe quantitative estimation of biodiversityin each study site. Mabuya specimens en-countered during these standardized sur-veys were not identified to species level.Records of reptile specimens from outsidethe transects were excluded from thequantitative biodiversity analysis, but wereused to describe the reptile fauna composi-tion of each study site (see "Reptile taxa ofthe study transects"). In general, at least 40hours were spent in the field in each studytransect.

We calculated both species diversityand species dominance of each area whichare important parameters for any biodi-versity analysis (DODD 1992).

Species diversity (£>) was calculatedusing MARGALEF'S Diversity Index (MA-GURRAN 1988):

DMg = (S-l)/\nN

where S represents the number of speciesand N the number of individuals (sampled)in the area under study.

Species dominance (d) was assessedusing the BERGER-PARKER Index (MAGUR-RAN 1988):

d = NmaxIN

where N represents the number of indi-



Table 2: Main habitat types present in study transects Tl to T10 of the Port Harcourt region (Rivers State, Ni-geria). NH - Number of habitat types; NI - Number of individuals; NT - Number of taxa; NTH - Number of transectsin which a given habitat type was found. Data under NI and NT exclusively represent the the results of the standard-ized sampling for the quantitative analyses.

Tab 2: Die wichtigsten Biotoptypen der Untersuchungstransekte Tl bis T10 im Port Harcourt Gebiet (RiversState, Nigeria). NH - Anzahl Biotoptypen; NI - Anzahl Individuen; NT - Anzahl Taxa; NTH - Anzahl Transekte, indenen ein gegebener Biotoptyp angetroffen wurde. Die Werte unter NI und NT sind ausschließlich Ergebnisse derstandardisierten Aufsammlungen für die quantitativen Analysen.

Habitat type / Biotoptyp

Primary dry Rainforest / PrimärwaldSecondary dry Rainforest / SekundärwaldShrubland / BuschlandPrimary Swamp-forest /Primärer SumpfwaldSecondary Swamp ForestSekundärer SumpfwaldHigh Mangrove / hohe MangroveLow Mangrove / niedrige MangroveFarmland and cultivations/landwirtschaftliche FlächenFreshwater body / Süßwasserkörper

NHNINT

Tl

-XX

--

Y

X

4266-7

T2

--X

--

Y

-

2113-4

T3

XX-

Y

--

Y

X

5485-6

T4

--X

--

Y

2364-5

T5

--X

-X

Y

3131-2

T6

-X-

xY

--

Y

Y

5264-5

T7

-XX

Y

-X

Y

5817-8

T8

---

X-

1435-6

T9

-XX

--

Y

X

4907-8

T10

XXX

Y

-X

Y

X

7527-8

NTH

267

1

4

13

9

5

NI

1911957

15

52

437

107

7

NT

7138

13

53

3

3

viduals and Nmax the number of individualsof the most abundant species (sampled) inthe area under study.

An increase in the value of \ld (the re-ciprocal form of the index) indicates an in-crease in diversity and a decrease in domi-nance (MAGURRAN 1988). In order to deter-mine whether inclusion of juveniles influ-enced the values of diversity and dominance,separate indices were calculated (i) with ju-veniles included and (ii) excluded. Since non-parametric matrix correlation Mantel tests didnot show evident differences between corre-sponding indices (i) and (ii) in any study tran-sect (in all cases: P > 0.4), the pertinent indi-ces were pooled. Recaptured individuals wereexcluded from calculations of diversity anddominance.

Several methodological factors mayhave biased the results: e. g., vegetation andlandscape characteristics permit easier local-

ization of reptiles in secondar}' forests than inprimary forests. Thus, the lower inunber oftaxa detected in the latter might reflect ratherfield constraints than true differences.

Notwithstanding that weak points maylimit the accuracy of our method, the presentinvestigation is one of the few quantitativeapproaches to study the reptile fauna of anAfrotropical rainforest region, and the firstone for Nigeria (comp. SCOTT 1982).

