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Reproductive Biology of the West African Savannah Frog Hyperolius Nasutus GÜNTHER, 1864

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Reproductive Biology of the West African Savannah Frog Hyperolius Nasutus GÜNTHER, 1864 ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Herpetozoa Jahr/Year: 2006 Band/Volume: 19_1_2 Autor(en)/Author(s): Rödel Mark-Oliver, Lampert Kurt, Linsenmair Karl Eduard Artikel/Article: Reproductive biology of the West African savannah frog Hyperolius nasutus Günther, 1864 (Amphibia: Anura: Hyperoliidae) Fortpflanzungsbiologie des westafrikanischen Savannenfrosches Hyperolius nasutus Günther, 1864 (Amphibia: Anura: Hyperoliidae) 3-12 ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at HERPETOZOA 19 (1/2): 3 - 12 Wien, 30. Juli 2006 Reproductive biology of the West African savannah frog Hyperolius nasutus GÜNTHER, 1864 (Amphibia: Anura: Hyperoliidae) Fortpflanzungsbiologie des westafrikanischen Savannenfrosches Hyperolius nasutus GÜNTHER, 1864 (Amphibia: Anura: Hyperoliidae) MARK-OLIVER RODEL & KATHRIN P. LAMPERT & K. EDUARD LINSENMAIR KURZFASSUNG Wir untersuchten das Paarungsgeschehen bei Hyperolius nasutus GÜNTHER, 1864 an einem temporären Savannentümpel im Comoé-Nationalpark, Elfenbeinküste. Hyperolius nasutus ist eine Art mit ausgedehnter Re- produktionszeit ('prolonged breeder'), die etwa 2-5 Monate nach dem Einsetzen der Regenzeit beginnt. Rufende Männchen saßen bevorzugt auf Seggen, die im Wasser wuchsen. Andere syntope Riedfroscharten wählten andere Rufwarten. Wir fanden Hinweise darauf, daß bei günstigen Wetterbedingungen innerhalb einer Regenzeit repro- duzierende Männchen zweier aufeinanderfolgender Generationen zeitlich versetzt am Gewässer auftreten können. Die Größe (Länge und Masse) der Männchen erlaubte keine Vorhersagen über deren Paarungserfolg. Weibchen entschieden sich im Wahlversuch nicht bevorzugt für das größere Männchen. Im Durchschnitt betrug die Masse des Männchens 80% der Masse des laichreifen Weibchens, mit dem es verpaart war. Mögliche Erklärungen dafür, daß Weibchen nicht die größten Männchen wählten, werden diskutiert. ABSTRACT We studied the mating activities of Hyperolius nasutus GÜNTHER, 1864 at a temporary savannah pond in the Comoé National Park, Ivory Coast. Hyperolius nasutus is a prolonged breeder that started reproducing 2-5 months after the beginning of the rainy season. At that pond, males preferably called from sedges growing in shallow water, whereas other syntopic hyperoliid species chose different calling sites. We found evidence that two con- secutive generations of reproducing male H. nasutus were sequentially present at the pond within one rainy sea- son. Neither male body length nor mass predicted male mating success. Females did not preferentially select larg- er over smaller males. On average, a female mated with a male which had about 80% of her own body mass. We discuss possible explanations for this behavior. KEY WORDS Amphibia: Anura: Hyperoliidae: Hyperolius nasutus, female choice, reproduction, population ecology, behavior, savannah, Comoé National Park, Ivory Coast, West Africa INTRODUCTION Most West African savannah anurans the explosive breeding species, the number seem to pursue one of two different repro- of male and female individuals at a pond is ductive strategies: species that spawn within more or less balanced or slightly male-biased, one to three days after a rainfall event (ex- Anurans that mainly reproduce towards the plosive breeders) and species whose breed- middle and the end of the rainy season are ing period extends more or less continuously almost exclusively prolonged breeders. At over weeks or even months (prolonged that time reproduction is less dependent on a breeders; RODEL 2000). Under the climatic definite rainfall event. The males congregate conditions of the study area, explosive breed- in large numbers almost every night in cho- ing is largely restricted to the beginning of ruses around the breeding ponds while the the rainy season (RODEL 1998, 2000). At this number of females present is comparatively time both amount and temporal pattern of small. In some prolonged breeders calling precipitation are highly unpredictable, how- and spawning sites are identical (RODEL ever rainfall is essential for reproduction. In 1998). In this case, female choice of the mat- ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at M.