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Social Monogamy in the Noki Or Dassie-Rat (Petromus Typicus) in Namibia

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Social Monogamy in the Noki Or Dassie-Rat (Petromus Typicus) in Namibia ARTICLE IN PRESS www.elsevier.de/mambio Original investigation Social monogamy in the noki or dassie-rat (Petromus typicus) in Namibia By G.B. Rathbun and Carolyn D. Rathbun Department of Ornithology and Mammalogy, California Academy of Sciences, San Francisco, CA, USA Receipt of Ms. 25.5.2005 Acceptance of Ms. 27.1.2006 Abstract The noki or dassie-rat (Petromus typicus) is a little-studied diurnal hystricognath rodent endemic to rocky outcrops in south-western Africa. An intermittent study in Namibia of 11 radio-tagged individuals, with over 250 h of direct observation, was done between 2000 and 2004 to document their basic natural history, including their social organization. The study was terminated when noki numbers collapsed, probably due to predation by an expanding population of black mongooses (Galerella nigrata) that was unintentionally provisioned with food at a nearby eco-tourist lodge. Male and female adults were distributed as monogamous pairs on home ranges that were similar in size and shape. The pair bond was strong, with many coordinated behaviours. Reproduction was seasonal with litters of single highly precocial young. Multiple generations remained on the parental home range without significant aggression. Most aggression was between adult males, while adult females showed tolerance towards each other and intruding adult males. Although noki social monogamy is probably the result of male mate guarding, considerable paternal care was observed, including allogrooming and vigilance against predators. Nokis have a relatively low metabolic rate and numerous thermoregulatory behaviours that suggest that nocturnal huddling by the male with his mate and young may be an important factor in the evolution of their social monogamy. r 2006 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. Key words: Petromus, dassie-rat, monogamy, Namibia, social organization Introduction The noki or dassie-rat (Petromus typicus A. mentioned in a published study by George Smith, 1831) belongs to the monospecific and Crowther (1981). hystricognath rodent family Petromuridae. Nokis are endemic to the southwest arid We use the common name ‘‘noki’’ because it biogeographical region of Africa (Meester avoids the confusion by many people with 1965) that stretches from extreme south- dassies (hyraxes in the mammalian order western Angola south through Namibia into Hyracoidea) or true rats (rodent families north-western South Africa. They are re- Muridae and Cricetidae). Noki is derived stricted to rocky or boulder-strewn habitats, from a Khoekhoegowab (=Hottentot) especially along the Namibian escarpment dialect (Shortridge 1942) and was first zone with its numerous mountains, cliff faces, 1616-5047/$ - see front matter r 2006 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2006.01.008 Mamm. biol. 71 (2006) 4 Á 203–213 ARTICLE IN PRESS 204 G.B. Rathbun, C.D. Rathbun inselbergs, and kopjes or rock outcrops Annual mean rainfall at Omaruru Prison is (Coetzee 2002). The aridity of the escarpment 292.9 mm, with virtually all of this falling during and closely related Namib Desert is at least the months of November through April. Annual 15 million years old (Ward and Corbett 1990) average minimum and maximum temperatures at the prison are 11.4–31.0 1C, with May through and the noki apparently has had an ancient August being the coolest as well as driest months association with these biomes (Meester 1965; (Rathbun and Rathbun 2005). Se´ne´gas 2004), as demonstrated by several After determining the suitability of the study site in morphological adaptations to living in rock June 2000, we radio-tracked and observed nokis crevices (George and Crowther 1981; Skinner during four subsequent periods that covered the and Chimimba 2005). These features include seasonal climatic extremes: 25 December 2000–5 a flattened cranium, flexible ribs, and dorso- January 2001, 5 September–21 November 2001, lateral mammae. 24 April–7 July 2002, and 11 May–23 July 2003. Apart from general natural history observa- tions (e.g., Skinner and Chimimba 2005) and Trapping and marking reports based mostly on opportunistic ob- servations (e.g., Coetzee 1983) or museum We captured Petromus with Sherman model XLF15 (11.5 Â 10 Â 30.5 cm folding aluminium) collections (Coetzee 2002), there are only two and Tomahawk model 201 (40.5 Â 13 Â 13 cm published field studies of noki ecology (With- single door) live traps set during daylight hours ers 1979; George and Crowther 1981). and baited with fresh carrots or apples. To prevent Recently, reproduction and behaviour of hyperthermia in captured animals we positioned captive nokis have been studied (Mess 2002; traps in the shade or avoided trapping during Mess and Ade 2005) in association with mid-day. research on their placentation (Mess 2003). Trapping nokis was difficult because a few