Statistical analyses

Statistical tests were done by means ofSAS® (Statistical Analysis System) and STA-TISTICA® PC packages. All tests were two-tailed, with alpha set at 5%. Data were testedfor normal distribution by Kosmogorov-Smirnov tests before applying parametric sta-tistical procedures, otherwise non-parametricstatistical tests were used.

RESULTS AND DISCUSSION

Reptile taxa of the region

Based on the methods applied, weregistered the presence of the following 60reptile taxa on the territory under study (3crocodilians, 7 chelonians, 13 saurians, 37

snakes). The list includes only taxa whichwere observed by the authors (i. e. inter-view results are not considered).

Crocodiles: Crocodylus niloticus suchusGEOFFROY, 1807, C. c. cataphractus CUVIER,1789, Osteolaemus t. tetraspis COPE, 1861.


146 GODFREY e. AKANI & LUCA LUISELLI & EDOARDO POUTANO

Turtles: Pelomedusa subrufa olivacea(SCHWEIGGER, 1812), Pelusios c. castaneus(SCHWEIGGER, 1812), P. niger DUMÉRIL&BlBRON, 1835), Trionyx triunguis (FORS-KAL, 1775), Kinixys belliana nogueyi (LA-TASTE, 1886), K. erosa (SCHWEIGGER, 1812),K. homeana BELL, 1827.

Saurians: Varanus niloticus ornatus(DAUDIN, 1803), Chamaeleo g. gracilis HAL-LOWELL, 1842, Ch. owenii GRAY, 1831,Agama a. agama LINNAEUS, 1758, Hemi-dactylus brooki angulatus HALLOWELL,1854, H. fasciatus fasciatus GRAY, 1845,H. mabouia (MOREAU DE JONNÈS, 1818),Lygodactylus conraui conraui TORNIER,1902, L. fischeri BOULENGER, 1890, Ma-buy a affinis (GRAY, 1839), M. maculilabrismaculilabris (GRAY, 1845), Mochlus (Rio-pa) femandii (BURTON, 1836), Panaspistogoensis WERNER, 1902.

Snakes: Calabaha reinhardti (SCHLE-GEL, 1848), Python regius (SHAW, 1802),P. sebae (GMELIN, 1788), Dispholidus t.typus (SMITH, 1829), Thelotornis kirtlandii(HALLOWELL, 1844), Rhamnophis a. aethi-opissa GÜNTHER, 1862, Grayia smythii(LEACH, 1818), Philothamnus h: heteroder-mus (HALLOWELL, 1857), Natriciteres fuli-ginoides (GÜNTHER, 1858), N. v. variegata(PETERS, 1861), Afronatrix anoscopus(COPE, 1861), Mehelya crossi (BOULENGER,1895), M. guirali (MOCQUARD, 1887), Mpoensis (SMITH, 1847), Gastropyxis sma-ragdina (SCHLEGEL, 1837), Hapsidophryslineatus FISCHER, 1856, Aparallactus m.modestus (GÜNTHER, 1859), Lamprophisvirgatus (HALLOWELL, 1854), Bothrophthal-mus I. lineatus (PETERS, 1863), Meizodoncoronatus (SCHLEGEL, 1837), Toxicodryasblandingi (HALLOWELL, 1844), T. pulveru-lenta (FISCHER, 1856), Dipsadoboa du-chesnii (BOULENGER, 1901), Crotaphopeltishotamboeia (LAURENTI, 1768), Psammophisphillipsi (HALLOWELL, 1844), Polemoncollar is brevi or (WITTE & LAURENT, 1947),Dasypeltis fasciata SMITH, 1849, Den-droaspis j . jamesoni (TRAILL, 1843), Najamelanoleuca HALLOWELL, 1857, N n. nigri-collis REINHARDT, 1843, Pseudohaje goldii(BOULENGER, 1895), Atheris squamiger(HALLOWELL, 1854), Causus maculatus(HALLOWELL, 1842), Bitis nasicorni s (SHAW,1802), B. g. gabonica (DUMÉRIL & BlBRON,1845), Atractaspis aterrima GÜNTHER, 1863,