-O. RODEL & K. P. LAMPERT & K. E. LINSENMAIR ing partner might be influenced by the quali- duction during the rainy season (RODEL ty of the site. In other species males call at 2000) in that larger choruses usually form some distance from potential spawning sites during August / September in the second and do not offer any site information for half of the regional rainy season. Since its females. If the male provides no direct ben- reproductive period lasts several months, //. efits in terms of increased breeding success, nasutus is a prolonged breeder. Eggs are we would expect females to select indirect attached to submerged vegetation (RODEL benefits such as "good" genes (FISHER 1958) 2000). Little is known about the biology of or high genetic variability (POMIANKOWSKI the species. In this study, we aimed to gain 1988). general information on the reproductive Hyperolius nasutus GÜNTHER, 1864 is biology of H. nasutus including data on the a common West African savannah frog. In possibility of differential choice of males by southern Comoé National Park, Ivory Coast, females. H. nasutus is the last species to begin repro- MATERIALS AND METHODS Study site information on geography, geology and vegetation of the CNP. The study was conducted in the rainy seasons of 1996 and 1999 in the Comoé Study animals National Park (CNP, 3°07' - 4°26' W, 8°30' - 9°37' N), Ivory Coast. The CNP Hyperolius nasutus was believed to be comprises different vegetation zones from a widespread African savannah frog, rang- the Guinea to the Sudan savannah. Our ing from West into southern Africa (SCHIOTZ study site was within open tree savannah 1999; RODEL 2000). Recently CHANNING et (POILECOT 1991) in the northern part of the al. (2002) revised the sharp-nosed reed Guinea savannah. During the dry period frogs on the basis of advertisement calls. (November to March / April), open water They concluded that two species occur in normally persists only within the river West Africa: a forest dwelling species (H. beds. During the rainy season (mean nasutus) and a savannah form that they annual precipitation: 900-1,100 mm) a assigned to H. acuticeps AHL, 1931. The large number of ephemeral savannah latter species which should include H. lam- ponds form. Our specimens all originated ottei LAURENT, 1958 was believed to occur from the 'Hyperolius pond' (HP) appr. 5 in CNP. However, we never found any H. km North of the Comoé Research Station nasutus like frog in more than 20 West and (8°45,1'N, 3°49,0'W). This pond was Central African forest sites, which we inves- chosen because, based on our previous tigated between Guinea and Central African knowledge, this pond seemed to very well Republic. All H. nasutus specimens record- reflect a typical breeding site for West ed by us were exclusively found in savan- African H. nasutus populations (RODEL nah habitats. Hyperolius lamottei, although, 1998, 2000; K. P. LAMPERT, K. E. LINSEN- having a call very similar to that of//, nasu- MAIR & M.-O. RODEL unpubl. data). The tus from CNP, is clearly a morphologically HP is a large savannah pond of approxi- distinct species (RODEL & ERNST 2003) that mately 50 m x 15 m surface and a maxi- is not closely related to H. nasutus (samples mum water depth of one meter. Two to from various West African localities; J. three months after the onset of the rainy KOSUCH and M.-O. RODEL unpublished data season it harbors a rich vegetation that is from mitochondrial 16S rDNA). We agree mainly made up of different grass species with CHANNING et al. (2002) that the name and water lilies. A more detailed descrip- H. nasutus (sensu SCHIOTZ 1999; RODEL tion, including the amphibian species 2000) probably comprises several cryptic breeding in this pond is given by RODEL species. However, as long as the systemat- (1998). POILECOT (1991) provides more ics of this species group is not sufficiently ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at Reproductive biology of Hypewlius nasutus GÜNTHER, 1864 elucidated we continue to refer to CNP frogs In addition, two 8 m x 8 m sampling as H. nasutus. Voucher specimens from plots were searched for calling sites three CNP are stored in the collection of the times (28 and 30 May, 4 June 1996). In the Staatliches Museum für Naturkunde, Stutt- first plot, one quarter was covered with E. gart (SMNS 8968.1-2, 9679.1-11), the re- acutangula, the other three quarters were search collection of the senior author (MOR covered with various Poaceae, and a medi- 9678) and the Institute of Zoology at Mainz um sized tree (4 m high, crown diameter: 3 University (tissue samples). m). Plot 2 was half grown over with E. Other syntopic hyperoliid frogs record- acutangula and half with various Poaceae ed in this study with activity times that match and other herbs. In these plots all H nasu- those of//, nasutus, are either typical savan- tus males were registered including data on nah species: Hypewlius nitidulus PETERS, their density, type of calling site, its height 1875, Afrixalus vittiger (PETERS, 1876) and and distance to the edge of the pond. A. weidholzi (MERTENS, 1937), or normally in- Additionally, we measured
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