ann en hi i Tr 2 in za a tr Rat BR in 22 \ ERROR UNSEREN rt a Dal ae) iR nf aasA aan UBER ANA nase # “ Var Altar sc NEN Mh a ER ARTEN Ye NR aereA> DrErn Te De: ER An: 3 Veen ma han oa x “ ee Eee A ass r EEE ar year ESN Ah mm area Ra Ne . iR BEN Ye ih. Be Ba FR Srusı Ener a vr en rm ar ht EN Run, R “ ErEErueL PETER ERE z Y ur S ET Be ges BESPRE U GERLRATT uns PrRET BELZEIP ER E 7 N j a i Kr! . an IR) TE DET. 3 DEE ER LEE a), NER, PIMRPENELLETD DU EIS S> Ne & {on DC. I NOILNLILSNI N ano S31IUVYaAlII_ LIBRARIES —_ | E32; = (02) = sSs318V MS V/ e (0) Anus INSTITU SMITHSONIAN INSTITUTION INSTITU Ian NENosn NVYINOSHLINS S31UV: SL 3 INSTITU NI NVINOSHLINS, _S31 UV? EZ — Ne = S Bee mo I. oO: oO SLR 5 Q 2 7) 7) NN xD {2} = Sr I en = & == : > - = 7: = > = = GG = 77) 2 12) Y (77) © 7) -S SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWNS S31IUVY4EI1 LIBRARIES S a = * RN N >= ri DIE r a SS : &. r | 2, = = ZEN & = = 12 # & © NEN (= E we = = 4 Q m I = m % 5 = fe 6) = I == ar 2 Sl >= = NI Ss314vV4yg11 LIBRARIES SMITHSONIAN _INSTITUTION NOILNLILSNI_N zZ = a 3 5 z 2 = ee) — = X — ww ) En = -, = 2 = 2 | > 2 > > > = > z = 2 = a = = 0) ar ® = = Hi h = 5 = R -S „SMITHSONIAN INSTITUTION „NWINOSHIINS,, = =" = a = < = < = = zZ - zZ = z - _ = Z = = ZN Se = Den 7 2 5. 2 NI_NVINOSHLINS S31UVUA17 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_N N 4 e n z = zZ (77) u 77) wu 7 wi 77) = = = ee = «x = | = .< Zi < te < = B = = . = = = =) = ° a: [e) = =) z RN | z 2 > — zZ -S SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31UVY4811 s 4 re 2 rm > m & S .D ° == e) = = = ee) = {00} = 5 el 5 Ey = D 5 | > E > > E: 2 > > a nn ZN = - > E = > U = Mm N e z NI NVINOSHLINS S319V4 817 SMITHSONIAN INSTITUTION _NOILNLILSNI_ I = 2) zZ 2) z RR 2) = .ZS Su = .< = < en < zZ I ,fl fs‘ zZ = 2 SS I z oO je = e O «: a e) N, 7 oO & 2 2 2 N: 2 u = = z = = 2 BR = 7 ı = ma -S SMITHSONIAN _ INSTITUTION NOILNLILSNI NVINOSHLINS S31UVAUgI1 LIBRARIES S on z 2 NE 2 u 3877 3E = ES: = = | LT an i SON In . < F s c < = I X c <. a % 5 = 5 ns ö = ) « _ RR UB z — NI S3IUVYgıI1_LIBRARIES_SMITHSONIAN _ INSTITUTION NOILNLILSNI_N = 5 \ Be: 5 = (es) _ => G w = w E) n = = 0; E) = a > au ef F 7 Ri m a. m S m Pe m i m — > er | S4 „INSTITUTION, NOILNLILSNI _NVINOSHLINS S31Uvaa17 LIBRARIES S Ss an & N = “ EEE = m; an, 3 WA EN. 5 MM. FE 5 Mn = Ne a >iee 3: E (o EN =) > (& BE > RN, Y E \ 5 a m N JE” 2 m oc = nu m N m Nonne o = [@7) — (dp) = (07) = an \s sa1uvualıı “ „NOLALILSNI is 2 „@ = < < = = 4, — I 5 R I \ E . 2 R 5 5 \ En 2 , #7, 5 = | 6; = o, Eu Oo Gl r e = E z E 2 G 2 z (77) ei i (77) N_ INSTITUTION NOILNLLLSNI_ NVINOSHLUINS S31UVYAIT_LIBRARIES SMITHSONIAN_ | . = ZN 6 = 5 oe & = RAIN R u Eu = 5 a IN < 3 <; = Ian PAIN x cc 3 si I "Da 5 a 5 = _ = Em zZ 37 " = S„S31UVUAIT LIBRARIES” SMITHSONIAN” INSTITUTION NOLLNLILSNI_NVINOSHLINS” | (©) = o er _ 5 SER 5 ver & .—— & >= & N = 5 E 5 | > e N Sı2 bb > > > = EU Ss r E: z = 'z = E N = > m a r = m = SI un = 7) = 7) z 7) I = N_ INSTITUTION NOILNLILSNI_ NVINOSHLINS S31yvygı7 LIBRARIES SMITHSONIAN | zZ (02) 2 (2) un ”o zZ < = < = < = < | ö 5 .ö Br S = S [7] . 0 RT) 72) 7) .@ N) I. Ne! I [®) I IN O I = 2, E= < 3 S 2 Lad = >" = >' = IS > = (77) zZ (77) z (77) m 2 = LIBRARIES SMITHSONIAN_ INSTITUTION NOLLNLLLSNI_ NYINOSHLINS” E 6 u = u z 1 x „Z = en! m Zi X BE < < u < = = m ai 2 En = u. N = ” 2 = = = u 203 N _ INSTITUTION NOILNLILSNI NVINOSHLINS SI3IUVYAIT LIBRARIES SMITHSONIAN | L = Zu Se p4 r> zZ 1 «u = o = er 2 m a = 2 E e = » = m > p > re % > 3 In. = = zZ r a % z m 2 n z 2 SS31UVUAIT LIBRARIES, B NOILALILSNI = RR ie < = z = = zZ SA 3 a = 'z 3 F % 2 NN 3 oO: Se ) T oO’ 2? N 2 27) (70) 7) (97) 2 ee. = = E = E a. 2.20 05 = N_ INSTITUTION NOILNLILSNI_NVINOSHLINS S31UVYSAIT_LIBRARIES SMITHSONIAN_| ‚o an a u = ” z a ee a = 4 — 4 e\ < z\3 2 ur m = er x — X c Be e = fs) Sn — — [e) {ee} — [00] I 2 | er = er | vo > Gr = s_S31uvual1_LIBRARI ES_ SMITHSONIAN_ INSTITUTION NOILNLILSNIZ „S S u u ° eı = 5 e) = DAfp, D 2 ei) > e > > LAD 2 > = E Do = 4 ZA - En == 5 = > ,%f £ = = = = z m 2) m u = un = 7) z | 7) r z | un z N INSTITUTION „NVINOSHLIWNS S31UVyaıT LIBRARIES, .ıl IR N u 2 ME. = am 2 z U. 5 3 ae 2 Mn. 2 U. 3 ma = BER f0 ZEITSCHRIFT FÜR SAÄUGETIERKUNDE INTERNATIONAL JOURNAL OF MAMMALIAN BIOLOGY Organ der Deutschen Gesellschaft für Säugetierkunde Volume 356, 1991 ISSN 0044-3468 Herausgeber / Editors P. J. H. van Bree, Amsterdam — W. Fiedler, Wien — H. Frick, München — W. Herre, Kiel - H.-G. Klös, Berlin — H.-J. Kuhn, Göttingen - E. Kulzer, Tübingen —- B. Lanza, Florenz — W. Maier, Tübingen — J. Niethammer, Bonn — Anne E. Rasa, Bonn — H. Reichstein, Kiel — M. Röhrs, Hannover — H. Schliemann, Hamburg — D. Starck, Frankfurt a. M. —- F. Strauß, Bern - E. Thenius, Wien — P. Vogel, Lausanne Schriftleitung/Editorial Office D. Kruska, Kiel — P. Langer, Gießen Mit 141 Abbildungen v Verlag Paul Parey Hamburg und Berlin u 17 5 % ni D | m Lo er 1223 } ee rat 7 Be: . u an Wissenschaftliche Originalarbeiten ALCANTARA, M.; TELLERfA, J. L.: Habitat selection of the Wood mouse (Apodemus sylvaticus) in cereal steppes of Central Spain. - Habitatwahl von Waldmäusen (Apodemus sylvaticus) ın Gesneidesteppen Zentralspaniens er... 00 u haeel en ne en ee le Le N AscaRAY, C. M.; PERRIN, M. R.; McLacHtan, A.; Eıs, S. F.: Population ecology of the Hairy- footed gerbil, Gerbillurus paeba, in a coastal dunefield of South Africa. — Populationsökolo- gie des Rauhfußgerbills Gerbillurus paeba in einem Dünenfeld an der Küste von Südafrika . . BRANDL, R.; BEZZEL, E.; REICHHOLF, ]J.; VÖLKL, W.: Population dynamics of the Red squirrel in Bavaria. - Populationsdynamik des Eichhörnchens in Bayern ........ 2.2.20... BRITTON-DAVIDIAN, JANICE; VAHDATI, MEHRNOUCH; BENMEHDI, FATIMA; GROS, PAULE; NANCE, VALERIE; CROSET, H.; GUERASSIMOV, S.; TRIANTAPHYLLIDIS, C.: Genetic differ- entiation in four species of Apodemus from Southern Europe: A. sylvaticus, A. flavicollis, A. agrarins and A. mystacinus (Muridae, Rodentia). — Genetische Differenzierung bei vier Apodemus-Arten in Südeuropa: A. sylvaticus, A. flavicollis, A. agrarius und A. mystacinus (Mluneidlae; None) In FE TEN RE SR SL EN ER E RORRE EN BRÜNNER, H.; NEET, C. R.: A parapatric scenery: The distribution and ecology of Sorex araneus and $. coronatus (Insectivora, Soricidae) in southwestern Germany. — Die parapatrische Szenerie: Die Verbreitung und Okologie von Sorex araneus und Sorex coronatus (Insecti- oraWSorierdae in Südwestdeutschland 2 2.0. 0 zen een le Felle BRUORTON, M. R.; PERRIN, M. R.: Comparative gut morphometrics of Vervet (Cercopithecus aethiops) and Samango (C. mitis erythrarchus) monkeys. — Vergleichend-morphometrische Untersuchungen am Verdauungskanal von Grünaffen (Cercopithecus aethiops) und Diadem- alten (CET ET RENNER EEE EL AN OR a EEE COMPARATORE, VIVIANA M.; MACEIRA, N. O.; BuscH, CrısTına: Habitat relations in Cteno- mys talarum (Caviomorpha, Octodontidae) in a natural grassland. — Habitat-Beziehungen von Ctenomys talarum (Caviomorpha, Octodontidae) auf natürlichem Grasland ....... Errıson, G. T. H.; SKINNER, J. D.: Thermoregulation and torpor ın African woodland dormice, Graphiurus murinus, following cold acclimation. — Thermoregulation und Torpor bei afrıkanischen Baumschläfern, Graphiurus murinus, nach Kältebelastung............ Espınosa, MARIA BEATRIZ; REIG, ©. A.: Cytogenetics and karyosystematics of South American oryzomyine rodents (Cricetidae, Sigmodontinae). Part III. - Zytogenetik und Karyosyste- matık von südamerikanischen Reisratten (Cricetidae, Sıgmodontinae) .... 2.2.2222... FLADERER, F. A.: Der erste Fund von Macaca (Cercopithecidae, Primates) im Jungpleistozän von Mitteleuropa. — The first find of Macaca (Cercopithecidae, Primates) in the Late Hleistocenesol Central Europe Ge FE nee Nena eigape GRANJON, L.; VAssaRT, M.; GRETH, A.; CRIBIU, E.-P.: Genetic study of Sand gazelles (Gazella subgutturosa marica) from Saudi Arabia. Chromosomal and isozymic data. — Genetische Untersuchungen an Sandgazellen (Gazella subgutturosa marıca) aus Saudi-Arabien. Chro- mosomalesundlelektrophoresische, Daten p e e nn HERZIG-STRASCHIL, BARBARA; HERZIG, A.; WINKLER, H.: A morphometric analysis of the skulls of Xerus mauris and Xerus princeps (Rodentia; Sciuridae). -— Morphometrische Analyse der Schädel von Xerus inauris und Xerus princeps (Rodentia, Sciuridae)........ InGoLD, P.; MARBACHER, H.: Dominance relationships and competition for resources among chamois Rupicapra rupicapra rupicapra ın female social groups. -— Dominanzbeziehungen und Konkurrenz um Resourcen in Gruppen von weiblichen Gemsen (Rupicapra rupicapra VODRGDNE) No ee ee en oa N A Eee REG JacoBsen, N. H. G.; NEwBERY, R. E.; DE WET, M. ].; VILJoEn, P. C.; PIETERSEn, E.: A contribution of the ecology of the Steppe pangolin Manis temminckü in the Transvaal. — Ein Beitrag zur Okologie des Steppenschuppentieres Manistemminckü in Transvaal ....... KeLT, D. A.; Parma, R. E.; GALLARDO, M. H.; Cook, J. A.: Chromosomal multiformity in Eligmodontia (Muridae, Sigmodontinae), and verification of the status of E. morgani. - Chromosomale Vielfalt bei Eligmodontia (Muridae, Sigmodontinae) und Bestätigung des STIUSVOML AN IONgaN En ne nee Korz, V.: Social relations and individual coping reactions in a captive group of Central American Agoutis (Dasyprocta punctata). — Soziale Beziehungen und individuelle Bewälti- gungsreaktionen in einer Gehegegruppe mittelamerikanischer Agutis (Dasyprocta punctata) . Kvam, T.: Reproduction in the European lynx, Lynx Iynx. — Fortpflanzung beim europäischen Lues, messen ora 06 men ol oc A ee MADE, J. VAN DER: Sexual bimodality in some recent pig populations and application of the findings to the study of fossils. - Geschlechtsbimodalität bei einigen rezenten Schweinepo- pulationen und Anwendung der Befunde für das Studium an Fossilien... ........... Near, B. R.: Seasonal changes in reproduction and diet of the Bushveld gerbil, Tatera leucogaster (Muridae: Rodentia), in Zimbabwe. — Saisonale Schwankungen in der Fortpflan- zung und in der Nahrungszusammensetzung beim Bushveld Gerbil Tatera lencogaster (Wusiel aeaRodentia)unyZmbabwer nennen 25 1912 41 306 272 169 177, 88 94 D92 207 146 Pessöa, LEıLa; Dos Reıs, $. F.: Cranial infraspecifie differentiation in Proechimys iheringi Thomas (Rodentia: Echimyidae). — Intraspezifische Schädelvariabilität bei Proechimys ihe- m eoihomasi(Rodentıa-Echimyıdae) PEssÖA, LEıLA MARIA; Dos Reıs, $. F.: The contribution of cranıal indeterminate growth to non-geographic variation in adult Proechimys albispinus (Is. Geoffroy) (Rodentia: Echimyi- dae). -— Der Anteil des unbegrenzten Schädelwachstums an der Variabilität von Schädeln gleicher Herkunft bei adulten Proechimys albispinus (Is. Geoffroy) (Rodentia: Echimyidae) . POGLAYEN-NEUWALL, 1.: Notes on reproduction of captive Bassariscus sumicrasti (Procyoni- dae). — Bemerkungen zur Fortpflanzung von Bassarıscus sumichrasti (Procyonidae) in Gelangenschalt 2... ca We. ee POGLAYEN-NEUWALL, 1.; SHIVELY, J. N.: Testicular cycles of the Ringtail, Bassariscus astutus (Carnivora: Procyonidae). — Testicularzyklen des Katzenfretts, Bassariscus astutus (Carni- vora® Proeyonidae)”. nun a. en Pe A N u PRINSLOO, PAULETTE; ROBINSON, T. J.: Comparative cytogenetics of the Hyracoidea: chromo- somes of two Hyrax species from South Africa. - Vergleichende Cytogenetik der Hyraco- idea: Chromosomen von zwei Schlierer Arten aus>Südalrıka REır, URSULA; KLinGeL, H.: Hiding behaviour in wild Gerenuk (Zitocranius walleri) fawns. — Abliegeverhalten bert@erenuk- Kützent(Wtocranıny wall) a ROPER, T. J.; TAIT, A. I; CHRISTIAN, $. F.; FEE, D.: Excavation of three badger (Meles meles L.) setts_ Die Strukturidreier BauerdesDachses(Melesıneles) re SANTOS, T.; TELLERTA, J. L.: An experiment to test the consumption of arboreal food by Wood mouse Apodemus sylaticns. — Eın Experiment zur Ermittlung der Nahrungsaufnahme der Waldmaus Apodemusisylvanıens) au Baumenn a Snıpes, R. L.; KRIETE, A.: Quantitative investigation of the area and volume in different compartments of the intestine of 18 mammalıan species. - Quantitative Untersuchung an Gesamtoberfläche und Volumen verschiedener Darmabschnitte von 18 Säugetierarten ..... . SOMSOOR, S$.; STEINER, H. M.: Zur Größe des Aktionsraumes von Microtus arvalis (Pallas, 1779). — Home-range s1zelo 1 Mierotnstanoahs Wallasyle770) SOMSOOKR, $.; STEINER, H. M.: Fangmethoden und Geschlechterverhältnis in Stichproben von Feldmauspopulationen, Microtus arvalıs (Pallas, 1779). — Sex ratio of Microtus arvalıs (Ballas, 1779) samplesieaushtiniehtferenugy,pesto ker ps STORCH, G.; HABERSETZER, J.: Rückverlagerte Choanen und akzessorische Bulla tympanica bei rezenten Vermilingua und Eurotamandua aus dem Eozän von Messel (Mammalia: Xenar- thra). — Posteriorly displaced internal nares and supplementary auditory bulla in extant Vermilingua and Eurotamandua from the Eocene of Messel (Mammalıa: X enarthra) ee SUCHENTRUNK, F.; WILLING, R.; HARTL, G. B.: On eye lens weights and other age criteria of the Brown hare (Lepus europaeus Pallas, 1778). — Über Augenlinsengewichte und andere Altersmerkmale beim Feldhasen (Lepus europaeus Pallas, 1778) ................ TAYLoR, P. ]J.; CAMPBELL, G. K.; MEESTER, J. A. J.; van Dyk, DEBORAH: A study of allozyme evolution in African mongooses (Viverridae: Herpestinae). - Untersuchungen zur Evolution afrıkanischer Mangusten (Viverridae: Herpestinae) aufgrund von Allozymen ......... VAN WAEREBEEKR, K.; CANTO, ].; GONZALEZ, ].; OPORTO, J.; BRITO, J. L.: Southern Right wahle dolphins, Zissodelphis peronii, off the Pacific coast of South America. - Das Vorkommen des südlichen Glattdelphins Zissodelphis peronui vor der Pazifik-Küste von Südamerika... ... VrruLLo, A. D.; Cook, J. A.: The role of sperm morphology in the evolution of Tuco-Tucos, Ctenomys (Rodentia, Ctenomyidae): confirmation of results from Bolivian species. — Die Rolle von Spermienmorphologie in der Evolution der Kammratten, Ctenomys (Rodentia, Otenomyidae) . „3. «2 aber dt an ve a N e VOLLETH, MARIANNE; TIDEMANN, C. R.: The origin of the Australian Vespertilioninae bats, as indicated by chromosomal studies. -— Die phylogenetische Herkunft der australischen Vespertilionnaemachreytogenetischen Ersebn1Sse ng WIRTZ, P.; OLDEKOP, G.: Time budgets of Waterbuck (Kobus ellipsiprymnus) of different age, sex and social status. — Zeitbudgets bei Wasserböcken (Kobus elhpsiprymnus) von unter- schredlichem Alter @eschlechtund So zul statu se sr En Wissenschaftliche Kurzmitteilungen ANDERA, M.: First records of Microtus (Pitymys) felteni ın Albanien. — Erste Nachweise von Mhcrotusi (atyıny)S)Keltenun Ab anıene ANDREASSEN, H.-P.; BONDRUP-NIELSEN, $.: A comparison of the effects of a moss diet and a varıed diet on the growth of juvenile Wood lemmings, Myopus schisticolor (Lilljeb.). — Ein Vergleich über die Effekte von Moos- und Mischnahrung auf das Wachstum von jungen Waldlemmingen,Mpopusischistzeolon lLilljeb>) rer re re ee 34 219 195 722 3a 159 129 19 225 200 a) 27 365 135 284 339 324 48 380 378 ANsSORGE, H.: Diphyodont dentition of first premolar in the Red fox Vulpes vulpes (Linng, 1758). - Diphyodonte Dentition des ersten Prämolaren beim Rotfuchs Vulpes vulpes (Linng, TEE) ee nn a ee 188 ROOKMAAKRER, L. C.: The scientific name of the bontebok. — Der wissenschaftliche Name für dien Buncbock » 5 sinken Sa a 2 a ER 190 BARRETTE, CYRILLE: Fully formed first lower premolars ın a White-tailed deer (Odocoilens virginianus Zimmermann, 1780). — Vollständig ausgebildete erste untere Prämolaren bei einem Weißwedelhirsch (Odocozleus virginianus Zimmermann, 1780)... .. 2.22.22... 121 BAvER, K.: Ein Fund des pliozän/pleistozänen Marderhundes Nyctereutes megamastoides (Pomel, 1843) in Österreich. - A first find of the pliocene to villafranchian raccoon dog INgerenentesmesamastoidesBomel, 1843)... 0... 2... onen nn.ne 245 Espınosa, MARIA BEATRIZ; VITULLO, A. D.; MErAnNI, Marıa $.: Chromosomes of the Argentine Andean mouse, Akodon andınus (Cricetidae: Sigmodontinae). - Chromosomen der argentinischen Andenmaus Akodon andinus (Cricetidae: Sigmodontinae) ......... 124 Fıstanı, A.: First record of a fossil Cervus elaphus L., 1758, from Albania. - Erster Nachweis Emessossilenk@erunstelaphusıe, 1758 aus Albanien rn... 248 HEINRICH, D.: Some remarks on sıze differences of northwest German game populations from Neolithic to modern times. — Einige Bemerkungen über Größenunterschiede bei nordwest- deurschenWuldpopulauionenseitdemiNeolithikum . 2 In... .noeooneecen.n,. 59 JAEcK, KATHrın: Anomalies of the upper incisors in the genus Microtus (Cricetidae, Rodentia). — Anomalien der oberen Schneidezähne innerhalb der Gattung Microtus (Cricetidae, Ro- demme) ara 0 0 ee er NER 2 BROT a EEE 254 PALOMARES, F.: Use of an active badger sett by Egyptian mongooses, Herpestes ichneumon, in Southwest Spain. — Benutzung eines bewohnten Dachsbaus durch Ichneumone, Herpestes hm emo IOISUd est. Spanien in en ee 119 QUMSIYEH, M. B.: Karyotype of the East European hedgehog, Erinaceus concolor, from Jordan. — Der Karyotyp beim osteuropäischen Igel, Erinaceus concolor, aus Jordanien ........ 375 ROTHE, H.; KoENnIG, A.: Sex ratio in newborn Common marmosets (Calhthrix jacchus): no indication for a functional germ cell chimerism. — Geschlechterverhältnis bei neugeborenen Weißbüschelaffen (Callıthrıx jacchus): keine Anzeichen von funktionalem Keimzellenchi- MITSNUS #6 ee ee EN er DE ANGE 318 SAAVEDRA, BARBARA; SIMONETTI, J. A.: Archaeological evidence of Pudu pudu (Cervidae) in central Chile. - Archäologischer Beleg von Pudu pudu (Cervidae) in Zentralchile ..... ... 