A. corpulenta leucura MOCQUARD, 1885.

Various other taxa, the presence ofwhich in the study area is either verified byother sources or is strongly to be suspected,were not found during our survey: e.g.Melanoseps occidentalis (PETERS, 1877)which was captured by ROMER (1953) inthe Port Harcourt area; Chamaeleo crista-tus STUTCHBURY, 1837 which was capturedby us in the surroundings of Eket (Akwa-Ibom State, approximately 100 km east ofPort Harcourt) and by REID (1986) in thesurroundings of Calabar (Cross River State,about 170 km east of Port Harcourt) and,thus, could potentially be present in thestudied region. With regard to snakes,BUTLER & REID (1986) cited Rhinotyphlopscrossii (BOULENGER, 1893) and fyphlopspunctatus (LEACH, 1819) as being presentin the Calabar area of south-eastern Nige-ria. These taxa might thus be found in thePort Harcourt region. However, T. puncta-tus is a savannah species, unlikely to occurin Calabar, where the forest species T. con-gestus (DUMÉRIL & BIBRON, 1844) is ratherto be expected (B. HUGHES, personal com-munication). Specimens of T. congestus,captured in the Niger Delta, are in factstored in the collection of the British Mu-seum (Natural History), London (B.HUGHES, pers. comm.). Other snake speciesmight have "escaped" our attention: Atract-aspis i. irregularis (REINHARDT, 1843),Pseudohaje nigra GÜNTHER, 1858, Thra-sops flavigularis (HALLOWELL, 1852), T.occidentalis PARKER, 1940, and Dasypeltisscabra (LINNAEUS, 1758). Dendroaspisviridis (HALLOWELL, 1844) was reported tooccur in the Niger Delta (VILLIERS, 1975);the same is true for the savannah form ofthe Nile Monitor Varanus n. niloticus (LIN-NAEUS, 1766) (BÖHME & ZIEGLER 1977)but these taxa are unlikely to be present inthe Niger Delta.

Species diversity of the reptile faunaof Rivers State is remarkably high sinceboth Afrotropical widespread taxa (e.g.Pelomedusa subrufa, Naja melanoleuca, N.nigricollis, Dispholidus typus) and taxawhich are typically confined to the rainforests of West Africa (e.g. Calabaria rein-hardti) are present.

Most of the above-cited reptile taxaare widespread and locally abundant in theeastern Niger Delta region. However, thelarge crocodiles {Crocodylus cataphractus



and C. niloticus) appear to be rare and veryendangered (AKANI & al. 1999), whileother taxa, which are regularly found,might soon suffer rapid population declinebecause of habitat loss and human huntingactivities (Kinixys spp., Python sebae).

Reptile taxa of the study transects

The habitat types found in the variousstudy transects are roughly presented intable 2 (for a more detailed description, seePOLITANO 1997).

Transect 1 (TI) - Agama agama,Mabuya affinis, M. maculilabhs, Mochlusfernandii, Python sebae, P. regius, Dis-pholidus typus, Psammophis phillipsi,Gastropyxis smaragdina, Grayia smythii,Crotaphopeltis hotamboeia, Dendroaspisjamesoni, Naja nigricollis.

Transect 2 (T2) - Agama agama,Mabuya affin is, Mochlus fernandii, Pythonregius, Psammophis phillipsi, Crotapho-peltis hotamboeia, Naja nigricollis.

Transect 3 (T3) - Osteolaemus tetras-pis, Pelusios castaneus, P. niger, Pelomedusasubrufa, Kinixys belliana, K. homeana,Agama agama, Mabuya affinis, M. maculi-labris, Mochlus fernandii, Panaspis togoen-sis, Hemidactylus fasciatus, H. mabouia, Va-ranus niloticus, Chamaeleo owenii, Ch. gra-cilis gracilis, Calabaria reinhardti, Pythonsebae, P. regius, Dispholidus typus, Toxico-dryas blandingi, T. pulverulenta, Thelotorniskirtlandii, Grayia smythii, Afronatrix ano-scopus, Natriciteres fuliginoides, N. varie-gata, Dasypeltis fasciata, Crotaphopeltis ho-tamboeia, Psammophis phillipsi, Gastropyxissmaragdina, Philothamnus heterodermus,Pseudohaje goldii, Dendroaspis jamesoni,Naja melanoleuca, Naja nigricollis, Athenssquamiger, Bitis nasicornis, B. gabonica.

