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P00791-P00794.Pdf July2000] ShortCommunications 791 SHARPE,R. B. 1909.A hand-listof the generaand spe- Rhodopechys,Carpodacus, Pinicola, Loxia, Uragus, ciesof birds, vol. 5. Taylorand Francis,London. Urocynchramus,and Propyrrhula.American Mu- SIBLEY,C. G., AND B. L. MONROE,Jr. 1990. Distribu- seum Novitates 1786:1-37. tion and taxonomyof birds of the world. Yale VAURIE, C. 1959. The birds of the Palearctic fauna. University Press,New Haven, Connecticut. Passeriformes.H. E & G. Witherby,London. SUSHKIN,P. P. 1927.On the anatomyand classifica- WOLTERS,H. E. 1979. Die Vogelarten der Erde, vol. 4. tion of the weaver-birds. Bulletin of the Ameri- Paul Parey,Hamburg and Berlin, Germany. can Museum of Natural History 57:1-32. Zu$I, R. L. 1978.The interorbitalsepturn in cardue- SWOFFORD,D. L. 1998.PAUP*. Phylogenetic analysis us- line finches. Bulletin of the British Ornitholo- ing parsimony(*and other methods), version 4.0. gists' Club 98:5-10. Sinauer Associates,Sunderland, Massachusetts. VAURIE, C. 1956. Systematic notes on Palearctic Received26 October1998, accepted 23 November1999. birds,no. 20. Fringillidae:The generaLeucosticte, AssociateEditor: A. ]. Baker The Auk 117(3):791-794, 2000 Anatomical and Nutritional Adaptations of the Speckled Mousebird (Colius striatus) COLLEENt. DOWNS,• J. OLAF WIRMINGHAUS,2 AND MICHAEL J. LAWES Schoolof Botanyand Zoology, University of Natal,Private Bag X01, Scottsville3209, SouthAfrica Folivoryis a rare phenomenonin birds that has creasingbody mass, small folivores have proportion- evolvedindependently in severallineages. It hasbeen ally higher metabolicrequirements relative to their reportedin ratites(Withers 1983, Herd and Dawson gut capacitythan do large folivores(Demment and 1984), anatids(Buchsbaum et al. 1986, Dawson et al. VanSoest1985). Consequently,small avian folivores 1989),ptarmigan (Gasaway et al. 1975),the Kakapo have greaterproblems of energy acquisitionthan (Strigopshabroptilus; Oliver 1955, Powleslandet al. theirlarger avian counterparts and thus are expected 1992),and somespecies of Saltator(Bosque et al. 1999). to be rare. The Hoatzin (Opisthocomushoazin) is uniqueamong Mousebirds,order Coliiformes,comprise six spe- birdsin beingthe only documentedforegut fermenter ciesthat are endemicto sub-SaharanAfrica (Maclean andthe only obligate avian folivore (Grajal et al. 1989). 1993).Despite the paucityof species,mousebirds are Althoughfolivory is commonin mammals(Chivers remarkablysuccessful and haveradiated into many and Langer1994), the evolutionof folivoryin birdsis habitatsin Africa, occurringfrom harsh desert to constrainedby bodymass and the highmass-specific moist savanna. The folivorous habits of these birds energyrequirements of endothermyand flight, despite were first observedin the SpeckledMousebird (Col- the apparentunlimited supply of freshfoliage in na- ius striatus;Rowan 1967), althoughpeculiarities of ture.Klasing (1998) defines an avianfolivore as one that their thermoregulatoryability were noticedprior to concentrateson leaves;however, the categories from an this (McAtee 1947). Body temperaturesof mouse- obligatefolivore to a facultativefolivore are poorlyde- birds are correlatedwith ambienttemperature fluc- fined. tuations,which is a putativereason for their noctur- Folivoryis associatedwith reducedfood quality nal huddling behavior(Rowan 1967). Average body (Chiversand Langer 1994). Digestioncosts for fer- mass of each speciesis about 50 g, making mouse- mentingfolivores are high owing to gut specializa- birds amongthe smallestof folivorousbirds. Con- tions (i.e. a fermentation chamber) and associated sequently,we investigatedtheir gastrointestinal tract bacterial micro flora for the breakdown of cellulose and digestivephysiology and suggesthow adapta- and release of volatile fatty acids (VFA; VanSoest tionsin thesetraits permit the combinationof small 1983).Energy from a folivorousdiet is releasedslow- size and folivory in birds. To determinethe extentof ly, requiringa behavioraland physiologicallifestyle morphologicaland physiologicaladaptations, we that minimizesenergy expenditure. Because energy madeoutgroup comparisons with the Purple-crested requirementsper unit body massincrease with de- Turaco(Tauraco porphyreolophus ), which is a largefor- est frugivore,and with the Hoatzin. Methods.