252 Bekanntmachungen SCHEN 5 u 0 2 0.8 a Be ee ee A EN 64, 126, 381 Buchbesprechungen SCHLECHTE a N a le Re 1271922565383 This journal is covered by Biosciences Information Service of Biological Abstracts, and by Current Con- tents (Series Agriculture, Biology, and Environmental Sciences) of Institute for Scientific Information Die in dieser Zeitschrift veröffentlichten Beiträge sind urheberrechtlich geschützt. Die dadurch begründeten Rechte, insbesondere die der Übersetzung, des Nachdrucks, des Vortrags, der Entnahme von Abbildungen und Tabellen, der Funk- und Fernsehsendung, der Mikroverfilmung oder der Vervielfältigung auf anderen Wegen, bleiben, auch bei nur auszugsweiser Verwertung, vorbehalten. Das Vervielfältigen dieser Zeitschrift ist auch ım Einzelfall grundsätzlich verboten. Die Herstellung einer Kopie eines einzelnen Beitrages oder von Teilen eines Beitrages ist auch im Einzelfall nur in den Grenzen der gesetzlichen Bestimmungen des Urheberrechtsgesetzes der Bundesrepublik Deutschland vom 9. September 1965 ın der Fassung vom 24. Juni 1985 zulässig. Sie ist grundsätzlich vergütungs- pflichtig. Zuwiderhandlungen unterliegen den Strafbestimmungen des Urheberrechtsgesetzes. Ge- setzlich zulässige Vervielfältigungen sind mit einem Vermerk über die Quelle und den Vervielfältiger zu kennzeichnen. © 1991 Paul Parey. Verlag: Paul Parey, Hamburg und Berlin. Anschriften: Spitalerstraße 12, W-2000 Hamburg 1; Seelbuschring 9-17, W-1000 Berlin 42, Bundesrepublik Deutschland. Printed in Germany by Westholsteinische Verlagsdruckerei Boyens & Co., Heide/Holstein ISSN 0044-3468 56 (1-6) 1-384 (1991) Be: R iltasttar See babe } nn laut bhn j Sa unkurk V a A a, f BR | | 01.56 (1), 1-64, Februar 1991 ISSN 0044-3468 C21274F ZEITSCHRIFT FÜR SAUGETIERKUNDE INTERNATIONAL JOURNAL OF MAMMALIAN BIOLOGY Irgan der Deutschen Gesellschaft für Säugetierkunde ur 3rünner, H.; Neet, C. R.: A parapatric scenery: The distribution and ecology of Sorex araneus and S. coronatus (Insectivora, Soricidae) in southwestern Germany. — Die parapatrische Szenerie: Die Verbreitung und Okologie von Sorex araneus and Sorex coronatus (Insectivora, Soricidae) in Südwestdeutschland 1 3randl, R.; Bezzel, E.; Reichholf, J.; Völkl, W.: Population dynamics of the Red squirrel in Bavaria. — Populations- dynamik des Eichhörnchens in Bayern 10 jantos, T.; Telleria, J. L.: An experiment to test the consumption of arboreal food by Wood mouse Apodemus sylvaticus. - Ein Experiment zur Ermittlung der Nahrungsaufnahme der Waldmaus (Apodemus sylvaticus) auf Bäumen 19 3ritton-Davidian, Janice; Vahdati, Mehrnouch; Benmehdi, Fatima; Gros, Paule; Nance, Valerie; Croset, H.; Guerassimov, S.; Triantaphyllidis, C.: Genetic differentiation in four species of Apodemus from Southern Europe: A. sylvaticus, A. flavicollis, A. agrarius and A. mystacinus (Muridae, Rodentia). —- Genetische Differenzierung bei vier Apodemus-Arten in Südeuropa: A. sylvaticus, A. flavicollis, A. agrariuss und A. mystacinus (Muridae, Rodentia) 25 >essöa, Leila, M.; Dos Reis, S. F.: Cranial infraspecific differentiation in Proechimys iheringi Thomas (Rodentia: Echimyidae). — Intraspezifische Schädelvariabilität bei Proechimys iheringi Thomas (Rodentia: Echimyidae) 34 zlison, G. T. H.; Skinner, J. D.: Thermoregulation and torpor in African woodland dormice, Graphiurus murinus, following cold acclimation. - Thermoregulation und Torpor bei afrikanischen Baumschläfern, Graphiurus muri- nus, nach Kältebelastung Wirtz, P.; Oldekop, G.: Time budgets of Waterbuck (Kobus ellipsiprymnus) of different age, sex and social status. — Zeitbudgets bei Wasserböcken (Kobus ellipsiprymnus) von unterschiedlichem Alter, Geschlecht und Sozialstatus 48 Wissenschaftliche Kurzmitteilungen Heinrich, D.: Some remarks on size differences of northwest German game populations from Neolithic to modern times. — Einige Bemerkungen über Größenunterschiede bei nordwestdeutschen Wildpopulationen seit dem Neolithikum 59 Bekanntmachung : ENNTHSONIAN JUN 27 1991 LIBRARIES Verlag Paul Parey Hamburg und Berlin HERAUSGEBERYJEDITORS P. J. H. van BREE, Amsterdam - W. FIEDLER, Wien — H. Frıck, München - W. HERRE, Kiel - H.-G. Krös, Berlin - H.-]. Kunn, Göttingen —- E. KuLzer, Tübingen - B. Lanza, Florenz - W. 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Säugetierkunde 56 (1991) 1-9 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 A parapatric scenery: the distribution and ecology of Sorex araneus and S. coronatus (Insectivora, Soricidae) in southwestern Germany By H. BrÜnNER and C. R. NEET Institut für Biologie I (Zoologie), Universität Freiburg, FRG, and Institut de Zoologie et d’Ecologie Animale, Universite de Lausanne, Switzerland Receipt of Ms. 7. 6. 1990 Acceptance of Ms. 31. 7. 1990 Abstract Studied the distribution of Sorex araneus and S. coronatus in southwestern Germany along a transect from the Upper Rhine Valley to the highest elevation, the Feldberg, in the Black Forest. 'The species were determined by polyacrylamide gel electrophoresis of total blood proteins. In the study area, the two species occurred in parapatry and showed an alternating altitude distributional pattern. S. araneus was found in the bottom of the Upper Rhine Valley and in the heights of the Feldberg, while S. coronatus was present in the lower and intermediate altitude levels of the Black Forest. In the Zastler Valley, S. coronatus advances 400 m farther uphill than in the St. Wilhelmer Valley where the climate is relatively cold and inclement. Two contact zones were found, situated at different altitudes according to the valley. In the study area, the species appeared to be altitudinal vicariads. The ecology and distributional history of S. araneus and $. coronatus are discussed in comparison with available data on their respective areas of distribution. It is argued that $. aranens occupies regions with rather continental climatic characters either on dry or wet soils. On the other hand, S$. coronatus prefers Atlantic climates and balanced soil humidity. The competitive interaction between S. coronatus and S. araneus and its biogeographical consequences are discussed. Introduction During the last 40 years, it gradually became obvious that in central and western Europe, the taxon Sorex araneus included a second, cryptic species: Sorex coronatus (e.g. HAUSSER et al. 1985; NEET 1989a; Hausser 1990; HaAusser et al. 1990). The two species are actually defined by their karyotypes (MEyLan and Hausser 1978) and have been shown to be clearcut species that do not hybridize (NEET and HAusser 1989). Since the first records of $. coronatus in Germany (OLERT 1973; SCHWAMMBERGER 1976), some information has been published on its distribution in the Rhineland and Westphalia by HAnDweErk (1987). However, only very few data are available for southern Germany (BRÜNNER and HoFFRICHTER 1987; Braun and Kıschnick 1987; KnocH 1989). In this paper we present new information on the distribution and ecology of S. araneus and $. coronatus in southwestern Germany (Freiburg region) and discuss their parapatric distribution in Europe, making special reference to the comparison of the distribution of the two species in southwestern Germany and western Switzerland. - Material and methods Shrews have been trapped in the dense, herbaceous vegetation found along small rivers or in similar vegetation along woodland paths. The trapping sites lie on a transect from the Kaiserstuhl in the Upper Rhine Valley to the Feldberg in the Black Forest. 100 live traps (Sherman, Tallahassee, USA) baited with a mixture of sardines in oil and rolled oats were exposed during 23 trapping dates in 19 trapping sites. A total amount of 87 shrews was captured and determined biochemically. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0001 $ 02.50/0 2 H. Brünner and C. R. Neet For electrophoretic determination, a blood sample of about 2 ul was taken from the tail. Blood samples were diluted ın a buffer solution, analysed according to a standard technique (HAussEr and ZUBER 1983) and ıdentified by the characteristic albumin patterns (NEET and Hausser 1989). As the electrophoretic technique has been tested in Switzerland only, karyological analyses were undertaken with some of the specimens of S. araneus and S. coronatus ın order to ascertain the reliability of the electrophoretic method in southwestern Germany (see NEET 1989a, for a discussion of the limitations of the applicability of the technique). Karyotypes were prepared in the laboratory from air dried mitotic chromosomes taken from bone marrow cells. The preparations were either Giemsa stained in a4 % solution (BAKER et al. 1982) or G- banded (SEABRIGHT 1971). All the S. araneus specimens analysed from southwestern Germany belonged to the “Vaud” karyotype with 22-24 (NF = 40) meta- and acrocentric autosomes, i.e. the same karyotype as the one used to test the biochemical determination techniques (NEET 1989a; NEET and Hausser 1989). The individuals of $S. coronatus all had 20 metacentric autosomes (NF = 44). Results In the study area, $. araneus and S. coronatus appear to be parapatrically distributed, ı.e. ın conjunct contact, without wıde zones of sympatry (Fig. 1). The corresponding lıst of trapping sites includes the altitudes at which the two species were caught (Table 1). Only one species, $. araneus, was found in the Upper Rhine Valley near Freiburg (Mooswald) and on the Kaiserstuhl. On the lower slopes of the Black Forest (near Ehrenkirchen) and of the Zartener Becken (near Freiburg), $. coronatus was the only species to be captured. In the Feldberg region, $. coronatus was the only species found up to 730 m in the Zastler Valley, and up to 700 m in the Bruggatal and in the St. Wilhelmer Valley. The upper limit of distribution of $. coronatus is at about 1050 m ın the Zastler Valley and 700 m in the St. Wilhelmer Valley. S. araneus ıs found between 900 m and 1450 m in the Zastler Valley and between 790 m and up to the top at the Feldberg (1496 m) in the St. Wilhelmer Valley. As one would expect according to thıs altitudinal segregation pattern where S. araneus ıs found in the lowlands and higher mountain altitudes, while $. coronatus ınhabits middle range altıtudes, two contact zones were found. In these zones, as in other contact zones studied ın Switzerland (NEET and Hausser 1990), the two species may coexist. The first contact zone is sıtuated in the middle of the Zastler Valley, at 900 m, and the second one on the Schönberg, a foothill of the Black forest, at 400 m. Discussion Distribution of S. araneus and S. coronatus in southwestern Europe The results presented here strongly suggest a parapatric Aislinion a che ma species ın southwestern Germany. This ıs consistent with the general pattern described ın central and western Europe by Hausser et al. (1985). The alternating altitude distribution found here also corresponds to the one found in western Switzerland (Hausser 1978) and confirms current views on the ecology of the two species. As a matter of fact, $. coronatus inhabits wide areas of northwestern Spain and most parts of France. In these countries, $. araneus is restricted to higher altıtudes (Pyrenees, Massıf central, Alpes), and the parapatric distribution ıs clearcut (HAusser et al. 1985). Moving towards the east and the north of Europe, the parapatrıcal distribution turns out to be progressively realized on a meso-distributional level and becomes more and more diffieult to visualize on a large scale. In Switzerland, S. coronatus is mainly present in the lowlands, the lower parts of the Jura and the Alps. In some places this species may, however, extend up to 1400 m of altitude. $. aranens ıs the dominant species above 800 m, but also occurs at low altitudes ın wet habitats such as the borders of the Lake ot Neuchätel. Although a simple histogram of their altıtudinal distributions suggests a wide The distribution and ecology of Sorex araneus and 5. coronatus 3 zone of sympatry, all detailed distributional studies have confirmed strict parapatry with contact zones never exceeding a few hundred meters (HAusser 1978; HAUSSER and Bourquin 1988; NEET 1989a; NEET and Hausser 1990). In Belgium, S. coronatus is the most common species in the western parts of the country. $. aranens appears in the eastern lowlands and the medium altıtudes ın the south of the country (Mys et al. 1985). In Holland, $. araneus occurs alone in the depressions in the north but the two species are believed to co-occur in the remainder of the country (LocH 1977). There ıs, however, some evidence that $. araneus is mainly distributed in areas below sea level, while $. coronatus occurs ın areas above this level (Hausser pers. comm.). In northeastern Germany the two species co-occur over a large area: Niederrheinische Tiefebene, Mittel- rhein, westfälische Bucht, Rheinisches Schiefergebirge (HUTTERER and VIERHAUS 1984; HANDWERK 1987). One can sum up at this point by stating that $. coronatus ıs found ın areas with a balanced Atlantic climate, while $. araneus occurs in colder or wetter habitats, and that the parapatric distribution ıs decreasingly distinct as one proceeds towards the north of Europe. Distributional ecology of S. araneus and S. coronatus in southwestern Germany In southern Germany three climatic zones have been differentiated (LIEHL and Sıck 1984): 1. the warm, dry and continental Upper Rhine Valley; 2. the humid lower-altitude foothills of the Black Forest (up to 600 m), with an Atlantic climate; and 3. the heights of the Black Forest (up to 1500 m) with a typically subalpıne climate and abundant rainfall. These zones closely correspond to the altitudinal distribution zones of the two species S$. araneus and S. coronatus. The fırst species ıs found in the Upper Rhine Valley and above 790 m in the Black Forest, while the second ıs mainly found at low altitudes of the Black Forest. Interestingly, a correspondance between altıtudınal zones of vegetation and the distribution of the two species has also been demonstrated in western Switzerland (NEET 1989a). The glacier-formed St. Wilhelmer Valley ıs wide with high nocturnal eradiation. It opens with a gradual slope into the Bruggatal almost right angles at a very narrow point (Fig. 1). Thus, the cold air is often congested providing a rough clımate. Moreover, the valley is on a lee position with rather low rainfall. In contrast, the Zastler Valley ıs narrow and steep sloping with several steps. It describes a slıght bow and opens widely into the Bruggatal (Fig. 1). Thus, the cold aır from the Feldberg heights can discharge quickly without any obstacle. As it ıs in a windward position, higher rainfall is noted (SCHWABE- KRATOCHWIL and BOGENRIEDER, pers. comm.). In the St. Wilhelmer Valley, S. coronatus was trapped up to an altıtude of 700 m, in contrast to the clımatıcally favoured Zastler Valley, where ıt can be found up to 1050 m. An interesting parallelism with this observation has been reported for a contact zone of the Sorex species in a valley of the Valaısan Alps between Val d’Illiez and Champery (MEyLAn 1964; Orr 1968). There, $. coronatus occurred on the southeastern slopes of the valley about 1 km farther uphill than on the other sıde of the river at the bottom of the northwestern slopes. It can be supposed that the banks along the southeastern slopes are also climatıcally favoured, thus enabling S. coronatus to reach higher into the Valley. Other sımilar examples have been found for the distribution of the two species in Switzerland (Hausser and BourQuin 1988). Another distributional pattern, similar to the one found in our study area, can be expected to be found for the two Sorex species in northern Baden, where S. araneus was trapped ın the Hardt Forest near Karlsruhe (BRÜNNER unpub.) and in several locations in the south of Rastatt (NIETHAMMER pers. comm.). $. coronatus was found in a beech forest at 300 m on the western slopes of the Black Forest. In this area, 26 individuals of S. 4 H. Brünner and C. R. Neet 0 1 2km Fig. 1. Trapping sites and distribution of $S. araneus (black circles) and $. coronatus (white circles) in the study area. The contact zone is given by a black and white circle. A: Freiburg region, in southwestern Germany, B: Feldberg region in the Black Forest. Locality numbers correspond to Table 1 coronatus and 3 of S. araneus were captured (Braun and KıscHnIck 1987). The shrews might have been taken from a contact zone but, at present, no additional data are available for this region. Whereas the distribution of the Sorex species ın the Black Forest corresponds to the situation throughout Europe described above, the occurrence of S. araneus in the Upper Rhine Valley, where it has already been found ın Alsace by Hausser (1978), and the sımultaneous absence of $. coronatus reveal a possible significance of drought and heat factors of the continental climate for the distributional ecology of the two species. An additional indication comes from the Catalan parts of the Pyrenees where the climate ıs dry, warm and Mediterranean-like and where $. araneus descends to low altitudes, while S. The distribution and ecology of Sorex araneus and $. coronatus 5 coronatus ıs only found at 800-1500 m, although it is wıdespread ın the Atlantic climate of northwestern Spain (LOPEZ-FUSTER et al. 1985). Ecological comparison of the two species In contrast to $. coronatus having a relatively limited distribution in western Europe, S$. araneus has a large distributional range from the Pyrenees to Lake Baikal in Sıberia with its distributional centre in the continental east. As a species, S. aranens reveals greater ecological valency (Hausser and BourQUIn 1988). This point is supported by some aspects of the evolutionary biogeography of the two shrew species. The British mainland, for example, with typical Atlantic climate, is only inhabited by $. araneus since $. coronatus did not arrıve before its separation from the continent after the last glaciations. This species also did not succeed ın passing the Alps to northern Italy. In these two regions S. araneus occupies all suitable habitats in the plains and in the mountain areas. As shown by Hausser and Bourquin (1988), in Switzerland S. araneus would be able to exist ın most biotopes of the country. Its absence ın the lowlands can more convincingly be explained by competition