Transect 4 (T4) - Pelusios castaneus,Pelomedusa subrufa, Trionyx triunguis,Kinixys homeana, Chamaeleo owenii, Aga-ma agama, Mabuya affinis, M. maculi lab-ris, Mochlus fernandii, Hemidactylus fas-ciatus, Varanus niloticus, Calabaria rein-hardti, Python regius, P. sebae, Dispholidustypus, Toxicodryas blandingi, T. pulveru-lenta, Grayia smythii, Afronatrix anoscopus, Natriciteres fuliginoides, N. variegata,Psammophis phillipsi, Gastropyxis smarag-dina, Crotaphopeltis hotamboeia, Pseudo-haje goldii, Dendroaspis jamesoni, Naja nig-ricollis, N. melanoleuca, Bitis gabonica.

Table 3: Reptile species including numbers ofspecimens (n) and their proportional shares (%n) asfound in the quantitative biodiversity analysis in tran-sects Tl to T10.

Tab. 3: Die in der quantitativen Biodiversitât-sanalyse in den Transekten TI bis TIO festgestelltenReptilienarten, ihre Individuenzahlen (n) und derenProzentanteile (%n).

Tl

T2

T3

T4

T5T6

T7

T8

T9

T10

Species / ArtAgama agamaMabuya sp.Mochlus fernandiiDendroaspis jamesoniDispholidus typusGrayia smythii

Sum/Summe TlMabuya sp.Mochlus fernandiiAgama agama

Sum / Summe T2Mabuya sp.Mochlus fernandiiAgama agamaDendroaspis jamesoniNaja melanoleuca

Sum / Summe T3Mabuya sp.Mochlus fernandiiAgama agamaGastropyxis smaragdina

Sum / Summe T4Mabuya sp.Mabuya sp.Dendroaspis jamesoniNatriciteres variegataNaja nigricollis

Sum / Summe T6Agama agamaMabuya sp.Panaspis togoensisVaranus niloticusThelotornis kirtlandiiDendroaspis jamesoniNatriciteres variegata

Sum / Summe T7Mabuya sp.Agama agamaVaranus niloticusGrayia smythiiNaja nigricollis

Sum / Summe T8Mabuya sp.Agama agamaMochlus fernandiiPsammophis phillipsiGastropyxis smaragdinaNaja nigricollisPython regius

Sum / Summe T9Mabuya sp.Agama agamaVaranus niloticusGastropyxis smaragdinaNaja melanoleucaNaja nigricollisBitis gabonica

Sum/Summe T10

n2182211

2663211321

1212

4824183

361321311

26383231322

812117122

43473142321

9040423111

52

%n7.69

69.237.697.693.843.84100

54.5427.2718.18100

66.672.08

25.002.084.16100

66.662.78

22.228.33100100

80.7611.543.853.85100

46.9139.513.701.233.702.462.46100

48.8339.532.324.654.65100

52.2234.444.442.223.332.221.11100

76.927.693.855.771.921.921.92100


148 GODFREY e. AKANI & LUCA LUISELLI & EDOARDO POLITANO

Transect 5 (T5) - Crocodylus niloti-cus, Kinixys homeana, Agama agama,Mabuya affinis, Varanus niloticus, Pythonsebae, Grayia smythii, Natriciteres fuligi-noides, Dispholidus typus, Thelotornis kirt-landii, Gastropyxis smaragdina, Psamm-ophis phillipsi, Naja melanoleuca, N. nig-ricollis, Bitis gabonica.