--Wecollected 13 Speckled Mousebirds E-mail: [email protected] from Creighton(30ø02'S, 29ø46'E) and Pietermaritz- Deceased. burg (29ø36'S,30ø24'E), South Africa, in May 1994 792 ShortCommunications [Auk, Vol. 117 TABLE1. Body and gut dimensionsof Coliusstriatus, C. colius,Urocolius indicus, and Tauracoporphyreolophus. Valuesare œ+__ SE, with n in parentheses. Urocolius Tauracoporphyreolo- Parameter Coliusstriatus C. colius indicus phus Body mass(g) 50.7 ---2.27 (13) 38.8 _+0.4 (15) 50 (1) 318.1 _+41.49 (4) Esophaguslength (mm) 44.1 ___1.30 (13) 44.6 + 0.7 (27) 30 (1) 64.2 _+3.59 (4) Proventriculuslength (mm) 21.3 ---0.99 (12) 17.2 _+0.7 (27) 26 (1) 26.2 + 2.21 (4) Ventriculuslength (mm) 21.9 ---0.86 (13) 12.2 + 0.5 (27) 14 (1) 31.6 + 2.60 (4) Ventriculusbreadth (mm) 15.7 ---0.67 (12) 14.1 _+0.5 (27) 19 (1) 20.8 _+3.78 (4) Small intestine-cloacal(mm) 225.8 --- 7.54 (11) 169.0 _+4.7 (27) 260 (1) 401.0 _+38.15 (4) Total GIT length (mm) 312.4 + 8.59 (11) 244.0 _+4.7 (27) 330 (1) 522.9 _+38.36 (4) GIT:HB indexa 46.86 (11) 54.39 (1) 66.59 (4) GIT:mass indexb 61.42 (11) 70.65 (1) 59.09 (4) Lengthof smallintestine to cloaca/cuberoot of head-bodylength (mm). Lengthof smallintestine to cloaca/cuberoot of body mass(g). and dissectedthe fresh specimens.Body measure- were pigmentedblack. The ventriculusin situwas a ments and gastrointestinaltract (GIT) dimensions prominentorgan, lying to the lower left sideof the were made with vernier calipersand a steel ruler abdomenwith the anterior dorsal portion slightly (+0.5 mm). Under a dissectingmicroscope, we de- behindthe posteriorend of a liver lobeand the pos- terminedthe proportionof leaf material,fruit, and terior dorsalportion lying directlyagainst the dorsal seedin the ventriculusof eachspecimen. Sections of bodywall. Thebody wall wasnonmuscular here and the GIT of C. striatuswere fixed in 5% bufferedglu- the skin pigmentedblack, with featherson the ex- taraldehydeat 4øCas initial preparationfor scanning terior surface.The GIT length:bodylength index of electronmicroscopy. The remainingsections of the SpeckledMousebirds was lower than that of the Pur- GIT were fixed in Bouin's solution from which his- ple-crestedTuraco, whereas the GIT length:body tologicalsections were prepared;these were stained massindex was higher (Table1). with eosinhaematoxylin to showthe histologicalar- Both the Speckled Mousebird and the Purple- rangementof the gastricepithelia. Immediately after crestedTuraco showed a typical gastricplan with dissection,the gut contentsin the ventriculiof eight mucosa,submucosa, and muscularis.In theSpeckled C. striatuswere identified to food categoryas noted Mousebird,the musculaturewas less extensively de- aboveand then placedin clearscrew-top vials, fro- velopedand showedno elastictissue compared with zen, and analyzedfor VFAs using gas chromatog- the Purple-crestedTuraco. The main differencebe- raphyby the Animal Nutrition and Animal Produc- tweenthe two specieswas the extensivesurface area tion Institute, Irene, South Africa. Gut contents of the in the ventriculusof the SpeckledMousebird owing remainingindividuals were spreadon a glassslide to the extensive convolutions, whereas few convo- and viewed using a dissectingmicroscope and per- lutionswere presentin the Purple-crestedTuraco. centagesof food categoriespresent estimated. Scanningelectron microscopy revealed the ven- We dissectedone Red-facedMousebird (Urocolius in- triculus to be coveredby speciosemicrobial flora. dicus)and four Purple-crestedTuracos obtained from The microbial flora included sphericaland ovoid the Durban Natural Science Museum to obtain GIT di- bacteriaarranged in long chainsor pairs (possibly mensions. Furthermore, 27 White-backed Mousebirds Planococcusor Rumenococcus;Sneath et al. 1986) and (Coliuscolius) collected in Prieska(29ø39'S, 22ø46'E) and othersthat were rod-shaped.Further research is re- PrinceAlbert (33ø14'S, 22ø02'E) by A. McKechniewere quired to isolate,culture, and classifythe microor- dissectedfor GIT dimensions.Gross morphology of the ganismsin the foregutof mousebirds. GIT of all specimenswas examinedusing a dissecting We recordedthe presenceand concentrationof microscope. VFAs in the proventriculusand ventriculuscontents Results.--Thestructure of the GIT of the Speckled of SpeckledMousebirds: i.e. aceticacid, propionic Mousebirdconsisted of an esophagusleading into a acid,iso-butyric acid, n-butyric acid, iso-valeric acid, well-developedproventriculus that was composed of and n-valericacid (Table2). Small peaksindicating a muscularwall with a grosssmooth inner surface other VFAs were also seen. exceptfor a gracilereticulate pattern; no cropwas ev- The gut contentsof SpeckledMousebirds varied ident. The proventriculusled to the ventriculus, widely in the amountof leaf materialpresent (0 to which was simple,unilobular and thin-walled.The 100%).A high proportionof fruit was eatenwhen inner wall of the ventriculuswas a mat of adpressed leaf intakewas low. The meanproportions (n = 10) papillae.The surfacearea of the ventriculusof the of dietary categoriesin the ventriculuswere 43.2 +__ Speckled Mousebird is large becauseof extensive SE of 10.5% leaves, 54.3 + 10.4% fruit, and 2.5 + 2.5% convolutions.It also has some areas (>50%) that seeds.We
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