pressure going out from $. coronatus than by autoecological constraints (NEET and HAusser 1990). It should, however, be pointed out that $. araneus is not a simple species, but consists of several chromosomal types (e.g. SEARLE 1984; Hausser et al. 1985; HAusseEr et al. 1986; REUMER and MEyLAN 1986). To what extent are these races differentiated from an ecological point of view is not yet known and, obviously, this point may be of crucıal importance in explaining the apparently wide ecological valency of S. araneus. The principle stating that ecologically ıdentical species cannot coexist (GAUSE 1934) is now widely accepted (BEGon et al. 1986). As a consequence of the general similarıty of S. araneus and S. coronatus, competition is an important factor ın the lack of overlap of their distributional ranges (HAusser 1978; NEET and Hausser 1990). An ecological field analysıs ın Switzerland revealed that in two contact zones, the two species coexisted over a Table 1. List of trapping sites in southwestern Germany, classified by their altitudes Altitude Species n (ind.) Locality ( m a.s.l.) S. araneus/ S. coronatus 1/0 1/0 0/1 2/0 3/3 2/0 0/1 1450 1230 1050 900 900 790 730 700 600 540 420 400 390 360 350 300 270 Feldberg, near top Feldberg, Zastler Hut Upper Zastler Valley Upper St. Wilhelmer Valley Middle Zastler Valley Upper St. Wilhelmer Valley Lower Zastler Valley 0/7 Middle St. Wilhelmer Valley 0/1 Lower Zastler Valley 0/5 Bruggatal, near Oberried 0/2 Zartener Becken, near Oberried 1/1 Schönberg, near Ebringen 0/1 Zartener Becken, near Kirchzarten 0/2 Zartener Becken, near Freiburg 0/1 Schönberg, near Ebringen 0/3 Border of Black Forest, near Ehrenkirchen 3/0 Kaiserstuhl, Liliental 230 4/0 Mooswald, northwest of Freiburg 21115 42/0 Mooswald, southwest of Freiburg Black circles = $S. araneus, white circles = $. coronatus, black and white circle = contact zone. Locality numbers correspond to Fig. 1 DOoONONVURUDND- ® ® ®) © Ö © ©) ®) ©) ©) © Ö ©) ©) © ©) ® ® ® 6 H. Brünner and C. R. Neet limited zone of overlap by selecting different microhabitats, those of $. araneus having a thicker litter layer and higher soil humidity (NEET and Hausser 1990). It was also shown that $. araneus and S. coronatus occupy the same trophic niche in their zones of contact (NEET 1989a) and that equal intensities of ıntra- and interspecific competition for food as a limiting factor lead to interspecific territoriality (NEET 1989b). In other words, the two species do differ in terms of their ecoclimatic adaptations but are still too similar to coexist. This example thus illustrates the usefulness of the principle of limiting similarıty proposed by Mac ARTHUR and Levins (1967). If the high ecological similarity of $. araneus and S. coronatus does not allow sympatrical distribution, and although the contact zones we observe today seem to be relatively stabilized, one may suppose that, on an evolutionary time scale, one species will dominate the other and, with climatic changes, will replace it over an appreciable geographical space. This is how the distributional ecology of S. araneus and S. coronatus has been interpreted, since the actual distribution of the two species bears several indications that S. coronatus has forced $. araneus up to the north and east of Europe since the last glaciations. Moreover, the limited adaptations of both species to regionally or locally different climates in connection with mutual exclusion are considered to be characteristics of an early stage of ecological differentiation (HAUSSER 1984; HAussERr et al. 1985). The results presented here confirm the parapatric interpretation (e.g. HAussEr et al. 1985; Hausser and BoURQUIN 1988) especially since there is a clear similarity between the situation in southwestern Germany and that in western Switzerland (Table 2). In central and northern Germany, the situation is less clear and the distribution is more mosaic-like. In that area, the distribution of the two species is sometimes considered as sympatrical (HUTTERER pers. comm.). HANDWERK (1987) reported ratios of $. araneus: S. coronatus of 1:1t0 1:3 for the plains of the Niederrhein and the Cologne-Bonn region, and 2:1 ratios Table 2. Comparison of the distributional ecology of S. araneus and S. coronatus in southwestern Germany and western Switzerland Data for western Switzerland are taken from NEET 1989a Southwestern Germany Western Switzerland Altıtude range S. araneus 215-1450 m 380-1950 m S. coronatus 300-1050 m 490-1340 m Type of distribution parapatric parapatric Relative habitat preferences S. araneus cold (continental) wet and cold S. coronatus warm (Atlantic) dry and warm balanced humidity Breadth of contact zones 200-2000 m? 100-1000 m® Ecological relationship in the contact zones Habitat selection in response to interspecific competition Interspecific territorıality “The breadth of the contact zones was estimated as follows: in the Zastler Valley, the breadth was estimated to be equivalent to the length of the trapping area, ı. e. 200 m; in the St. Wilhelmer Valley the maximal breadth was estimated to be 01 2000 m, which corresponds to the distance between capture points 6 and 8 (Fig. 1). - ° Nerr (1989) indicates breadths around 100 m. However, ın patchy areas (woodlands alternating with grasslands) it is difficult to estimate a precise value (see NEET and HAusser 1990). However, maximal values around 1000 m have been estimated. The distribution and ecology of Sorex araneus and S. coronatus 7 for the adjacent regions of medium altıtude. NEET and Hausser (1990) consider that the work of Hanpwerk (1987) lacks sufficient detail to ascertain whether the two species are sympatrically or parapatrically distributed in that area. A detailed ecological study might reveal local differentiation, which may be very subtle, as in the Swiss contact zones. Nevertheless, an alternative hypotheses exists and is due to the simple fact that resource availability influences interspecific competition. In the contact zones studied by NEET and Hausser (1990), there is evidence that food was a limiting factor (NEET 1989a) and that competitive pressure induced interspecific territoriality (NEET 1989b). If there are no limiting ressources, the situation may change. An example is given in the work by LLEwELLYN and JENKINS (1987) on the cricetids Peromyscus maniculatus and P. trueı, where it was shown that there are seasonal changes in overlap of microhabitat niche, and that these changes may be explained by the degree of resource scarcity. In one of the contact zones studied in Switzerland, it was shown that the competitive pressure between S. araneus and S. coronatus ıs present throughout the year (NEET and Hausser 1990). However, the situation may be different in central and northern Germany. A seasonal variation in competitive pressures is a realistic hypothesis and may result in varıations of the degree of overlap between the two species and limited sympatry. Acknowledgements One of us (H. B.) gratefully received important impulse for this work from Prof. G. ÖscHE, University of Freiburg. We both thank Prof. P. VoGEL and Prof. J. HAaussEr from the University of Lausanne for their support, interest and help in this research. The electrophoretic determinations were started in the Institute of Zoology and Animal Ecology of the University of Lausanne and later performed in the laboratory of Prof. H. GUTZEIT, University of Freiburg, with friendly support of V. Grau. Traps were placed at disposal by M. Brau, Staatliches Museum für Naturkunde, Karlsruhe. The work was supported by Prof. FRIEDRICH KIEFER-FonDs, Badischer Landesverein für Natur- kunde und Naturschutz e.V. Many thanks are due to H. Brack for her invaluable help in the field. Zusammenfassung Eine parapatrische Szenerie: Die Verbreitung und Ökologie von Sorex araneus und Sorex coronatus (Insectivora, Soricidae) in Südwestdeutschland Die Verbreitung von Sorex aranenus und Sorex coronatus ın Südwestdeutschland wurde in einem Transekt vom Oberrheingraben zum Feldberg ım Schwarzwald untersucht. Die Artbestimmung geschah mittels einer Polyacrylamıid-Gelelektrophorese des Gesamtbluteiweißes. Im Untersuchungs- raum waren die beiden Arten parapatrisch verbreitet, und dies in Form einer alternierenden Höhen- verbreitung. $. araneus wurde im Oberrheintal und am oberen Feldberg angetroffen, $. coronatus in den unteren und mittleren Schwarzwaldlagen. Im klimatisch begünstigten Zastler Tal reicht die Verbreitung von S. coronatus 400 Höhenmeter weiter talaufwärts als im kälteren und rauheren St. Wilhelmer Tal. Zwei Kontaktzonen, eine obere und eine untere, wurden ausfindig gemacht. Im Untersuchungsgebiet können die beiden Arten als höhenvikariierende Arten bezeichnet werden. Die Ökologie und Verbreitungsgeschichte von $. araneus und S. coronatus wird anhand der bisher bekannt gewordenen Daten über ihre Lebensräume diskutiert. $. araneus besiedelt Gebiete mit eher kontinen- talem Charakter sowie Lebensräume mit nassen und trockenen Böden. Dagegen bevorzugt S. coronatus atlantisches Klima und ausgeglichene Bodenfeuchtigkeit. References BAKER, R. J.; Haıpuk, M. W.; RoBBıns, L. W.; CADENA, A.; Koop, B. F. (1982): Chromosomal studies of South American bats and their systematic implications. Special publ. Pymatuning Lab. Ecol. 6, 303-327. BEcon, M.; HARPER, J. L.; TownsenD, C. R. (1986): Ecology. Individuals, Populations and Communities. Oxford: Blackwell. BRAUN, M.; Kischnick, P. (1987): Spitzmäuse und ihre Nahrung in einem Buchenwald. Carolinea 45, 159-160, 8 H. Brünner and C. R. Neet BRÜNNER, H.; HOFFRICHTER, ©. (1987): Neue Fundorte der Alpenspitzmaus (Sorex alpinus Schinz, 1837) im Südschwarzwald. Mitt. bad. Landesver. Naturkunde u. Naturschutz N.F. 14, 403-408. Gause, G. F. (1934): The struggle for existence. Baltimore: Wıiliams and Wilkins Co.: (1934). New York: Hafner Publ. (Repr. 1964). 3 HANDWERK, J. (1987): Neue Daten zur Morphologie, Verbreitung und Okologie der Spitzmäuse Sorex araneus und $. coronatus ım Rheinland. Bonn. zool. Beitr. 38, 273-297. Hausser, J. (1978): Repartition en Suisse et en France de Sorex araneus L., 1758 et de Sorex coronatus Millet, 1828 (Mammalia, Insectivora). Mammalıa 42, 329-341. — (1984): Genetic drift and selection: Their respective weights in the morphological and genetic differentiation of four species of shrews in Southern Europe (Insectivora, Soricidae). Z. zool. Syst. Evolut.-forsch. 22, 302-320. — (1990): Sorex coronatus Millet, 1828. In: Handbuch der Säugetiere Europas. Ed. by J. NIETHAM- MER and F. Krapp. Wiesbaden: Aula-Verlag. Hausser, J.; Bouraumn, J.-D. (1988): Repartition de douze especes de mammiferes en Suisse. Atlas des Mammiferes de Suisse. Societe suisse pour l’Etude de la faune sauvage, Lausanne. HaAussER, J.; CATZEFLIS, F.; MEyLAN, A.; VOGEL, P. (1985): Speciation in the Sorex araneus complex (Mammalia: Insectivora). Acta Zool. Fennica 170, 125-130. HaussER, J.; DANNELID, E.; CATZEFLIS, F. (1986): Distribution of two karyotypic races of Sorex araneus (Insectivora, Soricidae) in Switzerland and the post-glacıial recolonization of the Valais: First results. Z. zool. Syst. Evolut.-forsch. 24, 307-314. HaAUuSSER, J.; HUTTERER, R.; VOGEL, P. (1990): Sorex araneus L. 1758. In: Handbuch der Säugetiere Europas. Ed. by J. NIETHAMMER and F. Krapp. Wiesbaden: Aula-Verlag. HaussER, J.; ZUBER, N. (1983): Determination specifique d’individus vıvants des deux especes jumelles Sorex araneus et $. coronatus, par deux techniques biochimiques (Insectivora, Soricidae). Rev. suisse Zool. 90, 857-862. HUTTERER, R.; VIERHAUS, H. (1984): Waldspitzmaus, Schabrackenspitzmaus. In: Die Säugetiere Westfalens. Ed. by R. SCHRÖPFER; R. FELDMANN and H. VIERHAUS. Münster: Abh. Westf. Mus. Naturkunde 4, 54-60. Knoch, D. (1989): Säugetiere im Belchengebiet. In: Der Belchen im Schwarzwald. Die Natur- und Landschaftsschutzgebiete Baden-Württembergs 13. Landesanstalt für Umweltschutz, Baden- Württemberg, 1159-1165. LiEHL, E.; Sick, W. D. (1984): Der Schwarzwald - Beiträge zur Landeskunde. 3 ed. Ver. Allemann. Inst. Freiburg ı. Br. 47. Bühl, Baden: Konkordıa. LLEWELLyYN, J. B.; JEnKIns, $. H. (1987): Patterns of niche shift in mice: Seasonal changes in microhabitat breadth and overlap. Am. Nat. 129, 365-381. LocH, R. (1977). A biometrical study of karyotypes A and B of Sorex araneus Linnaeus, 1758, in the Netherlands (Mammalıa, Insectivora). Lutra 19, 21-36. LOPEZ-FUSTER, M. J.; GOSALBEZ, ]J.; Sans-CoMA, V. (1985): Presencia y distribucion de Sorex coronatus Millet, 1828 (Insectivora, Mammalıa) en el NE iberico. P. Dept. Zool. Barcelona 11, 93-97. Mac ARTHUR, R. H.; Levins, R. (1967): The limiting sımilarity, convergence and divergence of coexisting species. Am. Nat. 101, 377-385. MeEyLan, A. (1964): Le polymorphisme chromosomique de Sorex araneus L. (Mammalia, Insectiv- ora). Rev. suisse Zool. 71, 903-983. MEyLAn, A.; HaussEr, J. (1978): Le type chromosomique A des Sorex du groupe araneus: Sorex coronatus Millet, 1828 (Mammalıa, Insectivora). Mammalıa 42, 115-122. Mys, B.; VAN DER STRAETEN, E.; VERHEYEN, W. (1985): The biometrical and morphological ıdentification and the distribution of Sorex araneus L., 1758 and $. coronatus Millet, 1828 in Belgium (Insectivora, Soricıdae). Lutra 28, 55-70. NEET, C. R. (1989a): Ecologie comparee et biog&ographie Eevolutive de deux especes parapatriques: Sorex araneus et S. coronatus (Mammalıa, Insectivora, Soricidae). Ph. D. thesis, University of Lausanne. — (1989b): Evaluation de la territorialite interspecifique entre Sorex araneus et $S. coronatus dans une zone de syntopie (Insectivora, Soricidae). Mammalıa 53, 329-335. NEET, C. R.; Hausser, J. (1989): Chromosomal rearrangements, speciation and reproductive isolation: the example of two karyotypic species of the genus Sorex. J. evol. Biol. 2, 373-378. — — (1990): Habitat selection in zones of parapatric contact between the common shrew Sorex araneus and Millet’s shrew S. coronatus. J. Anım. Ecol. 59, 235-250. OLERT, J. (1973): Cytologisch-morphologische Untersuchungen an der Waldspitzmaus (Sorex araneus Linne, 1758) und der Schabrackenspitzmaus (Sorex gemellus Ott, 1968. (Mammalıa — Insectivora). Veröffentl. Univ. Innsbruck 76, 1-73. OTT, J. (1968): Nachweis natürlicher reproduktiver Isolation zwischen Sorex gemellus sp. n. und Sorex araneus Linnaeus, 1758 ın der Schweiz (Mammalıa, Insectivora). Rev. suisse Zool. 75, 53-75. The distribution and ecology of Sorex araneus and S. coronatus 9 REUMER, J. W. F.; MEyLan, A. (1986): New developments in vertebrate cytotaxonomy IX: Chromo- some numbers in the order Insectivora (Mammalia). Genetica 70, 119-151. SCHWAMMBERGER, K. H. (1976): Nachweis der Schabrackenspitzmaus (Sorex gemellus Ott, 1968) in Westfalen. Natur u. Heimat 36, 66-69. SEABRIGHT, M. (1971): A rapid banding technique for human chromosomes. Lancet 2, 971-972. SEARLE, J. B. (1984): Three new karyotypic races of the common shrew Sorex araneus (Mammalıa: Insectivora) and a phylogeny. Syst. Zool. 33, 184-194. Authors’ addresses: HARALD BRÜNNER, Institut für Biologie I (Zoologie), Albertstraße 21a, Albert- Ludwigs-Universität, W-7800 Freiburg/Br., FRG, and Dr CorneLis R. NEET, Conservation de la faune, Chemin du Marquisat 1, CH-1025 St. Sulpice, Switzer- land Z. Säugetierkunde 56 (1991) 10-18 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Population dynamics of the Red squirrel in Bavaria By R. BranDı, E. BEZZEL, J. REICHHOLF and W. VÖöLkL Lehrstuhl für Tierökologie I, Universität Bayreuth, FRG, Institut für Vogelkunde, Garmisch-Partenkirchen, FRG and Zoologische Staatssammlung München, FRG Receipt of Ms. 23. 5. 1990 Acceptance of Ms. 30. 7. 1990 Abstract Describes the population dynamics of the red squirrel (Sciurus vulgaris) in Bavaria since the early 1970s. A decline ın the population of squirrels was found between 1970 and 1980, while recently an increase was recorded. The decline can be attributed to reproductive waves caused by mast years of coniferous trees. The recent increase can be explained by general forest decline, since damage to trees desynchronizes seed production and squirrels are provided with a constant food resource over a number of years. Weather conditions were found to be unimportant for the population dynamics of squirrels. For the beech marten (Martes foina), however, a negative correlation could be demonstrated between number of squirrels and this predator at one study site. Introduction Compared to birds the available data on long-term fluctuations of medium-sized mammals are very rare. This is due to the difficulties to trap and count these animals within a reasonable period of time. Even for diurnal species like the red squirrel (Sciurus vulgaris) only a limited amount of information ıs available. Long-term series are badly needed to document the full range of patterns in the population dynamics of mammals. In game biology there ıs a long tradition to use hunting bags or other indirect measures to document fluctuations in game animals (for a recent example see PoTTs et al. 1984). We followed this line of investigations and present indirect density indices to describe the dynamics of the red squirrel in Bavaria. We concentrate on the following questions: 1. Are there any long-term trends in the population of squirrels in Bavaria? In two reports we suggested that there was a decline in numbers of squirrels since the early 1970s (BrAanDL 1983; REICHHOLF 1983). 2. Is the dynamics of the squirrel populations similar in different regions of Bavaria? 