Transect 6 (T6) - Crocodylus niloti-cus, Osteolaemus tetraspis, Pelusios casta-neus, P. niger, Kinixys erosa, K. homeana,Mabuya affinis, M. maculilabris, Mochlusfernandii, Panaspis togoensis, Hemidacty-lus fasciatus, H. mabouia, Varanus niloti-cus, Chamaeleo owenii, Ch. gracilis, Cala-baria reinhardti, Python regius, P. sebae,Toxicodryas blandingi, Aparallactus mo-destus, Grayia smythii, Afronatrix anosco-pus, Natriciteres fuliginoides, N. variegata,Bothrophthalmus lineatus, Crotaphopeltishotamboeia, Gastropyxis smaragdina, Phi-lothamnus heterodermus, Rhamnophisaethiopissa, Psammophis phillipsi, Den-droaspis jamesoni, Pseudohaje goldii,Naja melanoleuca, N. nigricollis, Athenssquamiger, Bitis gabonica, B. nasicornis,Atractaspis aterrima.

Transect 7 (T7) - Agama agama, Ly-godactylus conraui, Mabuya affinis, M.maculilabris, Varanus niloticus, Pythonregius, P. sebae, Grayia smythii, Dispholi-dus typus, Psammophis phillipsi, Gastro-pyxis smaragdina, Crotaphopeltis hotam-boeia, Dendroaspis jamesoni, Naja nigri-collis, Bitis gabonica, Causus maculatus.

Transect 8 (T8) - Agama agama, Ma-buya affinis, Varanus niloticus, Python re-gius, P. sebae, Thelotornis kirtlandii, Toxi-codryas blandingi, T. pulverulenta, Afrona-trix anoscopus, Gastropyxis smaragdina,Psammophis phillipsi, Dendroaspis jame-soni, Naja nigricollis.

Transect 9 (T9) - Crocodylus cata-phractus, C. niloticus, Osteolaemus tetr-aspis, Pelomedusa subrufa, Pelusios casta-neus, P. niger, Kinixys belliana, K. erosa,K. homeana, Agama agama, Mabuya af-finis, M. maculilabris, Mochlus fernandii,Hemidactylus fasciatus, H. mabouia, Vara-nus niloticus, Chamaeleo owenii, Cala-baria reinhardti, Python sebae, P. regius,Thelotornis kirtlandii, Toxicodryas blan-dingi, Grayia smythii, Afronatrix anosco-pus, Natriciteres fuliginoides, N. variegata,Bothrophthalmus lineatus, Crotaphopeltis

hotamboeia, Aparallactus modestus, Dasy-peltis fasciata, Gastropyxis smaragdina,Psammophis phillipsi, Dendroaspis jame-soni, Pseudohaje goldii, Naja nigricollis,N. melanoleuca, Bitis gabonica, B. nasi-cornis.

Transect 10 (T10) - Pelusios casta-neus, P. niger, Agama agama, Mabuya af-finis, Mochlus fernandii, Varanus niloticus,Python regius, P. sebae, Dispholidus typus,Toxicodryas blandingi, Grayia smythii, Af-ronatrix anoscopus, Crotaphopeltis hotam-boeia, Gastropyxis smaragdina, Philo-thamnus heterodermus, Psammophis phil-lipsi, Naja melanoleuca, N. nigricollis.

Diversity of reptilian speciesand diversity of available habitats

High numbers of reptile taxa werefound in transects T3, T4, T6, and T9whereas the other transects housed muchsmall numbers of reptile taxa. These dataindicate that high taxonomic diversity wasassociated with the presence of rainforestpatches, whereas low taxonomic diversitywas associated with cultivated lands andareas subject to intensive human activity.

There was a positive correlation be-tween the number of environmental typesper study transect and the correspondingnumber of species found (r = 0.331, n = 10,P < 0.05; fig. 2). This general pattern wascaused by the snakes as the lizard speciesdiversity proved to be rather independenton the habitat heterogeneity of each studytransect (snakes versus lizards: ANCOVAto both slopes and ordinate intercepts, P <0.01). The numbers of crocodile and che-lonian taxa were too small to permit anystatistical comparison of this type.

Quantitative biodiversity parameters

Table 3 presents the results of thequantitative surveys along the study tran-sects Tl to T10; in table 4 their biodiver-sity indexes are entered.