Because of the paucity of long-term and individuals became trap happy and disrupted detailed field data and the limited distribu- attempts to capture trap shy individuals. Intruders were especially trap shy because of their wariness tion of nokis, there is considerable interest in and preoccupation with exploration. When food this near-endemic Namibian rodent (Griffin plants were plentiful during the wet season nokis 1998). The aim of this study was to investi- were especially difficult to trap, whereas during the gate the social behaviour and social organi- dry season they were more easily attracted to fresh zation of free-ranging nokis in Namibia. moist bait. We permanently marked nokis for subsequent identification with passive integrated transponder Material and methods chips (Destron-Fearing, South Saint Paul, MN) implanted subcutaneously (Schooley et al. 1993) Study site and study period between the shoulders, although care had to be Our study was done near the Erongo Wilderness taken because of their fragile skin (G. B. Rathbun Lodge (21127.679S, 15152.523E) on Okapekaha and C. D. Rathbun, unpublished data). For visual Farm, about 10 km west of Omaruru town in the identification we bleached pelage on different parts foothills of the Erongo Mountains of Namibia. of the body using a mixture of 20% hydrogen The site is 1240 m above sea level and is character- peroxide and commercial hair bleaching powder ized by huge rounded granite dikes that rise about mixed into a paste. We thoroughly washed the 100 m above the surrounding peneplain and smaller paste out of the fur with water after about 5 min. 10–20 m high granite outcrops or kopjes sur- We divided nokis into three age-classes. Adults, rounded by intruding fingers of the surrounding which were capable of reproduction, were over bushveld. The vegetation at the study site is 160 g. Subadults, which were independent but not composed of widely spaced low trees and bushes reproductive, were less than 160 g. Young, which interspersed with seasonally dense annual and were neonatal through weaning, were below 100 g. perennial forbs and bunch grasses. The dominant trees include Combretum apiculatum, Sterculia Radio tracking and visual observations africana, Terminalia prunoides, and Boscia albi- trunca and the more common bushes include We attached radio transmitters (Holohil Systems several species of Grewia, Croton gratissimus, Ltd., Carp, Ontario, Canada; model MD-2C, 2.2 g Dichrostachys cineria, and Mundulea sericea weight, 120-day battery life, 20-pound-test and (Rathbun and Rathbun 2005). 10-cm-long fishing leader wire whip antenna) with ARTICLE IN PRESS Social monogamy in the noki or dassie-rat 205 the standard Holohil mammal collars made of idea of the social organization of nokis. In antenna wire inside Tygon tubing. We radio- 2002, however, the four adults were clearly located nokis several times a day between 04:30 distributed as male–female pairs on highly and 22:30 h, but separated consecutive fixes by at congruent and exclusive home ranges, least 2 h, which was sufficient to reduce inter-fix and each pair had a single young (Tab. 1, autocorrelation. The movement of radio-tagged nokis, along with distortion of radio signals by Fig. 1B). The social structure in the other granite rock faces and kopjes, often made triangu- years can only be understood in the context lation difficult. Therefore, we used a combination of several factors (age-classes of individuals of homing and triangulation (Kenward 2001)to (Tab. 1), individual behaviours, recent mor- determine locations (fixes). We flagged locations tality, and the presence of unmarked trap-shy made at night and determined coordinates the next nokis) that are not apparent by considering day. Young nokis were not radio tagged so their the home range boundaries alone. locations were based on sightings alone. We In 2000, our study period was short and we calculated universal transverse Mercator (UTM) gathered relatively few location fixes (Tab. 1) coordinates for locations by determining the distance and azimuth to or from a ‘‘target’’ site and did not trap neighbouring nokis to the with known coordinates, which we determined with east (Fig. 1A versus B). Also, an unmarked a global positioning system (GPS) receiver at the adult female associated with adult male 209 beginning of our study. We then calculated was killed while still in a Tomahawk trap by coordinates for several other prominent target a black mongoose (Galerella nigrata)on26 kopjes that were visible within 100 m from any December, before we had time to determine spot on the study area with the computer software her spatial and social relationship to the program UTM CALC (O’Leary 1998). The preci- other nokis. sion of our laser rangefinder (Bushnell model In 2001, the number of nokis present on our 400X) and sighting compass (Brunton model 16- study site increased to six (Tab. 1), including FSM360LA-SME) was 71.0 m and 70.51, respec- tively. The accuracy of our location calculations three from the year before.
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