3. The main food of the red squirrel is the seeds of coniferous trees (WILTAFSKY 1978). Many studies have shown that the populations of squirrels follow seed production (Formosov 1933; PULLIAINEN 1984; REICHHOLF 1974). Is this pattern also evident within our sets of data? 4. The red squirrel is the prey of several predators such as the pine and beech marten (Martes martes, M. foina; STUBBE 1988). Do these species have some influence on the populations of the red squirrel? Material and methods In the following we name each data set by its geographical location (see Fig. 1): 1. Pressath: This data set consists of specimens delivered to a taxidermist between September and March from 1965/66 onwards to 1982/83 with a gap between 1967/68 and 1972/73 (550 individuals; Branpı 1983). The working period of the taxidermist was not constant over the years. We used the U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0010 $ 02.50/0 Population dynamics of the Red squirrel in Bavarıa 11 U] Bad Berneck Pressath 8 Nürnberg München Garmisch - Partenkirchen Fig. 1. Geographical location of the investigated areas in Bavaria number of mammals and birds to standardize red squirrel data and calculated percentages of red squirrels delivered to the taxidermist. Around Pressath the dominating types of vegetation are spruce forests (Picea abies) and at poorer sites pine forests (Pinus sylvestris). Altitudes range from about 400 to 600 m NN. 2. Bad Berneck: Similar to Pressath these data are specimens delivered to a taxidermist between 1970 and 1988 (922 individuals). This taxidermist worked over the entire year. We standardized the data because the overall working effort appeared to vary from year to year. The number of delivered birds was used as an independent measure of the working effort. Numbers of red squirrels are expressed in individuals delivered per 100 birds. Bad Berneck is situated within the Fichtelgebirge. Spruce forests are the dominating type of vegetation, and the altitudes range from 500 to 1000 m. At higher altitudes winters may be quite severe. For this area we have also data about weather, seed production of the spruce and dynamics of one potential predator. a. Monthly information of rainfall and temperature was available for Bayreuth, only 10 km from Bad Berneck. b. The forest authorities estimate the seed production in four categories: “Vollmast” = very high 12 R. Brandl, E. Bezzel, J. Reichholf and W. Völkl seed production, “Halbmast” = good seed production, “Sprengmast” = low seed production and no seed production. We ranked each year on this scale from O0 to 3 and allowed for intermediate values according to information of the forest authorities (available years 1971 to 1988). c. The number of martens delivered to the taxidermist (pine and beech marten) may be used as an indicator of the predator density. The pine marten is rather uncommon in the area around Bad Berneck, so we concentrate on the beech marten. Data were standardized similar to those for squirrel numbers. 3. Inn: These data are from squirrel sightings made during standard excursions between 1971 and 1982 (153 records; REICHHOLF 1983). 4. B12: Road kills are often good indicators of the population dynamics of medium-sized mammals. The B 12 data sets are the sums of road kills during a year along the federal highway B 12 between Munich and Bad Füssing (89 individuals; REICHHOLF 1983; 1976-1988). The killed martens were also counted along the same route. 5. Garmisch-Partenkirchen: Data are from squirrel sightings along three transects (1979 to 1988; 195 records), situated at the lower mountain forests about 800 to 900 m NN and were sampled twice each month. The dominating tree species is the spruce, but deciduous species of trees are also present. Winters may be severe with snow cover between November to April. Results Phenology The seasonal distribution of data (Inn, B 12, Garmisch-Partenkirchen) is plotted in Fig. 2. The phenological patterns are sımilar for Inn and Garmisch-Partenkirchen (r = 0.55; P< 0.05; one-tailed). No road kills were found during January and December ın contrast to the data based on squirrel sightings. All three plots showed a mınımum during July. 10 JR TMTAT MINEN TEAT SSTZOZENZED Fig. 2. Seasonal distribution of road kills (B12) and sighting records (Inn, Garm. = Garmisch- Partenkirchen) of the red squirrel, given as the sum of sightings or road kills recorded within each month Population dynamics of the Red squirrel in Bavarıa 13 40 Garm. 30 Inn N 20 10 B12 1964 1196825197.21197.6 21980771984 11988 Fig. 3. Population fluctuations of the red squirrel in 60 different areas of Bavarıa. For the areas Inn, B 12 and Garmisch-Partenkirchen (Garm.) the sum of all sightings or road kills within each year ıs given. The data from Bad Berneck and Pressath are relative % indices calculated as percentages of squirrels deli- vered per all anımals (Pressath) or squirrels delivered per 100 birds (Bad Berneck; see Material and methods) Bad Berneck Pressath 0 1964 1968 1972 1976 1980 1984 1988 YEAR Population dynamics In Fig. 3 all data sets are plotted across the years studied. The data from northern Bavaria (Bad Berneck and Pressath) show a clear decline in the squirrel populations. This decline ıs also apparent within the Inn data, but with an obvious peak in 1974. Since 1980 there seems to be a slight increase of squirrel populations, evident ın all data sets covering this period. Furthermore, the plots show a rough concordance in minor peaks (e.g. 1975, 1978, 1983, 1985). Table 1 presents the correlation coefficients between sampling areas: all correlation coefficients are positive and six from ten are significant. Table 1. Matrix of pairwise correlation coefficients (upper half of the matrix) between the squirrel density indices of Fig. 3 Each coefficient measures the sımilarity in the population dynamics of squirrels between a particular pair of geographical locations, considering only the years with data for both areas. The lower half of the matrix gives the significance levels of the correlation coefficients (one-tailed) 1 Inn 215112 3 Garmisch-P. 4 Bad Berneck 5 Pressath Squirrels and predators We concentrated on the data from Bad Berneck, because this is the longest time series. In Fig. 4A we removed the negative trend apparent in Fig. 3 by linear regression. Runs with several transformations produced identical results. Firstly, some very prominent peaks are evident by the resıduals, and secondly, there is a depression in squirrel densities between 1976 and 1980. This depression within the squirrel data seems to correlate with a maxımum in the population index of the beech marten (Fig. 4B). A stepwise multiple regression was 14 R. Brandl, E. Bezzel, J. Reichholf and W. Völkl performed with the squirrel data as dependent variable and years and predator density index as independent variables. The sequence of varıables entering the regression was 1. year and 2. predator with a significant negative regression coefficient (see also Table 2). Fig. 5A plots the residuals from a simple regression of squirrels to years (data in Fig. 4A) against the marten index (Fig. 4B): the negative correlation between squirrels and the beech marten is evident. 20 10 -10 0 1968 1972 1976 1980 1984 1988 40 B 30 20 10 1968 1972 1976 1980 1984 1988 Fig. 4. A: Residuals of the red squirrel density after 15 C removing the negative trend over the years investi- gated. B: Dynamics of the beech marten in the same > area, given as number of martens delivered per 100 birds (see Material and methods). C: Residuals of the ” red squirrel density after removing the trend over the 2 years examined and the influence of the marten (all N] a . 1968 1972 1976 1980 1984 1988 data from Bad Berneck, Fichtelgebirge) YEAR Residuals Bad Berneck Fig. 5. A: Scatter plot of the red squirrel (after removing the negative trend over the years ex- amined) and the populations of the beech marten (see Fıg. 4 A, B; area: Bad Berneck). B: Scatter 10 plot of road kills of squirrels and beech marten Martes foina along the highway B 12 Population dynamics of the Red squirrel in Bavaria 15 Table 2. Stepwise multiple regression with red squirrel index from Bad Berneck as dependent variable and year, marten population and mast index of the spruce in the previous year as independent variables A: sequence of varıables entering the regression Step Variable 1 year 2 marten 3 mast ındex B: regression equation after step 3 Variable Slope Stand.slope Year -1.532 Marten -0.533 Mast index al Constant 3065 ! note that P is two-tailed in Table 2; in the text we used a one-tailed probability. In Fig. 5B a scattergram was also plotted for the road kills of red squirrel and beech marten along the B12. Contrary to the area around Bad Berneck we found no negative correlation between predator and prey (r = 0.37; P > 0.2; two-tailed). Squirrels, food and weather For the data set from Bad Berneck information on seed yield of spruce was available (Fig. 6B). The residuals from the multiple regression after step two ın Table 2 are also shown. Seed index and residuals are significantly correlated with a time lag of one year (r = 0.48; p = 0.04; one-tailed; note that degrees of freedom have to account for the number of varıables used to calculate the residuals). Table 2 shows the stepwise multiple regression of squirrel population index and the independent varıables 1) year, 2) marten population and 3) mast index (one year time lag). The mast index entered the regression during the last step. The influence of monthly rainfall and temperature patterns on the residual (Fig. 4C) for time lags for up to two years were also tested. From 60 calculated correlation coefficients 20 A Residuals © -20 1IES TITTEN ITZEFANISEOTENIEATE 11988 6) Fıg. 6. A: Dynamics of the red squirrel after remov- ing the influence of years and beech marten. B: Fluctuations in seed production of the spruce (area ö : (0) Bad Berneck, Fichtelgebirge) 1968 1972 1976 1980 1984 1988 Seed Production 16 R. Brandl, E. Bezzel, ]J. Reichholf and W. Völkl only two were significant at the 5 % level, a result expected by chance alone. Therefore, it seems rather safe to conclude that weather conditions are unimportant for the dynamics of red squirrels. Discussion The seasonal distribution of the observational data and the road kills of red squirrels are quite different. During winter the red squirrel shows a reduced overall activity (ZWAHLEN 1975; Tonkın 1983). The reduced number of road kills are in accord with this fact. But why are so many sightings recorded during winter? The squirrel is a diurnal species with a long bimodal activity during summer, and a short unımodal pattern in winter (TONnkINn 1983). In winter the peak activity is during late morning. This activity pattern may correlate with the activity of observers. Squirrels search for food during winter in the vicinity of the dreys without making longer excursions to save energy (PULLIAINEN 1973). That is why during transect counts squirrels are easily detected by an observer. A reduced foraging range may decrease the probability of beeing killed on the road. Fig. 3 suggests a decline in the squirrel population in nearly all regions of Bavaria. Is this decline a natural phenomenon within the population dynamics of the squirrel, or is this decline the result of the environmental damage caused by man? From studies in boreal regions it is well known that squirrels increase their numbers after years of a high seed yield (Formosov 1933; PULLIAINEN 1984). At first glance, the correlation between seed production and squirrel numbers seems to be weak in Bavaria (Table 2), but we believe that this impression is incorrect. For example MÖckEL (1987) described a clear increase ın the red squirrel after a mast year ın the West-Erzgebirge and a decline to the original density within 5 years. The availability of food after a mast year seems to improve the condition of females and thereby the probability of reproduction (WAUTERS and DHONnDT 1989; GURNELL 1983) also under normal ambient conditions ın Central Europe. The increased probability of reproduction, an increased litter size as well as better survival of young produces a population explosion in squirrels. The maximum age of red squirrels is more than 10 years, and about 1 % of individuals may reach 5 years of age or older (WILTAFskY 1978). Therefore, the explosion needs around 5 years to fade, as long as adult survival is nearly independent of seed production by trees. 1971 was one of the most important mast years of spruce within the examined time span (REICHHOLF 1974). As most of our data sets start in 1971 or later we are unable to document the increase in squirrel numbers: our data only show the fadıng of the “population wave”. Furthermore, the minor mast years also produced smaller waves, and they all superposed. This scenario provides an explanation of the decline in the red squirrel since the early 1970s (Fig. 3). The negative regression coefficient of the varıable year in Table 2 describes in part the lasting effects of a very prominent “population wave”. The weak effect of our mast index in Table 2 is an artefact, because some of the mast-induced variation in squirrel numbers ıs covered by the variable year. We attribute the observed decline of the red squirrel to natural varıations of the seed production of coniferous trees. The good correlations between data sets (Table 1) may be a consequence of synchronized masts across Bavaria. SCHRÖDER et al. (1982) describe a simulation model for the capercaillie (Tetrao urogallus) similar to our explanation of the squirrel dynamics. Random variations in reproduction may impose long-term cycles in a species, only because adults are long-lived and independent from the factors influencing reproduction. Bad weather conditions are the cause in the capercaillie, seed production in the red squirrel. We would like to stress that the data in Fig. 3 and the sımulations of SCHRÖDER et al. (1982) should act as examples for investigators studying the dynamics of long-lived species. A time series of 20 years may be Population dynamics of the Red squirrel in Bavaria 17 too short for a complete analysis, because historical events outside the analysed time span may have lasting and confusing effects. The high level of the squirrel index at Pressath between 1965 and 1967 is inconsistent with the view that the dynamics of the squirrel is mainly influenced by seed production. Möcker’s (1987) data indicate a very low density during that period. Consequently, one may suggest that the population density of the red squirrel dropped below the level of the late 1960s after an intermediate increase induced by spruce mast ın 1971. Note that we have no data for the period around 1971 for Pressath (question mark ın Fig. 3). Perhaps some people would invoke forest decline as a factor, which may negatively influence forst anımals. Damage ot forests ıs frequent ın northern Bavarıa (SCHULZE 1989). Forest decline has been prominent since the late 1970s, but the squirrel populations have increased since 1980 (see Fig. 3; Garmisch-Partenkirchen and B 12). Furthermore, forest decline may have a positive effect on species depending on seeds. The synchronized production of seeds is reduced by the damage to trees. Every year some trees produce seeds and the food supply may be more constant compared to periods with synchronized, but unpredictable seed production. Around Garmisch-Partenkirchen nests of the crossbill (Loxia curvirostra) were found nearly each year in the late 1980s, whereas in the 1970s broods were only observed during mast years (BEZZEL unpubl.). In the Fichtelgebirge the beech marten is negatively correlated to squirrel density (Fig. 5A). This suggests that within certain circumstances predators have some influence on prey species (note that the B 12 data did not show such a correlation!). Squirrels are only an alternative prey of the beech marten and the densitiy of the marten is regulated by different factors: the population dynamics of the squirrel has no effect on marten populations. PULLIAINEN (1984) found no increase in the pine marten after an increase in squirrels. In turn, an increase in predator populations may, increase predation pressure on alternative prey species. This situation is sımilar to the “alternative prey hypothesis” of ANGELSTAM et al. (1984), which explains why certain anımal species show synchronized fluctuations in Fennoscandıa. Voles are known to fluctuate wıth a cycle of about 3 to 4 years and the dynamics of the predators (like the red fox Vulpes vulpes) are coupled to this prey. When the main prey declines the predators turn to alternative prey species inducing a cylce on these prey species. Our explanation implies no fluctuations in the main prey of the beech marten; we only hypothesize that an increased predator density should have an inhibiting impact on the alternative prey. This ıs sımilar to JANZEN’s (1976) explanation of the low reptile biomass in Africa. JANZENn believes that many predators are able to maintain high population sizes because of the large herbivore biomass. The carnivores impose predation pressure on minor prey species and lower the reptile biomass (JanzEn 1976; but see KREULEN 1979). Acknowledgements We thank F. van DEN HEUVEL, who provided the squirrel data from Bad Berneck. We are also grateful to the forest authorities of Bayreuth (spruce mast) and the Wetteramt Nürnberg (weather) who made their data available to us. Zusammenfassung Populationsdynamik des Eichhörnchens in Bayern Für fünf Gebiete Bayerns werden die Populationsschwankungen des Eichhörnchens (Sciurus vulgaris) dargestellt. Zwischen 1970 und 1980 war eine Abnahme der Eichhörnchen zu beobachten, seit 1980 hingegen kam es wieder zu einer leichten Zunahme. Die Abnahme ist wohl keine Folge menschlicher Eingriffe, sondern kann auf Reproduktionswellen zurückgeführt werden, die durch die Mastjahre der Fichte bedingt sind. Die Zunahme seit 1980 wird als Folge des Waldsterbens diskutiert, da die Schädigung der Nadelbäume zu einer Desynchronisation der Zapfenproduktion geführt hat. Damit stand den Eichhörnchen eine konstante Nahrungsquelle zur Verfügung. Wetterfaktoren scheinen die Dynamik der Eichhörnchenpopulation nicht zu beeinflussen. Dage- gen finden wir eine negative Korrelation zwischen Steinmarder (Martes foina) und Eichhörnchen. 