During our quantitative surveys, werecorded a total of 426 reptile individuals.In general, and despite the remarkable dif-ferences observed between the various tran-sects, lizards represented the majority ofthe reptile individuals. Among lizards, thescincids of the genus Mabuya were by far



Table 4: Values of MARGALEF's diversity index(D), and BERGER-PARKER'S species dominance index(d) for the study transects Tl to T10.

Tab. 4: Die Werte von MARGALEFs Diversi-tâtsindex (D) und BERGER-PARKERS Dominanzindex (d)in den Untersuchungstransekten TI bis T10.

Transect

TIT2T3T4T5T6T7IT8T9TIO

D

1.5380.8340.4650.8370.0000.9202.7331.0634.0501.800

d

0.6920.5450.6670.6671.0000.8070.4690.4880.5220.769

the most abundant ones, however, Agamawas dominating in some transects, espe-cially in the vicinity of small villages androads.

When we plotted the dominance in-dexes of reptile species in transects Tl toT10 (d in table 4) against the number ofhabitat types available in each transect,there was no significant correlation be-tween these variables (r = 0.253, n = 10, P> 0.2; fig. 3a). The absence of a signifi-cantly negative correlation between the twovariables is surprising, as one would ex-pect: the higher the number of habitat typesavailable, the lower the dominance of onesingle taxon versus all others. When weexcluded Mabuya from this analysis, theexpected negative correlation betweenthese variables was obtained (r = -0.38, n =9, P < 0.05). This shows that the expectedpattern was indeed present in the NigerDelta area, but was masked by the abun-dance and apparent habitat generalism ofMabuya. With regard to the habitat gen-eralism of Mabuya, the specimens of thisgenus were not distinguished and pooled inour analyses.

Mabuya lizards and Agama agamawere the most abundant species in almostall places of the Niger Delta territory, notonly along the transects accurately sur-veyed for quantitative biodiversity, but alsoin all other spots visited. In general,Mabuya species were the dominant lizardsin natural habitats, whereas Agama was thedominant lizard in towns and villages, in-cluding Port Harcourt City. In fact, synecy

of Agama and human populations is wellknown (BUTLER 1986; ANIBALDI & al.1998).

Impact of gas and oil transmissionand extraction systems

The following effects of gas and oiltransmission and extraction activitieswhich can adversely affect reptile popula-tions were observed in the Niger Delta.

(1) industrial effluents can contami-nate waste water; this risk is even higher inNigeria than elsewhere because of the habitof some local people to broach oil trans-mission pipes, and sell oil and fuel on theblack market; (2) habitat loss, fragmenta-tion or partial destruction due to deforesta-tion; (3) modification of the river character-istics (i. e. deviations of the courses, destruc-tion of the original vegetation on the banks,modification of the aquatic vegetation whenpipes are laid in the water); (4) mortality ofanimals caused by working persons andmachines; (5) essential acoustic distur-bance: presence of men (and their ma-chines) at work, and resulting habitat loss;(6) starting of new cultivations, and (7)opening of new access roads to forest andmangrove sites; this can stimulate people touse these prepared areas for their activities(i. e. cultivating, marketing, etc).

To avoid progressive impoverishmentof the reptilian diversity in the Niger Deltaarea, we would suggest that, as a generalrule, oil and gas companies should nolonger lay their oil and gas transmissioninstallations through or in the vicinity ofprimary and secondary rain forest patches,which house both the highest diversity oftaxa and the highest densities of reptilespecimens. River tracts and creaks sur-rounded by primary forest patches deservespecial attention because of the presence ofscattered populations of C. niloticus and C.cataphractus, which are among the mostthreatened reptile species in the NigerDelta (AKANI & al. 1999). In this regard,we strongly suggest to avoid any furtherinterference with the environ ment of theOrashi river course (from Emabu village tothe upper course) and of the Sambreiroriver course (from Degema and Abonnemavillages to the upper course), where pris-tine forest patches are still present.


150 GODFREY e. AKANI & LUCA LUISELLI & EDOARDO POUTANO

2 3 4 5 6

Number of Habitats per Transect

Fig. 2: Correlation between number of habitat types and number of reptile taxaas found in ten transects along the studied liquefied gas pipe-line in the eastern Niger Delta (Nigeria).