18 R. Brandl, E. Bezzel, J. Reichholf and W. Völkl References ANGELSTAM, P.; LINDSTRÖM, E.; WIDEn, P. (1984): Role of predation in short-term fluctuations of some birds and mammals in Fennoscandia. Oecologia 62, 199-208. Branpı, R. (1983): Populationstrend des Eichhörnchens Sciurus vulgaris L. in der Oberpfalz. Säugetierkundl. Mitt. 31, 76-78. ForMmosov, A. N. (1933): The crop of cedar nuts, invasions into Europa of the Siberian nutcracker (Nucifraga caryocatactes macrorhynchus Brehm) and fluctuations in numbers of the squirrel (Sciurus vulgaris L.). J. Anım. Ecol. 2, 70-81. GURNELL, J. (1983): Squirrel numbers and the abundance of tree seeds. Mammal Rev. 13, 133-148. Möckeı, R. (1987): Bemerkenswerter Massenwechsel des Eichhörnchens (Sciurus vulgarıs) im West- Erzgebirge. Säugetierkdl. Inf. 11, 489-493. Janzen, D. A. (1976): The dependence of reptile biomass by large herbivores. Am. Nat. 110, 371-400. KrEULEN, D. A. (1979): Factors affecting reptile biomass in African grasslands. Am. Nat. 114, 157-165. Ports, G. R.; TAPPER, $. C.; Hupson, P. J. (1984): Population fluctuations in red grouse: analysis of bag records and a simulation model. J. Anım. Ecol. 53, 21-36. PuLLiamen, E. (1973): Winter ecology of the red squirrel (Sciurus vulgaris L.) in northeastern Lapland. Ann. Zool. Fennicı. 10, 487494. PuLLıaınen, E. (1984): The predation system seed-squirrel-marten under subarctic conditions. Z. Säugetierkunde 49, 121-126. REICHHOLF, J. (1974): Die Auswirkungen der Fichtensamen-Rekordernte von 1971 auf die Populationen von Buntspecht und Eichhörnchen am Unteren Inn. Natur und Landschaft 49, 77-78. REICHHOLF, J. (1983): Bestandsentwicklung und Farbphasen des Eichhörnchens Sciurus vulgaris L. in Südbayern. Säugetierkundl. Mitt. 31, 73-75. . SCHRÖDER, W.; SCHRÖDER, ]J.; SCHERZINGER, W. (1982). Über die Rolle der Witterung in der Populationsdynamik des Auerhuhns (Tetrao urogallus). J. Orn. 123, 287-296. SCHULZE, E.-D. (1989): Air pollution and forest decline in a spruce (Picea abies) forest. Science 244, 776-783. STUBBE, H. (1988): Das Buch der Hege. Bd. 1: Haarwild. Berlin; VEB Deutscher Landwirtschafts- verlag. Tonkın, J. M. (1983): Activity patterns of the red squirrel (Sciurus vulgaris). Mammal Rev. 13, 99-111. WAUTERS, L.; DHONDT, A. (1989): Body weight, longevity and reproductive success in red squirrels (Scinrus vulgaris). J. Anım. Ecol. 58, 637-651. WILTAFskY, H. (1978): Sciurus vulgaris Linnaeus, 1758 — Eichhörnchen. In: Handbuch der Säugetiere Europas. Ed. by J. NIETHAMMER and F. Krapp. Wiesbaden: Akademische Verlagsgesellschatt. Bd. 1, 86-105. ZWAHLEN, R. (1975): Die lokomotorische Aktivität des Eichhörnchens (Sciurus vulgaris). Oecologia 22, 79-98. Authors’ addresses: Dr. RoLAnD BrAnDL and Dr. WOLFGANG VÖLKL, Lehrstuhl für Tierökologie I, Universität Bayreuth, Postfach 101251, W-8580 Bayreuth, FRG; Dr. EINHARD BEZZEL, Institut für Vogelkunde, Gsteigstr. 43, W-8100 Garmisch-Partenkir- chen, FRG; Prof. Dr. JosEr REICHHOLF, Zoologische Staatssammlung, Münch- hausenstr. 21, W-8000 München 60, FRG Z. Säugetierkunde 56 (1991) 19-24 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 An experiment to test the consumption of arboreal food by Wood mouse Apodemus sylvaticus By T. Santos and J. L. TELLERfA Departamento de Biologia Anımal (Zoologia), Universidad Complutense, Madrid, Espania Receipt of Ms. 24. 4. 1990 Acceptance of Ms. 31. 7. 1990 Abstract Analysed factors affecting the arboreal feeding activity of wintering wood mice in a set of woodlots ın central Spain. Stations were baited with acorns on branches, trunk and ground of 429 trees to test the arboreal behaviour of mice. Acorn intake on branches was positively and significantly related to accessibility of food but not to mice abundance. The best predictor of food consumption on branches was, however, the ground-trunk acorn intake, suggesting that this varıable was a good index of the scarceness of food. We conclude that morphological constraints in the arboreal ability of the species were evident. Introduction Wood mice (Apodemus sylvaticus) are recognized as expert climbers (HoLisovA 1969; LE Louarn and SAINT-GIRONS 1977; GURNELL 1985), but there ıs a lack of information about the factors determining this behaviour (MONTGOMERY and GurnELL 1985). The use of arboreal substrata may be subject to conflicting factors, such as predation risks, resource availability, and others (e.g. Kıng 1985; BRown 1989; SIMONETTI 1989). The trade-oft between rısks and benefits ın a particular microhabitat may change, however, as a result of increased pressures on other microhabitats nearby. For example, high population densities may force some individuals to use less preferred resources (FRETWELL 1972; GURNELL 1985). Thus, decreased availability of food, frequently related to peak densities (FLOWER- DEw 1985; Hansson 1985), may stimulate the exploratory behaviour of rodents (STICKLE 1979; MONTGOMERY and GURNELL 1985; PENNYCUIK and REISNER 1989). Morphological constraints and travel expenditures seem to be the another relevant factor influencing the selection of feeding microhabitats (WıEns 1973; GURNELL 1985; BROwNn 1989; SIMONETTI 1989). In this work, we explore the incidence of mice abundance and food accessibility on the arboreal feeding activity of wintering wood mice (Apodemus sylvaticus) in Holm oak woodlots of central Spain. The winter is a period wıth heavy food constraints for many temperate endotherms (NEwToN 1981; MERRIT 1984), so mice arboreality could be stimulated in this season. Study area The study area is located in Burgos province, central Spain. Average altitude is 850 m and phytoclima- tie conditions are inland mediterranean, with long and cold winters (Rıvas-MArTinEz 1981). The area has a deforested agricultural landscape, with extensive cereal cropping and sheep grazing as main usages. Some tracts of the original forests remain, with a dominant vegetation formed by Holm oaks (Quercus rotundifolia) and shrubs of the genera Cistus, Genista, Thymus and Lavandula. These woodlots support wintering populations of wood mouse, but mice are scarce during winter in the surrounding fields (TELLERTA et al. 1990; see also ALCAnTARA 1986). Previous trapping work showed that rodent species other than wood mice are scarce or absent in these forests; Common voles (Microtus arvalıs) settle some grassy sectors and Garden dormice (Ehomys quercinus) hibernate throughout the winter. Seventeen isolated woodlots, ranging from 0,1 to 280 ha (Table), were studied in order to evaluate the use of arboreal preys by mice. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0019 $ 02.50/0 20 T. Santos and J. L. Telleria The 1988 autumn crop of acorn, preceding the study winter, was very scarce; at the start of December only 0.8 % (n = 131) of the Holm oaks beared fruits and 3.2 % (n = 158) had shed acorns to the surrounding ground. By the 20th of January a high degree of depletion was evident, since tree and ground acorns were lacking in most woodlorts. Material and methods Food accessibility Holm oak half-acorns were pierced with wire and tied round branches with different diameters and at different distances from the tree basıs to test the relationship between the arboreal feeding efficiency of mice and food accessibility. The wood mouse is an omnivore, and includes fruits in its diet (OBRTEL and Houısova 1983; Hansson 1985), so that we could assume that Holm oak acorns should be profitable baits. Acorns were arranged during one dark night in each woodlot, and placed and controlled in the same order, so all acorns were exposed the same time. Two experiments were carried out in 1989, from 12 to 16 January and from 2 to 5 February. One half-acorn per tree were tied on 429 trees distributed among the 17 Holm oak forests. Numbers of acorns ranged from 6 to 49 in each experiment (Table), according to a logarıthmic function of forest area. Branch diameter was estimated visually for 380 acorns; distance from ground was estimated for each acorn as the shortest path from the base of the tree. Branches ranged from 0.3 to 10 cm in diameter, with the majority between 0.5 and 3cm (87.9 %). Distances ranged from 0.5 to 12m, those between 2 and 4m accounting for 78.5 %. Half-acorns were also placed on the trunk, 1.5 m above ground and on the base of the trunk, in the same trees, to assess the global food searching incidence by mice in each forest. As wintering mice mainly use underground burrows (DUFOUR 1978; MONTGOMERY and GURNELL 1985), accessibility of arboreal food should be negatively related to ground distance, and positively to branch diameter for morphological reasons. Since separate analysis of these two varıables can produce confounding results, a ditficulty index (ID) was estimated for each acorn as the product of distance by the inverse of diameter; ID ranged from 0.2 to 26.7, with values between 0.2 and 6 summing 92.1 %. We calculated mean values of distance, diameter and ID for each forest. Percentages of acorns preyed upon by mice were used as estimates of feeding incidence in the three positions sampled. Acorn use on ground was significantly associated with acorn use on trunks (r = 0.824, p < 0.001, 15 d.f.), therefore both positions were analysed together. Acorn intakes in January and February were positively correlated (r = 0,78, p < 0.001, 15 d.f.; see TELLERTA et al. 1990), so both samples were pooled. Preyed acorns were easily recognizable by teeth marks and fecal rests. The acorns preyed by birds or lost were not considered in the analyses. Mice abundance At the beginning of March 1989, 213 stations were provided with two snap-traps during two consecutive nights (SMITH et al. 1975); number of traps ranged from 12 to 196, according to woodlot area. Abundance indexes were obtained as the number of individuals captured by trap in each forest (Table; see TELLERfA et al. 1990, for more details on the trapping design). Wood mouse was the only species caught. Statistical analyses were made according to SokaL and RoHLF (1981) and Zar (1984). Mice abundance and acorn intake rates were arcsin transformed, and distance, dıameter and ID values logtransformed. Results Branch acorns were exclusively preyed upon in locations placed between 0.5 and 3.5 m from the base of the tree; 71 acorns arranged between 3.6 and 12 m remained untouched, most of these placed in the canopy edge. Acorns were consumed from branches of all diameters, although twigs of less than 1 cm supported the lowest predation (see Figure). Relationship between acorn predation rate and accessibility was first explored classıfying the acorns in 9 classes according to distance, diameter and ID. The results show that the proportion of preyed acorns decreased significantly with the increase of distance and ID, but a diameter response was not obvious (Figure). ID was the only variable among these three used in the following analyses, since it may be considered a synthetic index representative of acorn accessibility. An experiment to test the consumption of arboreal food by Wood mouse 2] Table. Area, mice abundance (No. individuals/trap x 100), number of acorns placed and controlled (N) and acorn intake rates (percentage of acorns preyed) in each forest woodlot mice intake area abundance ground branches According to relationships between the feeding incıdence on branches and the indepen- dent varıables estimated in each forest, ID accounted for more varıatıon (r = -0.569, p < 0.05; R? = 32.4 %) than mice abundance (r = 0.183, n.s.; R? = 3.3 %), but this was less than that accountable for acorn intake on ground and trunks (r = 0.711, p < 0.01; R’ = 50.6 %). Ihese results were supported by partial correlation analyses of feeding incıdence on branches with 1) mice abundance and ID: partial r values were 0.287 (n.s.) for abundance and -0.599 (p < 0.02) for ID; 2) with mice abundance and ground-trunk acorn intake: r = -0.299 (n.s.) and r = 0.731 (p < 0.01), respectively; and 3) with ground-trunk acorn ıntake and ID: r = 0.649 (p < 0.01) and r = -0.457 (n.s.), respectively. ACORN INTAKE ON BRANCHES 30 Ps:-0.905** 30 f3:-0.933%** 26 17 18 _90 09 225 33545 6 61 09 1 15 2 253 4 5 541 distance diameter ID Relationships of acorn intake on branches (in percentages) to distance (m), diameter (cm) and acorn accessibility (ID). r, is the Spearman’s rank correlation coefficient; **: p < 0.01 22 T. Santos and ]J. L. Telleria Discussion Mice consumed acorns placed in branches regardless of diameter and up to a distance 3.5 m from the tree base, but not further away. Thus, our results agreed with the arboreality of the species, although a limit in its arboreal exploration was apparent in our study. This could be directly due to constraints in movement ability of mice in trees, but other factors may operate simultaneously (CArAco 1980). In crowded conditions, some island populations of rodents increase their resource partitioning, with younger individuals obliged to live in suboptimal patches (GrLiwıcz 1984; see also Lomnick1 1988). Therefore, we should expect a rıse of food demands with the increase in mice density in the woodlots studied, and hence a searching increase towards the least accessible acorns. Since this expected relationship did not occur, it is plausible that even the higher densities obtained were very low and mice did not need to explore the most inaccessible acorns; actually, mice abundance accounted for a relatively low variance of the acorns consumed on ground and trunks (R? = 26.2 %). On the other hand, if resource levels differed among woodlots, the total acorn intake would be a better estimator of trophic stress than mice abundance. In fact, some small woodlots with “low” abundances of mice gave the highest intakes (Table). This view is consistent with our results, suggesting that under conditions of limited food supply mice could search for supplementary food above ground. Also, an increase of searching actıvity in the adjacent fields, including dispersal behaviour, could be an alternative response. Exploratory and dispersal ability, agricultural field use and habitat tolerance in the Wood mouse (JONGE and DiEnskeE 1979; JENSEN 1982; GEUSE et al. 1985; GurNnELL 1985; WOLTON and FLOWERDEW 1985), suggest that this response would be more profitable than an exploratory increment towards presumably little accessible substrata, such as branches, but at present experimen- tal evidence is lackıing. In conclusion, the findings of our experiment, namely, the clear limit observed in the ability of the Wood mouse to obtain arboreal food, and the association of acorn use on branches with ID, suggest morphological constraints of this species in the searching of tree food, and would indicate a negative yield in the handling of distant arboreal preys. At the same time, this study suggests that mice are capable of overcoming local situations of food scarcity on the ground by using alternative arboreal resources. In another context, the decreasing gradient observed in the feeding incidence of mice from ground to trunk-branches, implies potential consequences for forest avian species feeding mainly on the ground and trunks (e.g. pariforms; see ULFSTRAND 1977). The high densities of mice registered in island conditions in this and other works (e.g. GLıwIcz 1984; GEUSE et al. 1985), point out a probable competition between birds and rodents whose main biological effects are unknown at present, although composition and densities of avıian species could be affected. As JanzEn (1986) recently expressed, other topics than customary (area, distance to continent, habitat diversity, etc.) are involved ın island biology; the ones related to high densities of generalist anımals, such as wood mice (FLOWERDEW 1977; LE LOUARAN and SAINT-GIRONS 1977), represent an ımportant threat for stenoecious or scarce organısms. Acknowledgements We thank MAnUEL ALCANTARA for field assistance and TomAs SANTOS Rıncön for useful advice. JosE A. Diaz made helpful comments on a previous draft of the manuscript. This paper is a contribution to the project “Biology and distribution of Iberian forest vertebrates”, funded by the Spanish C.1.C.Y.T. (project PB-86-0006-C02). An experiment to test the consumption of arboreal food by Wood mouse 28 Zusammenfassung Ein Experiment zur Ermittlung der Nahrungsaufnahme der Waldmaus (Apodemus sylvaticus) auf Bäumen Untersucht wurden mehrere Faktoren, die die Aktivitäten von Waldmäusen bei der Nahrungssuche auf Bäumen beeinflussen. Dazu wurden in mehreren Waldparzellen in Zentralspanien während des Winters Eicheln am Boden, an Stämmen und auf Zweigen von 429 Bäumen angeboten. Der Konsum von Eicheln auf Zweigen war positiv und signifikant korreliert mit der Erreichbarkeit der Nahrung, aber nicht mit der Häufigkeit der Waldmäuse. Die Ficheln wurden vornehmlich von Boden und Stämmen aufgenommen, aber auch von Zweigen unterschiedlicher Stärke, allerdings nur bis zu 3,5 m Höhe. Dieses weist auf Grenzen der Art hin, den arboricolen Lebensraum zu nutzen. References ALCANTARA, M. (1986): Preferencias de häbitat del Ratön de Campo (Apodemus sylvaticus L.) en medios agricolas. Memoria Lic. Universidad Complutense, Madrid. Brown, J. S. (1989): Desert rodent community structure: a test of four mechanisms of coexistence. Ecol. Monogr. 59, 1-20. f CArRAco, T. (1980): On optimal selection in a multiattribute stochastic environment. Evol. Theor. 5, 127-133. Durous, B. (1978): Effect of external and internal factors upon burrowing and nest-building activities of the wood mouse (Apodemus sylvaticus L.) ın terrarıum conditions. Behav. Processes 3, 57-76. FLOWERDEW, J.R. (1977): Wood mouse Apodemus sylvaticus. In: The Handbook of British Mammals. Ed. by G. B. CorgET and H. N. SOUTHERN. Oxford: Blackwell, pp. 206-217. — (1985): The population dynamics of wood mice and yellow-necked mice. Symp. zool. Soc. Lond. 55, 315-338. FRETWELL, S. (1972): Populations in a seasonal environment. Princeton: Princeton Univ. Press. GEUSE, P.; BaucHat, V.; LE BOULENGE, E. (1985): Distribution and population dynamics of bank voles and wood mice in a patchy woodland ın central Belgium. Acta Zool. Fennica 173, 65-68. Giwicz, J. (1984): Rodents on islands: living in a crowd. Acta Zool. Fennica 172, 95-98. GURNELL, J. (1985): Woodland rodent communities. Symp. zool. Soc. Lond. 55, 377411. Hansson, L. (1985): The food of bank voles, wood mice and yellow-necked mice. Symp. zool. Soc. Lond. 55, 141-168. Horısova, V. (1969): Vertical movements of some small mammals in a forest. Zool. Listy 18, 121-141. JANZEn, D. H. (1986): The eternal external threat. In: Conservation Biology. The Science of Scarcity and Diversity. Ed. by M. E. Souz£. Sunderland: Sinauer, pp. 286-303. JENsEn, T. S. (1982): Seed production and outbreaks of noncyclic rodents populations in deciduous forests. Oecologia, Berlin, 54, 184-192. JONGE, G. DE; DIENSKE, H. (1979): Habitat and interspecific displacement of small mammals in The Netherlands. Netherlands J. Zool. 29, 177-214. Kıng, C. M. (1985): Interactions between woodlands rodents and their predators. Symp. zool. Soc. Lond. 55, 219-247. LE Lovarn, H.; SAINT-GIRoNSs, M.-C. (1977): Les Rongeurs de France. Parıs: I.N.R.A. LomnIckt, A. (1988): Population ecology of individuals. Princeton, N. J.: Princeton Univ. Press. MERRIT, J. F. (ed.) (1984): Winter ecology of small mammals. Pittsburgh: Spec. Publ. Carn. Mus. Nat. Hist. 10. MONTGOMERY, W.1.; GURNELL, J. (1985): The behaviour of Apodemus. Symp. zool. Soc. Lond. 55, 89-115. NEwTon, 1. (1981): The role of food in limiting bird numbers. In: The integrated study of bird populations. Ed. by H. Kromp and J. W. WOLDENDORP. Amsterdam: North Holland Publ. Comp, pp. 11-30. OBRTEL, R.; HoLisova, V. (1983): Winter and spring diets of three coexisting Apodemus spp. Folia Zoologica 32, 291-302. PEnNnycuIk, P. R.; REISNER, A. H. (1989): Food restriction: mechanisms limiting numbers and home site utilization in populations of house mice, Mus musculus. Oikos 55, 159-164. Rıvas-MARTINEZ, $. (1981): Les &tages bioclımatiques de la vegetation de la Peninsule Iberique. Anales Inst. Bot. Madrid 37, 251-268. SIMONETTI, J. A. (1989): Microhabitat use by small mammals in central Chile. Oikos 56, 309-318. SMITH, M. H.; GARDNER, R. H.; GENTRY, J.; Kaurman, D. W.; O’FARRELL, M. H. (1975): Density estimators of small mammal populations. In: Small mammals: Their productivity and population dynamics. Ed. by F. B. GoLı£v, K. Perrusewicz and L. Ryszkowskı. Cambridge: Cambridge Univ. Press, pp. 25-53. 24 T. Santos and ]J. L. Telleria SoKAL, R. R.; ROHLF, F. J. (1981): Biometry. 2nd ed. San Francisco: W. H. Freeman. STICKLE, L. F. (1979): Population ecology of house mice in unstable habitats. J. Anım. Ecol. 48, 871-887. TELLERSA, J. L.; SANTOS, T.; ALCANTARA, M. (1990): Abundance and food-searching intensity of wood mice (Apodemus sylvaticus) in fragmented forests. J. Mammalogy (in press). ULFSTRAND, $. (1977): Foraging niche dynamics and overlap in a guild of passerine birds in a south Swedish coniferous woodland. Oecologia, Berlin, 27, 23-45. WiıEns, J. A. (1973): Pattern and process in grassland bird communities. Ecol. Monogr. 43, 237-270. WOLToN, R.; FLOWERDEW, J. R. (1985): Spatial distribution and movements of wood mice, yellow- necked mice and bank voles. Symp. zool. Soc. Lond. 55, 249-275. ZAR, J. H. (1984): Biostatistical Analysıs. 2nd. ed. London: Prentice-Hall. Authors’ address: TomAs Santos and Jos£ Luıs TELLERfA, Departamento de Biologia Animal (Zoologia), Facultad de Biologia, Universidad Complutense, E-28040 Madrid, Espana Z. Säugetierkunde 56 (1991) 25-33 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Genetic differentiation in four species of Apodemus from Southern Europe: A. sylIvaticus, A. flavicollis, A. agrarius and A. mystacinus (Muridae, Rodentia) By JAnIcE BRITTON-DAVIDIAN, MEHRNOUCH VAHDATI, FATIMA BENMEHDI, PAULE GROS, VALERIE NAnc£, H. CROSET, $. GUERASSIMOV and C. TRIANTAPHYLLIDIS Laboratoire de Genetique, Institut des Sciences de !’Evolution, USTL, Montpellier, France Receipt of Ms. 15. 5. 1990 Acceptance of Ms. 10. 8. 1990 Abstract Genic varıability was estimated for four Apodemus species from Southern Europe by electrophoresis at 20 loci. Genic divergence data agree well with the subgeneric classification, A. agrarıns (subgenus Apodemus) being very distant from the three other species (subgenus Sylvaemus), as was previously shown by GEMMERkE (1980) using 11 locı. However, the large dıvergence between these two groups suggests that their taxonomic relationships should be revised. The morphological overlap between A. sylvaticus and A. flavicollis is confirmed throughout Southern Europe which stresses the use of biochemical methods for unambiguous identification of specimens. The karyological analysis of samples of A. sylvaticus, A. flavicollis and A. agrarins revealed chromosomal varıation in only one individual (A. flavicollis wıth 2 n = 49 ın Bulgarıa). Introduction The genus Apodemus ıs one of the most widespread noncommensal rodent groups ın the Paleartic. In the Western part of their range, the use of biochemical genetics has proven extremely useful in unambiguously discriminating morphologically similar species. This has been the case for A. sylvaticus and A. flavicollis, ın particular, which although easily distinguishable in Central and Northern Europe, show a morphological overlap in the more southern areas. That this overlap is due to clinal variation in sıze and pelage color following opposite trends in both species and not to hybridization, has been shown by a number of authors (NIETHAMMER and Krarp 1978; GEMMEKE 1980; BENMEHDI et al. 1980; NascETTI and Fırıppuccı 1984). This discrete morphological varıation seems to be a general trait within the subgenus Sylvaemus (A. mystacinus excepted), although genic differentiation as measured by electrophoretic methods is quite extensive. Recent studies of more eastern populations suggest that this subgenus will most likely reveal a complex of species (NIETHAMMER 1969; DARVICHE et al. 1979; GEMMERE and NIETHAMMER 1982). Karyotypie varıability ıs also well documented within the subgenera Sylvaemus and Apodemus. All species carry 48 chromosomes but differ in the NF number showing that chromosomal evolution has proceeded mainly by pericentric inversions (KrAaL 1970; SOLDATOVIC et al. 1975; for a review see ZımA and Krau 1984). The evolutionary relationship between Apodemus species was previously investigated at 11 locı by GEMmMERE (1980). The genetic differentiation between Apodemus species belonging to the two subgenera Sylvaemus (A. sylvaticus, A. flavicollis, A. mystacinus) and Apodemus (A. agrarins) is here extended to populations from Greece, Bulgaria and Spain for which morphological, chromosomal and allozymic data at 20 loci are presented. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0025 $ 02.50/0 26 Janıce Britton-Davidıan et al. Material and methods Specimens belonging to four species of the genus Apodemus (A. sylvaticus, A. flavicollis, A. mystacinus and A. agrarins) were live-trapped in localities (see Figure 1 for names) from five southern European countries. Nine populations were analyzed for genic varıability: France (1), Italy (2), Greece (3, 4, 5, 6), Spain (7) and Bulgarıa (8). Starch gel electrophoresis techniques are described in PASTEUR et al. (1987). A total of 20 locı were analysed: alcohol dehydrogenase (Adh), alpha-glycerophosphate dehydrogenase (alpha-Gpd), albumin (Alb), amylase (Amy-1), glutamate oxaloacetate transaminase (Got-1), glucose phosphate isomerase (Gpi), hemoglobin (Hbb), ısocitrate dehydrogenase (/dh-1 and Idh-2), lactate dehydrogenase (Ldh-1 and Ldh-2, respectively coding for the A and B subunits; see EnGEL et al. 1973), the regulator gene for Zdh-2 ın red cells (Ldr), NAD-dependent malate dehydrogenase (Mdh-1 and Mdh-2), NADP-dependent malate dehydrogenase (Mod-1), nucleoside phosphorylase (Np), phosphoglucose dehydrogenase (Pgd), phosphoglucomutase (Pgm), sorbitol dehydrogenase (Sdh) and superoxide dismutase (Sod). Allelic designations were determined by comparison to the mobility of the most frequent allele in A. sylvatıcus which was arbitrarily assıgned the value 100. The HBB pattern in Apodemus consists in two spots, the slower one showing no variation, so the genetic varıability scored refers only to variation ın mobility of the faster migrating spot. Genetic varıabılity measures, genetic distances (NEI 1978) and the UPGMA phenogram were computed and performed using the BIOSYS-1 program of SWOFFORD and SELANDER (1981). . sylvaticus . flavicollis . agrarius . mystacinus Fig. 1. Distribution of sampled localities and technique of analysıs: empty symbols = allozymes; dark symbols = morphology; dotted symbols = chromosomes. Localities: France (1. St Cyr-les-Colons, 9. Gardiole, 10. Avene, 21. Mauguio, 22. Iseron, 25. Cuxac, 26. Banyuls); Italy (2. Ticino region near Pavia); Greece (3. Gallikos, 4. Strimonikon, 5. Doiranı, 6. Prosotsanı hills, 16. Aggitis river, 17. Prosotsani river, 18. University farm Thessaloniki, 19. Avas); Spain (7. Calonge, 11. Granada); Bulgaria (8. Plovdiv, 12. Karnobat, 13. Vranıa, 14. General Toshevo, 23. Orizare); Germany (20. Lübeck) and Austria (24. Burgenland) Morphological data were collected on specimens from France (1, 9, 10), Italy (2), Spain (7, 11), Bulgaria (8, 12, 13, 14), Greece (3, 15, 16, 17, 18, 19) and Germany (20). The chromosomal study was performed on field mice from difterent localıties: A. sylvatıcus from France (9:1 male, 21:1 male, 22:1 female, 26:1 male), Spain (11:1 female), Bulgaria (13:1 female, 23:1 female) and Austria (24:1 female); A. flavicollis from France (25:1 female) and Bulgaria (14:1 male, 13:1 male); A. agrarius from Greece (4:1 male, 1 female). The karyotypes were established using the classical air-drying technique. No results were available for A. mystacınus. In both the morphological and chromosomal study, all mice, except for A. agrarins and A. mystacinus which were unambiguously identified in the field, were ascribed to A. sylvatıcus or A. flavicollis on the basis of their LDH-2 alleles. The specimens are deposited as pickled carcasses at the Institut des Sciences de l’Evolution. Genetic dıfferentiation in four species of Apodemus from Southern Europe 27 Table 1. Allele frequencies at the 19 variable loci Species / Locality 28 Janıce Britton-Davidıan et al. Table 1 (continued) Species / Locality 1.00 0.00 1 0.00 1.00 0.00 AS=A. sylvaticus; AF = A. flavicollis; AA = A. agrarıns; AM = A. mystacinus. FR = France; IT = Italy; GR = Greece; SP = Spain; BU = Bulgaria. Results Genetic variability and differentiation Genetic varıability parameters (Table 2) were computed from the allelic frequencies (Table 1). Only one locus (/dh-2) was found monomorphic for the same allele in the four Apodemus species. Twelve locı (Alb, Amy-I, Got-1, c+l.c, Hbb, Idh-1, Lah-1, Ldh-2, Mdh-1, Pgd, Sdh and Sod) discriminated at least two of the four species, whereas two locı were species diagnostic: Mod-! and Np. All varıiability parameters were highest for A. sylvatıcus (mean H = 0.08, mean Po.o5 = 21% and mean A = 1.3) than for the other three species which ranged between 10 %-15 % for the rate of polymorphism and 4-6 % for mean heterozygosity. However, where sample sizes are small, this measure of variability may not be representative of the species as a whole. Genetic differentiation in four species of Apodemus from Southern Europe 2) Genetic distances (Table 3) calculated Table 2. Genetic variability measures according to NEI (1978) were used to gen- erate aUUPGMA phenogram (Fig. 2). The Locality latter agrees well with expectations ın that A. sylvaticus and A. flavicollis cluster to- A. sylvaticus gether, and both of these wıth A. mysta- a cinus. A. agrarius, on the other hand is set en 6) well apart from the first three species, Spain (7) yielding a mean genetic distance of 1.28. Mean Intraspecific genetic distances were com- A. flavicollis puted for A. sylvaticus and A. agrarius only France (1) and yielded respectively a mean of 0.045 A. agrarins and 0.067 which falls within the value gen- Greece (5) erally recorded for subspecific genetic Bulgaria (8) differentiation. Mean A. mystacınus Greece (6) 2 lei Morphological discrimination mean number of alleles; P = Morphological data are presented ın Table polymorphic locı (5 % level); H mean 4 and Figure 3. Whereas discrimination of heterozygosity (NEı 1978 except for A. mys- tacınus for which the sample size was too A. agrarins and 4. ST IE small). Numbers in parenthesis refer to phological grounds is immediate, this ıs not Inealliice the case between A. sylvatıcus and A. flavicollis which show a large overlap ın body lengths. Additionally, although the presence of a complete collar is species diagnos- tic, its absence is not since 35 % of the A. flavicollis we captured exhibited only a more or less large chest spot ın lieu of a collar. France A. sylvaticus Italy Greece Spain A. flavicollis France A. mystacinus Greece . agrarius Greece Bulgaria Nei's (1978) genic distance 1.4 1.2 1.0 0.8 0.6 0.4 0.2 009 Fıg. 2. UPGMA phenogram computed from genic distances Chromosomal variability The chromosomal analysis of specimens from each species corresponded to results from previous studies (Krar 1970; SOLDATOVIC et al. 1975; BEKASOva et al. 1980; ZımA 1984). The karyotypes of A. sylvaticus and A. flavicollis both carrıed 48 acrocentric chromosomes except for the A. flavicollis from Bulgarıa (14. General Toshevo) which had 49 chromosomes showing an additional small acrocentric chromosome. Such varıation is common in this species and has been attributed to the presence of supernumerary chromosomes (SOLDATOVIG et al. 1975; Zıma 1984; Zıma and Krar 1984). A. agrarıns (2n 30 Janice Britton-Davidian et al. Table 3. Genetic distance coefficients (NEı 1978) Locality . AS-FR -—-- AS-IT 0.016 = —— AS-GR 0.065 0.067 === AS-SP 0.039 0.063 0.022 AF-FR 0.601 0.548 0.478 ; ===> AA-GR 1.236 1.284 11415 x 1.446 -—-- AA-BU 15133 1.166 1.170 ; 16522 0.067 = . AM-GR 0.876 0.866 0.932 . 0.823 12592 LS 1 2R 38 Zi Io 6. 7 8 AS = A. sylvaticus; AF = A. flavicollis; AA = A. agrarius; AM = A. mystacinus. FR = France; IT = Italy; GR = Greece; SP = Spain; BU = Bulgarıa. Table 4. Morphological data for samples of three Apodemus species Numbers after countries refer to localities sampled Chest spot Locality BL DE (mm) (mm) A. sylvaticus Italy (2) 93(3) 7765) 0.84(0.07) Bulgaria (8. 12-14) 96(6) 81(4) 23(0) 0.86(0.05) Spain (7. 11) 93(5) 88(7) 22(1) 0.96(0.07) Greece (3. 15-18) 107(2) 91(2) 23(0) 0.85(0.02) France (1) 84(2) 792) 21(1) 0.93(0.02) (9) 97(2) 84(3) 21(1) 0.87(0.03) Mean 97(2) 84(1) 22(0) 0.88(0.01) A. flavicollis Greece (16-17) Yo) ee 2 ee Bulgaria (8. 12-14) 98(7) 86(1) 22(2) 0.88(0.08) France (10) 94(5) 95(4) 23(1) 1.03(0.04) (1) 94(5) 99(5) 24(0) 1.06(0.03) Mean 95(3) 958) 23(0) 1.01(0.03) A. agrarıns Greece (5) 11265) 78(2) 22(1) 0.69(0.02) Germany (20) 6 102(5) 70(12) 19(1) 0.69(0.17) Mean 108(4) 756) 21(1) 0.69(0.05) BL = body length; TL = tail length; HF = hind foot length; chest spot color: no, small (Sm) or large (L) spot; Col = complete collar. STandard errors in parenthesis. = 48) showed four pairs of small metacentric chromosomes which increased the NF to 56. No other chromosomal variability was found within samples. Discussion Interspecific genetic differentiation at 20 loci is presented for four species of the genus Apodemus. Previous studies on these same species were made by GEMMERE (1980) but the electrophoretic survey included only 11 loci. More recent and extensive work was performed by Nascerri and Fırıppuccı (1984) on 27 loci but was restricted to the subgenus Sylvaemus. The overview of these three independent studies shows that the Genetic differentiation in four species of Apodemus from Southern Europe 3 Li 120 110 100 den 90 Bas 25 (@) oO 00 ©) 80 70 BL(mm) 70 80 90 100 110 120 Fig. 3. Body (BL) and tail length (TL) distribution. Empty circle = A. sylvaticus; dark circle = A. flavicollis. Envelopes show the morphological varıability of samples from Bulgaria and Greece for both species overall specific discrimination (the number of diagnostic locı between two species) is sımilar and often more important with the set of locı and the electrophoretic buffers we used. This reasoning is based on the assumption that the differences in scoring results are probably more related to the techniques used than to varıability of the biological material. These differences in discriminating capacıty do not, however, alter the phylogenetic relationships as determined by the genetic distances. Our results agree wıth previous work on this genus (GEMMERE 1980) ın that A. sylvaticus, A. flavicollis and A. mystacinus belonging to the same subgenus Sylvaemus cluster together whereas A. agrarius, which represents a different subgenus (Apodemus) ıs exterior to this group. That morphological discrimination is difficult and even impossible between specimens of A. sylvaticus and A. flavicollis inhabiting Southern Europe has previously been shown for France (BENMEHDI et al. 1980), Italy (NascETTI and FıLıppuccı 1984) and Germany (EnGEL et al. 1973; GEMMERE 1980) and is here extended to Bulgarıa and Greece. Previous results are here again confirmed in that in all cases of morphological ambiguity, the biochemical analysis allowed to assıgn the specimens to either species and showed the absence of any introgression between them. It is therefore suggested that field specimens be identified by electrophoretic methods (on albumin for example, DEBROT and MERMOD 1977; GEMMERE 1981) or by using the morphological criteria put forth by FıLıppucct etal. (1984) which enabled to discriminate at least 95 % of Italian specimens and should be tested elsewhere. This