Abb. 2: Die Beziehung zwischen der Anzahl der Biotoptypen und der Reptilientaxain zehn Untersuchungstransekten entlang einer Flüssiggas-Pipeline im östlichen Niger Delta (Nigeria).

The mangroves represent another es-sential habitat type in the ecosystem of theeastern Niger Delta. However, the reptilespecies diversity (number of taxa) found inthis habitat type was relatively small incomparison with that of the forest patches(e.g. compare transect T8 with transect T3)and, contrary to what we observed in forestpatches, there is not a single taxon which isfound exclusively in mangrove formations.

Considering that the extension ofmangrove formations is still enormous inthe Niger Delta region (SINGH & al. 1995;POLITANO 1997), we think that - at the lo-cal scale of the Delta - mangroves are cur-rently less threatened than the rainforests.Where ever it is absolutely necessary totake a choice between mangrove and rain-forest (and always in conformity with the

needs of defending the mangrove habitatwhich is one of the most threatened in thewhole world), we suggest that environmen-talists working for the main oil companiesshould give priority to the conservation ofthe few remaining rainforest patches. Inany case, where possible and as a generalrule, already altered dry land should beused for planning industry installations inthis fragile ecosystem, also because of therisks of large scale pollution to aquatic bi-ota when water bodies are used.

Enormous areas of mangroves withconspicuous reptile populations are presentalong the New Calabar river (from the seato approximately Orubiri village), theBonny river, and the Sambreiro river (fromthe sea to approximately Idama village).

ACKNOWLEDGMENTS

We thank "Aquater S. p. A", "Ecosystem s. r. L",and "T. S. K. J. Nigeria Ltd.", Agip-Petroli S. p. A,NAOC ltd., Inst. DEMETRA, for financial support ofthis research and for logistic support in Nigeria. Manypersons of our scientific staff (ENI-Agip group Envi-ronmental Dept.) co-worked with us in the field, andkindly provided original materials and observations.

Amongst them, we express our sincere thank to DrFRANCESCO M. ANGELICI, Dr BRUNO CESCÛN, DrOTONYE L. DA VIES, Dr BENJAMIN EKEKE, Dr IVANAGRIMOD, Dr LEONARDO RAGGI, and Dr ANTONIOSlGISMONDI. Dr BERRY HUGHES (London) and DrULRICH JUGER (Darmstadt) provided useful informationon taxonomy of some Nigerian species.



2 3 4 5 6 7

Number of Habitats per Transect

B

2 3 4 5 6Number of Habitats per Transect

Fig. 3. Correlation between number of habitat types and dominance index of reptiles as found in ten transectsalong the studied liquefied gas pipe-line in the eastern Niger Delta (Nigeria).

A - all taxa included in computation; B - genus Mabuya excluded from computation.Abb 3: Die Beziehung zwischen der Anzahl verfugbarer Biotoptypen und dem Dominanzindex für Reptilien

in zehn Untersuchungstransekten entlang einer Flüssiggas-Pipeline im östlichen Niger Delta (Nigeria).A - unter Einbeziehung und B - unter Ausschluß der Gattung Mabuya von der Berechnung.


152 GODFREY e. AKANI & LUCA LUISELLI & EDOARDO POLITANO

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DATE OF SUBMISSION: March 9th, 1998 Corresponding editor: Heinz Grillitsch

AUTHORS: Prof. Dr. GODFREY C. AKANI, Department of Biological Sciences, The Rivers State University ofScience and Technology, P.M.B. 5080, Port Harcourt, Rivers State, Nigeria; Dr LUCA LUISELLI, Institute of En-vironmental Studies DEMETRA, Via dei Cochi 48, 1-00133 Roma, Italy [fax: +39-6-8411964). Dr EDOARDOPOLITANO, E.N.I. Environmental Department ("Aquater S.p.A."), via Mirabello 53, 1-61047 S. Lorenzo in Campo,Italy [fax:+39-721-731376].


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