study shows that the separation of the four Apodemus species studied into two subgenera is supported by the biochemical distance data. Within the subgenus Sylvaemus, A. sylvaticus and A. flavicollis are remarkable in that they represent morphologically and chromosomally very similar species with relatively large genic distances. The particular position of A. agrarius is worth commenting on, however. The very important genic distance between this species and those of the subgenus Sylvaemus is 92 Janice Britton-Davıdian et al. probably an underestimate, being at the limit of the discriminating power of electrophore- tic methods. In fact, IsKANDAR and BONHOMME (1984) showed that sequential elec- trophoresis allowed to uncover 27 % more alleles between the two subgenera whereas no additional variation was revealed between the three species of the subgenus Sylvaemus. These data then suggest that A. agrarıns ıs probably even more distantly related to the Sylvaemus species group than what we indicate herein. Based on these data, we agree with BONHOMME et al. (1985) in suggesting that a taxonomıic revision of this group be made. It is probable that the two current subgenera will be elevated to a genus rank since Apodemus appeared not to be more closely related to Sy/vaemnus than to other murid (BONHOMME et al. 1985) or even arvicolid (data not based on sequential electrophoresis: GILL et al. 1987) genera. For this analysis, it would be imperative that biochemical data be collected for species belonging to the third subgenus (Alsomys) in order to correctly establish the evolutionary relationships within the Apodemus complex. Acknowledgements We wish to thank F. BONHOMME, J. CATALAN, ]J. CASSAING, F. CATZEFLIS, ]. GROBERT, D. ISKANDAR, U. Jünes, P. Orsınt, F. Portevin and L. THALER for their help in gathering and processing specimens and discussions. Field trips were financially supported by a NATO grant and Franco-Bulgarian cooperation agreements. Zusammenfassung Genetische Differenzierung bei vier Apodemus-Arten in Südeuropa: A. sylvaticus, A. flavicollis, A. agrarius und A. mystacinus (Muridae, Rodentia) Die genetische Variabilität von vier südeuropäischen Apodemus-Arten wurde durch Elektrophorese von Proteinen geschätzt, die über 20 Genloci kodiert werden. Die nach Neı (1978) berechneten Abstandswerte entsprechen ungefähr den bisherigen Vorstellungen von der abgestuften Verwandt- schaft dieser Arten. So unterscheidet sich A. agrarins (Untergattung Apodemus) beträchtlich von den drei anderen, in der Untergattung Sylvaemus zusammengefaßten Arten. Apodemus sylvaticus und A. flavicollis überschneiden sich in ihren morphologischen Merkmalen in Südeuropa stark, lassen sich aber gelelektrophoretisch stets einwandfrei bestimmen. Alle vier Arten haben gewöhnlich 48 Chro- mosomen. Nur 1A. flavicollis von 27 aus Bulgarien hatte mit einem kleinen zusätzlichen akrozentri- schen Element insgesamt 49 Chromosomen. Literature BEKAsOva, T. S.; VORONTSov, N. N.; KOROBITSYNA, K. V.; KoRABLEV, V.P. (1980): B-chromosomes and comparative karyology of the mice of the genus Apodemus. Genetica 52/53, 3343. BENMEHDI, F.; BRITTON-DAVIDIAN, J.; THALER, L. (1980): Premier apport de la genetique biochimi- que des populations ä la systematique des mulots de France continentale et de Corse. Biochem. Genet. Ecol. 8, 309-315. BONHOMME, F.; ISKANDAR, D.; THALER, L.; PETTER, F. (1985): Electromorphs and phylogeny in muroid rodents. In: Evolutionary Relationships among Rodents. Ed. by W. P. LuckErTT and ].-L. HARTENBERGER. London: Plenum Publishing Corp. 671-683. DARVICHE, D.; BENMEHDI, F.; BRITTON-DAVIDIAn, J.; THALER, L. (1979): Donnees preliminaires sur la systematique biochimique des Genres Mus et Apodemus en Iran. Mammalıa 43, 427-430. DEBROT, $.; MERMOD, C. (1977): Chimiotaxonomie du genre Apodemus Kaup, 1829 (Rodentia, Muridae). Rev. Suisse Zool. 28, 521-526. EnGEL, W.; VoGEL, W.; VoıcuLsscv, 1.; RopErs, H.-H.; ZEnzEs, M. T.; BENDER, K. (1973): Cytogenetic and biochemical differences between Apodemus sylvaticus and Apodemus flavicollis, possibly responsible for the failure to interbreed. Comp. Biochem. Physiol. 44B, 1165-1173. Fırıppuccı, M. G.; CrISTALDI, M.; Tızı, L.; Contouı, L. (1984): Datı morfologici e morphomerrici in popolazioni dı Apodemus (Sylvaemus) dell’Italia centro-meridionale determinati elet- troforeticamente. Ric. Biol. Selv. 9, 85-125. GEMMEKE, J. (1980): Proteinvariation und Taxonomie in der Gattung Apodemus (Mammalıa, Roden- tia). Z. Säugetierkunde 45, 348-365. GEMMEKE, H. (1981): Albuminunterschiede bei Wald- und Gelbhalsmäusen (Apodemus sylvaticus und A. flavicolls, Mammalia, Rodentia) auch in getrockneten Muskeln und Bälgen elek- trophoretisch nachweisbar. Z. Säugetierkunde 46, 124-125. Genetic differentiation in four species of Apodemus from Southern Europe 389 GEMMEKE, H.; NIETHAMMER, ]. (1981): Die Waldmäuse Apodemus sylvaticus und A. flavicollis vom Monte Gargano (Süditalien). Z. Säugetierkunde 46, 162-168. GEMMEKE, H.; NIETHAMMER, J. (1982): Zur Charakterisierung der Waldmäuse (Apodemus) Nepals. Z. Säugetierkunde 47, 33-38. GiLL, A.; PETRov, B.; Zıvkovid, $.; Rımsa, D. (1987): Biochemical comparisons in Yougoslavian rodents of the families Arvicolidae and Muridae. Z. Säugetierkunde 52, 247-256. ISKANDAR, D.; BONHOMME, F. (1984): Variabilite electrophoretique totale a 11 locı structuraux chez les rongeurs murides (Muridae, Rodentia). Can. J. Genet. Cytol. 26, 622-627. Krar, B. (1970): Chromosome studies in two subgenera of the genus Apodemus. Zool. Listy 19, 119-134. NAscETTI, G.; FıLıppuccı, M. G. (1984): Genetic variability and divergence in Italian populations of Apodemus sylvaticus and Apodemus flavicollis (Rodentia, Muridae). Ric. Biol. Selv. 9, 75-83. Neı, M. (1978): Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89, 583-590. NIETHAMMER, J. (1969): Die Waldmaus, Apodemus sylvaticus (Linne, 1758) ın Afghanistan. Säugetierkdl. Mitt. 17, 121-128. NIETHAMMER, J.; Krapp, F. (1978): Handbuch der Säugetiere Europas. Wiesbaden: Akademische Verlagsges. pp. 476. PASTEUR, N.; PASTEUR, G.; BONHOMME, F.; CATALAN, J.; BRITTON-DAVIDIAN, J. (1987): Manuel d’electrophorese appliquee A la genetique des populations. Paris: Editions Techniques et Documentation. pp. 232. SOLDATOVIGC, B.; SavIC, 1.; SETH, P.; REICHSTEIN, J.; TOLKSDORF, M. (1975): Comparative karyologi- cal study of the genus Apodemus (Kaup, 1829). Acta Veterinaria, Beograd, 25, 1-10. SWOFFORD, D. L.; SELANDER, R. B. (1981): BIOSYS-1: aFORTRAN program for the comprehensive analysıs of electrophoretice data in population genetics and systematics. J. Hered. 72, 281-283. Zıma, J. (1984). Chromosomes of certain small mammals from Southern Bohemia and the Sumava Mts (CSSR). Folia Zool. 33, 133-141. ZımA, J.; Krar, B. (1984): Karyotypes of European Mammals II. Acta Nat. Acad. Sci. Bohemos- lovacae, Brno 18, 34-35. Authors’ addresses: JaANICE BRITTON-DAVIDIAN, MEHRNOUCH VAHDATI, FATIMA BENMEHDI and VALERIE NanceE Laboratoire de Genetique, Institut des Sciences de l’Evolution, USTL, Place E. Bataillon, F-34095 Montpellier Cedex 5, France; PAULE GRoS and HENRI CROSET, Laboratoire d’Eco-ethologie, Institut des Sciences de l’Evolution, USTL, Place E. Bataillon, F-34095 Montpelier Cedex 5, France; SVETOSLAV GUERASSIMOV, Zoology Institute, Bulgarıan Academy of Sciences, Bd Rouski 1, Sofia 1000, Bulgaria; CosTAs TRIANTAPHYLLIDIS, Arıstotelian Univer- sity of Thessaloniki, School of Sciences, Division of Genetics, Development and Molecular Biology, GR-54006 Thessaloniki, Greece Z. Säugetierkunde 56 (1991) 34-40 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Cranial infraspecific differentiation in Proechimys iheringi Thomas (Rodentia: Echimyidae) By LeırLa M. PessöA and S$. F. Dos Reıs Departamento de Zoologia, IBS, Universidade Federal do Rio de Janeiro and Departamento de Parasitologia, IB, Universidade Estadual de Campinas, Brazil Receipt of Ms. 6. 3. 1990 Acceptance of Ms. 8. 6. 1990 Abstract Proechimys iheringi Thomas ıs an echimyid rodent occurring in eastern Brazil from the states of Bahia to Säo Paulo. MOOJENn (1948) recognized sıx subspecies of P. iheringi primarily on the basis of clinal variation in the number of cheekteeth countertfolds. In this study we analyzed infraspecific differentia- tion in 13 morphometric cranıal characters ın three populations assıgnable to three subspecies of P. iheringi. Cranıal dımensions vary clinally increasing from north to south and morphomerric differ- entiation is correlated with geographic distance. The congruence between the clinal variation in cheekteeth counterfolds and cranıal traits indicates that the trinomial nomenclature should not be applied to P. iheringı. Introduction Proechimys iheringi Thomas is an echimyid rodent which occurs ın eastern Brazil, ranging from Bahıa to Säo Paulo (MooJEn 1948). In a detailed analysıs of infraspecific differentia- tion ın P. iheringi MOOJEN (1948) detected varıation in several skull traits including incisive foramen, tympanıc bulla, mesopterigoid fossa, palatine foramen, and vomerine sheath. In spite of the varıation in these characters, MOOJEn (1948) relied prımarily on the number of cheekteeth counterfolds, which varies clınally increasing from north to south, to recognize sıx subspecies ın P. zheringi, namely P. ı. denigratus from Bahia, P. :. gratiosus, P. 1. paratus, and P. i. panema from Espirito Santo, P. 1. bonafidei from Rio de Janeiro, and P. :. iheringi from Säo Paulo (Fig. 1). The subspecific structure in P. zheringı ıs thus based on a trait that varıes on a cline and whose differentiation ıs correlated wıth geographic distance (MoojJeEn 1948). In this paper, we analyzed cranial varıation in three populations of P. iheringi assignable to the following subspecies: P. 1. denigratus, P. i. gratiosus, P. i. bonafıdei from the states of Bahia, Espirito Santo, and Rio de Janeiro, respectively. The primary objective of thıs study was to determine whether the pattern of differentiation in cranıal quantitative traits is congruent with the clınal varıation in cheekteeth counterfolds and to address the question of recognition of infraspecific units ın P. iheringı. Material and methods A total of 54 specimens of P. iheringi available in the mammal collection of the Museu Nacional (Rıo de Janeiro) was examined in this study. All specimens were classıfied to one of the 10 age categories defined by Parron and RoGers (1983) for P. brevicanda on the basıs of tooth eruption and occlusal surface wear criteria. This procedure was employed in order to control the ontogenetic source of variation, and 42 specimens from age classes 8-10 were selected for the analysıs of geographic varıation because they were adults by the criteria of Patron and RocGers (1983). The specimens analyzed in this study represent samples collected at the following localities: Ilheus, state of Bahia (13° 01’ S, 40°01’ W; n = 16), Santa Tereza, state of Espirito Santo (19° 55’ S, 40°36’ W; U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0034 $ 02.50/0 Cranial infraspecıfic variation in Proechimys iheringi Thomas 35 Fig. 1. Ranges for the subspecies of Proechimys iheringi Thomas according to MooJEn (1948). The stars denote localıty samples analyzed in this study n = 18) and Teresöpolos, state of Rio de Janeiro (22° 26’ S, 42° 36’ W; n = 8). Sexes were pooled in the analysıs of geographic varıation to increase sample sizes. Twelve cranıal measurements defined in Parron and RoGers (1983) in addition to one mandibular measurement were taken with electronic digital calipers accurate to 0.01 mm, as follows: palatal length A (PL), zygomatic breadth (ZB), nasal length (NL), interorbital constriction (IC), rostral breadth (RB), diastema length (DL), rostral depth (RD), skull length (SL), basal length (BL), rostral length (RL), maxillary breadth (MB), pospalatal length (PP), and mandibular length (ML) (Fig. 2). Cranial character variation ın P. iberingi was analyzed by univariate and multivariate procedures (Sokar and ROHLF 1981; NEFF and Marcus 1980). Cranial characters were tested for significant differences among localities by univariate analysis of variance (AnovA). Significant characters were tested for maximally non-significant subsets of means employing Ryan-Einot-Gabriel-Welsch (REGWF) test on the main effect represented by the variable localıty. 36 Leila M. Pessöa and S. F. dos Reis Fig. 2. Thirteen measurements taken on the skulls of Proechimys iheringi Thomas (see text for explanation of measurement abbreviations) The pattern of variation in multivarıate character space was analyzed by principal components analysis (NEFF and Marcus 1980) and size-independent canonical discriminant analysis (STRAUSS 1985). The first pooled among-group principal component was used to study size variation among populations of P. iheringi. Scores derived from the fırst principal component were used as a measure of individual overall cranıal sıze and were tested by unıvariate anovA for heterogeneity among the populations. Size-independent canonical discriminant analysis was employed to analyze patterns of discrimina- tion and ordination among the P. iheringi populations. This procedure removes the effect of size variation within groups by performing canonical discriminant analysıs on the residuals obtained from the regressions of each log-transformed character separately on the first pooled within-group principal component (STRAuss 1985). This procedure was employed because P. iheringi shows post-ontogenetic growth (indeterminate growth) that generates size variation within populations (PessöA 1989) that may confound the analysıs of geographic varıation (THORPE 1983). Scores derived from canonical discriminant analysis were plotted to assess the pattern of ordination and discrimination among the populations of P. iheringi. Canonical loadings were expressed as bivarıate correlations calculated between original character values and scores on the canonical variates (Strauss 1985). Statistical analyses were performed using SAS-PC Version 6, the current microcomputer edition of the Statistical Analysıs System (SAS Institute, 1988). Cranial infraspecıfic variation in Proechimys iheringi Thomas 97. Results All cranial characters, except for palatal length and diastema length, increase in mean size from north to south, i.e. from the state of Bahia to the state of Rio de Janeiro (Table 1). Univariate analysis of variance indicates that all cranial traits differ significantly in the three populations of P. iheringi (Table 1). Nevertheless, the pattern of inter-locality differentia- tion in cranial traits is not uniform as indicated by the REGWF procedure (Table 1). Cranial Table 1. Standard statistics for 13 cranial characters (in mm) in three populations (BA, ES, and RJ) of Proechimys iheringi Thomas Character Palatal length Zygomatic breadth Nasal length Interorbital constriction Rostral breadth Diastema length Rostral depth Skull length Basal length Rostral length Maxillary breadth Mean (SD) ES 15.82 (0.79) BA 24.02 (1.21) BA 15.91 (0.98 ) BA 11.06 (0.57) BA 7.03 (0.58) ES 10.16 (0.80) ) ) ) BA 9.36 (0.52 BA 46.27 (1.81 BA 34.74 (1.67 Mean (SD) BA 16.30 (0.79) ES 25.32 (1.12) ES 17.47 (0.99) ES 11.53 (0.96) ES 7.36 (0.58) RJ 10.98 (0.63) ES 10.05 (0.83) ES 49.89 (2.31 ES 34.84 (1.55 BA 19.39 (0.99) BA 8.17 (0.42) ES 8.45 (0.88 Mean (SD) RJ 16.80 (0.76) RJ 26.08 (1.18) RJ 19.33 (1.27) RJ 12.47 (0.55) RJ 8.26 (0.43) BA 11.15 (0.78) RJ 10.87 (0.63) RJ 53.00 (2.66) RJ 37.22 (1.57) RJ 23.28 (1.35) RJ 9.09 (0.51) E A) 9.58 28.41 9.50 12.97. 7.38 18:24 26.87 7.30 22.26 5.28 B 0.0201 0.0004 0.0001 0.0005 0.0001 0.0020 0.0001 0.0001 0.0022 0.0001 0.0096 ) ) ES 21.43 (1.71) ) ) 27.20 7.87 0.0001 0.0014 Pos-Palatal length Mandibular length BA 2.72 (0.75) BA 23.87 (1.23) ES 23.40 (0.94 ( RJ 24.58 (1.26) ES 24.70 (1.38) ( RJ 26.20 (1.54) Statistics given are mean, standard deviation, F-value of an analysıs of varıance and associated probability levels (P), and results of REGWF analysıs. Lines below population means connect nonsignificant subsets. BA = Bahia, ES = Espirito Santo, and RJ = Rio de Janeiro. characters such as interorbital constriction, rostral breadth, basal length, maxillary breadth, and mandibular length are not significantly different ın the populations of Bahia and Espirito Santo, although individuals in both populations are significantly smaller than those ın the population from Rio de Janeiro. The three populations differ statistically in several cranıal traits including nasal length, rostral depth, skull length, rostral length, and pos-palatal length. Palatal length, zygomatic breadth, and diastema length have unique patterns of varıiation among the three populations (Table 1). In order to analyze size varıation among populations of P. ıheringi in multivariate character space, the first pooled among-group principal component (PAGPc-1) was extracted from the covarıance matrix of log-transformed character values. PAGPc-1 can be interpreted as a general sıze factor since all vector coefficients are positive and have significant correlations with log-transformed character values (Strauss 1985) (Table 2). The scores from principal components analysis for the individuals in the three populations of P. iheringi can then be used as a measure of multivariate cranial size. Mean score values increase from north to south in the populations of Bahia (-0.0729), Espirito Santo (0.0056), and Rıo de Janeiro (0.1163) and these mean values were shown to be highly significant different by an univariate anovA (F = 16.48; P < 0.0001). The pattern of ordination and discrimination of P. iheringi populations was assessed by 38 Table 2. Principal component and canonical variate loadings for 13 cranial morphometric Leila M. Pessöa and S. F. dos Reis characters in Proechimys iheringi Thomas Character PAGPC-1 Palatal length Zygomatic breadth Nasal length Interorbital constriction Rostral breadth Diastema length Rostral depth Skull length Basal length Rostral length Maxillary breadth Pos-Palatal length Mandibular length Percent of variance 0.160 0.212 0.380 0.134 0.339 0.178 0.357 0.285 94195 0.401 0.306 0.242 0.247 CV-1 0.845** -0.083ns -0.650** -0.652** —0.053ns 0.966” —-0.253ns -0.764** 0.590** -0.709** O.211ns -0.667** 0849 89.79 CV-2 0.06/ns 0.416” 0.238ns 0.205ns 0.341” 0.002ns —0.116ns —-0.078ns 0.201ns —0.132ns —0.050ns —0.045ns —0.153ns 10.21 explained 737 Canonical varıate loadıngs are expressed as vector correlations between log-transformed character values and canonical scores. PAGPC-1 is the pooled among-group first principal component. r is the Pearson product-moment correlation coefficient between cranial characters and the first principal component. * P<0.05; ** P<0.0001; n = non significant SVi Fıg. 3. Scatterplot of individual scores from a size-independent canonical discriminant analysis from three populations of Proechimys iheringi. Bahia (triangles), Espirito Santo (circles), and Rio de Janeiro (squares) sıze-independent canonical discriminant analysıs. The first canonical varıate (CV-1) ex- plains 89.79 % of the total among-group varıation while CV-2 accounts for the remaining 10.21 %. The plot of canonical varıate scores shows that the three populations of P. iheringi occupy different positions in the reduced space of canonical varıates (Fig. 3). The population from Bahıa is discriminated from the populations from Espirito Santo and Rio de Janeiro along CV-1, whereas individual scores for the latter populations have non- overlapping distributions along CV-2. Figure 3 also indicates that the populations from Espirito Santo and Rıo de Janeiro, which are the geographically closest, are also mor- phometrically more sımilar. These populations are more differentiated morphomertrically from the population of Bahia which is geographically farther. The vectors of correlations between canonical varıates and original log-transformed character values indicate that the population from Bahia differs from those of Espirito Santo and Rio de Janeiro ın most cranial measures of length (Table 2). Canonical variate 1 correlations indicate a contrast between palatal length, diastema length, basal length, and mandibular length with positive significant correlations and nasal length, interorbital Cranial infraspecific varıation in Proechimys iheringi Thomas 39 constriction, skull length, rostral length, and pos-palatal length with negative significant correlations. On the other hand, the population from Espirito Santo differs from that of Rio de Janeiro in zygomatic breadth with a negative significant correlation and rostral breath with a positive significant correlation (Table 2). Discussion The univariate statistical analysis of cranıal character varıation did not produce a consistent pattern of inter-locality population differentiation in P. iheringi, although most characters vary in a cline increasing in size from north to south. The lack of consistency ın character trends observed for P. iheringi ın this study is acommon result, whenever morphomertric characters are analyzed univarıately (BAKER 1980; THORPE 1983; MAcEDO and MARES 1987). The multivariate procedures employed produced a much clearer picture of the nature and extent of inter-locality ditferentiation in P. zheringi. Multivarıate cranıal size, esti- mated by mean score values derived from principal components analysis, increases from north to south confirming the clıne observed for most cranıal morphometric traits ın P. iheringi in the univariate analysıs. This clıne follows the same direction of the gradient in cheekteeth counterfolds observed by MoojJEn (1948). The pattern of ordination of P. iheringi populations in the space of canonıcal varıates further indicates a correlation between morphometric and geographic distance confirming MooJeEn’s (1948) observations based on qualitative arguments. The analysıs of geographic differentiation in ?. iheringi reported in thıs paper ıs based upon population samples representing three subspecies among the sıx forms recognized by MoojJEn (1948). We believe nevertheless that our findings are representative of the overall pattern of varıatıon ın P. ıheringi since the three subspecies we analyzed are distributed over most of the range of thıs species (Fig. 1). Our results, in addition to MOOJEN’s (1948) findings, indicate that the varıation in P. iheringi ıs geographically structured in a clıne of increasing cranıal dimensions and number of cheekteeth counterfolds from northern to southern populations. The recognition of subspecies on the basıs of clinal varıation, as is the case for P. iheringi, has been criticızed primarily due to the continuous nature of the variation expressed ın a cline (BARROWCLOUGH 1982; THORPE 1987). The validity of the recognition of subspecific units in P. iheringi can be questioned since the application of trinomials has been considered suitable to describe character varıatıon that do not sımply form clines but rather diagnoses groups of populations indicating the existence of inde- pendent infraspecific units (BARROWCLOUGH 1982; THORPE 1987; SMITH and PATTON 1988; PATTon and SMITH 1989). Our preliminary study indicates that, on the basis of the pattern of cranıal varıation, the use of the subspecific nomenclature is not justified for P. iheringi. Nevertheless, other character systems should be surveyed to assess the nature and structure of variation in order to understand the process of differentiation in this species and determine whether independent evolutionary units (sensu SMITH and PATToN 1988; PATTON and SMITH 1989) should be recognized for P. iheringı. Acknowledgements We thank Drs. A. S. Age and I. Sazıma for critically reading the manuscript and contributing to its improvement. Dr. U. Caramaschi kindly allowed the examination of museum specimens under his care. We are indebted to M. F. PEssöA for drawing the skulls and J. A. DE OLıveira for making the photographs. This study was supported by grants from CNPq (402265/87.4.20), FAPESP (88/2227- 4, 89/0772-1, and 89/3405-0), and FAP (503/89). Work by L.M.P. and S.F.R. was partially supported by graduate and research fellowships from CNPg, respectively. 40 Leila M. Pessöa and S. F. dos Reis Zusammenfassung Intraspezifische Schädelvariabilität bei Proechimys iheringi Thomas (Rodentia: Echimyidae) Schädel von Igelratten (Proechimys iheringi) aus drei Populationen, die drei der sechs beschriebenen Unterarten dieser Art zuzuordnen sind, wurden in 13 Maßen verglichen. Danach nimmt die Schädelgröße von Norden nach Süden zu. Die allgemeinen morphometrischen Abstände zwischen den Populationen sind mit ihren geographischen Abständen korreliert. Da sich auch die Zahl der Schmelzfalten in gleicher Richtung klinal ändert, läßt sich eine Gliederung von Proechimys iheringi in Unterarten mit diesen Merkmalen nicht rechtfertigen. Literature SAS Institute Inc. (1988): SAS/STAT User’s Guide, Release 6.03 Ed. Cary, NC. BAkKER, A. J. (1980) Morphomertric differentiation in New Zealand populations of the house-sparrow (Passer domesticus). Evolution 34, 638-653. BARROWCLOUCH, G. F. (1982): Geographic varıation, predictiveness, and subspecies. Auk 99, 601-603. Mac£no, H. R.; Mares, M. A. (1987): Geographic varıation in the South American cricetine rodent Bolomys lasiurus. J. Mammalogy 68, 578-594. MOoOoJEN, J. (1948): Speciation ın the Brazilian spiny rats (Genus Proechimys, Family Echimyidae). Univ. Kans. Publ., Mus. Nat. Hist. 1, 301-406. NErr, N. A.; Marcus, L. F. (1980): A survey of multivarıate methods for systematics. New York: privately published. PATTon, J. L.; ROGERs, M. A. (1983): Systematic implications of non-geographic variation in the spiny rats genus Proechimys (Echimyidae). Z. Säugetierkunde 48, 363-370. PaTTon, J. L.; SMITH, M. F. (1989): Population structure and the genetic and morphologic divergence among pocket gopher species (Genus Thomomys). In: Speciation and its Consequences. Ed. by D. OTTE and J. A. EnDLER. Sunderland: Sinauer Ass. 284-306. Pessöa, L. M. (1989): Diferenciagäo cranıana infraespecifica em Proechimys iheringi Thomas e P. dimidiatus (Guenther) (Mammalia: Rodentia: Echimyidae). Unpub. MSc diss., Univ. Fed. do Rio de Janeiro. SMITH, M. F.; Parron, J. L. (1988): Subspecies of pocket gophers: Causal basis for geographic differentiation in Thomomys bottae. Syst. Zool. 37, 163-178. SokaL, R. R.; ROHLEF, F. ]J. (1981): Biometry. 2nd. ed. San Francisco: Freeman. Strauss, R. E. (1985): Static allometry and varıation in body form in the South American catfısh genus Corydoras (Callichthyidae). Syst. Zool. 34, 381-396. THORBE, R. S. (1983): A review of numerical methods for recognising and analysıng racıal differentia- tion. In: Numercial Taxonomy. Ed. by J. FELSENSTEIN. Berlin: Springer Verlag. 404423. — (1987): Geographic varıation: A synthesis of cause, data, pattern and congruence in relation to subspecies, multivariate analyses and phylogeny. Bull. Zool . 54, 3-11. Authors’ addresses: LeıLa M. PessöA, Departamento de Zoologia, CCS, Universidade Federal do Rio de Janeiro, BR-21941, Rio de Janeiro, R], Brazil; $S. F. Dos Reıs, Depar- tamento de Parasitologia, IB, Universidade Estadual de Campinas, €7P.26109 BR-13081, Campinas, SP, Brazil Z. Säugetierkunde 56 (1991) 41-47 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Thermoregulation and torpor in African woodland dormice, Graphiurus murinus, following cold acclimation By G.T. H. Errison and J. D. SKINNER Mammal Research Institute, University of Pretoria, Pretoria, South Africa Receipt of Ms. 28. 11. 1989 Acceptance of Ms. 14. 9. 1990 Abstract Studied thermoregulation of 3 adult woodland dormice (Graphiurus murinus) following acclimation to 15 °C and 10 °C. The dormice entered hibernation under these conditions characterised by a fall ın body temperature to within 1°C of ambient, and a prolonged reduction in oxygen consumption measured over 24 h at 10 °C and 5 °C. Non-shivering thermogenic capacity exceeded that required to re-attain homeothermy by 20 %, but would permit these dormice to maintain normothermia down to -5 °C. The cycle of body mass displayed by G. murinus in the present study is characteristic of hibernating temperate dormice, but was not observed in specimens collected throughout the year in southern Africa. It therefore appears that woodland dormice may not enter hibernation throughout their distribution during winter, but may exhibit facultative torpor under adverse climatic conditions. Introduction Afrıcan woodland dormice, Graphiurus murinus, are small, arboreal rodents found throughout large areas of tropıcal and subtropical Afrıca (SMITHERS 1983). Although EISENTRAUT (1962) could not elicit torpor in G. murinus from West Africa (Cameroon), SMITHERS (1983) suggested that this species shows sıgns of lethargy during cold weather ın South Africa. Following the discovery of a torpıd woodland dormouse during rodent trapping by the authors outside Pretoria ın March 1987, the following study was initiated to ıinvestigate thermoregulation and torpor in G. murinus under controlled laboratory conditions. Material and methods Three Afrıcan woodland dormice (2 adult females, 1 adult male) were collected at Vaalkop Dam Nature Reserve (25°23’ 27°28’E) during April 1989. Within one week of capture the dormice were transferred to a windowless climate chamber (Specht Scientific, Johannesburg) at Pretoria University (25°45'S 28°12’E) wherein photoperiod and temperature could be accurately controlled. Throughout the study the photoperiod within the chamber was maintained at 10 h light: 14 h dark (lights on at 07h00 and off at 17h00) which closely resembled the photoperiod prevailing at Vaalkop Dam during mid-winter. The dormice were initially acclımated for 50 days at an ambient temperature (T,) of 15 °C followed by a further 50 days at T, = 10°C. The dormice were housed separately in standard laboratory rat cages containing sawdust, shredded paper and a 500 ml plastic beaker which served as a nest box. Water, sunflower seeds, rat cubes and rabbit pellets (Epol, Vereniging) were provided ad libitum, a diet occasionally supplemented with pieces of fresh apple. The body mass of each dormouse was recorded upon capture and subsequently at regular intervals throughout the study. In addition, rectal temperatures (T,) were measured between 07h00-13h00 on five occasions during acclimation to both T, = 15 °C and T, = 10 °C. T, was determined by inserting a chromel alumel thermocouple (K-Type: Fluke, Everett), attached to a digital thermometer, approxi- mately 2 cm into the rectum for a period not exceeding 15 sec. To avoid undue perturbation, body mass and rectal temperature were recorded at intervals of 3 days or more, and at all other times the dormice remained undisturbed within the climate chamber. Following 50 days acclimation at 10°C the oxygen consumption (VO;) of the dormice was measured as an indication of metabolic rate (MR) using an open-circuit system (as described by Depocas and Harr 1957, and Hırr 1971). The dormice were placed in perspex metabolic chambers U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0041 $ 02.50/0 42 G. T. H. Ellison and ]J. D. Skinner through which a flow of dried air (Silica Gel: Holpro, Johannesburg) was passed at a rate of 600 ml x min”!. The chambers were immersed in a constant temperature water bath (Labotec, Isando), and a chromel alumel thermocouple within the chamber was used for monitoring chamber tempera- ture (T.). Following full equilibration VO, was recorded using an Ametek S-3A/I oxygen analyser (Applied Electrochemistry, Pittsburgh) linked to a multi-channel data logger (Grant Instruments, Chambridge). The oxygen analyser was calıbrated before and after measurement, and VO, was corrected to standard temperature and pressure, dry (STPD). Average daily metabolic rate (ADMR) was measured over 48 h in large (13 I) metabolic chambers. The dormice were transfered to these chambers, together with their bedding and food, 20 h prior to the start of measurement, and were subsequently subjected to 24 h at T. = 10 °C followed by 24 hat T.=5°C. Aır leaving the chambers was connected to a three-way valve and time-switch (Air/Water 350 KPA: Ascoreg, Johannesburg), so that VO; from two dormice could be monitored simultane- ously every 30 min using a single oxygen analyser. At the end of measurement the mass and rectal temperature of each dormouse was recorded. Resting metabolic rate (RMR) was measured following one hour’s exposure to T.=5 °C and 10 °C within smaller metabolic chambers (700 ml). VO, was monitored over 30 min after which rectal temperature and body mass were recorded. Maximum oxygen consumption (VO,,,.x) following an injection of noradrenaline (NA) was used as an indication of non-shivering thermogenetic capacity (as described by HELDMAIER 1971). The dormice were anaesthetised with sodium pentobarbitone (70 mg/kg intraperitoneally. Sagatal: May- baker, Port Elizabeth) after which a chromel alumel thermocouple (J-Iype: Grant Instruments, Cambridge) was inserted deep into the rectum to measure T, and secured to the base of the tail with tape. Following a subcutaneous injection of 1.5 mg/kg NA (HELDMAIER 1971) the dormice were transfered to a small metabolic chamber (700 ml) at T. = 30°C and VO, and T}, were recorded simultaneously at one minute intervals for 60 min using the multichannel data logger. VOzmax and Tumax were taken as the highest values of VO, and T, recorded following NA injection. All results are presented as mean + one standard deviation unless otherwise stated. Results Body mass Following acclımation, the mean body mass of dormice in the present study entered a cycle comprising a rapıd period of weight gain followed by a slower sequence of weight loss. From a mean capture mass of 29.1 # 4.3 gthe dormice weighed 43.2 # 4.3 g after 25 days in captivity, an increase of almost 50 %. Following a gradual decline the dormice returned to capture mass by Day 65, after which they retained a stable mean body mass for the remainder of the study. Body temperature All the measurements of T, taken during acclımation to T, = 15 °C and 10 °C were pooled and displayed in two histograms (Fig. 1). Under both T, regimes the dormice were predominantly torpid, with the majority of T}, measurements fallıng at, or slightly above T,. At these Typs the dormice were stiff and displayed the lack of responsivity and coordination characteristic of deep torpor. In addition, the level of hypothermia recorded was dependant upon T,, being lower ae 17, - 1O,@zthanrarıT, — SS CHE measurements fell within the normothermic range for mammals (33 °C-36 °C) which suggests that the dormice were usually in torpor between 07h00 and 13h00 when T}, was recorded. Metabolism At 10°C the mean RMR of the dormice was 3.74 + 0.14 mlO,xg'xh (T, = 34.7+0.4°C) and increased slightly at 5°C to 3.99+0.15 mlO,xg"'xh’' (T, = 34.1 #0.1 °C). In contrast, VO, measured over 24 h at these temperatures remained at low levels for prolonged periods, punctuated with brief increases in VO, lasting between 1 h and 6 h (Fig. 2). Excluding these incidents of increased metabolism, the mean ADMR Thermoregulation and torpor in Graphiurus murinus 43 frequency frequency 10 15 20 25 30 35 1,(00) Fig. 1. Rectal temperatures recorded between 07h00 and 13h00 during acclimation to T, = 15°C and 10°C 12.19.@7(0.27.= 0.06 ml®©, Xen x<hr)) was only 7.3% RMR and at 5°€ (0.52 # 0.26 mlO,x g"'xh"') was 13.0 % RMR (Fig. 3). Despite the modest rise in ADMR following exposure to 5°C, the dormice exhibited a mean T, of 6.0 #0.3°C at the end of measurement. The male dormouse died under anaesthesia during the measurement of NST, whilst the females displayed a mean VO, max of 5.75 # 0.67 mlO,x g"' x h"! following NA injection, accompanied by mean Tpmax Of 40.6 # 1.0 °C. When compared to ADMR and RMR in Fig. 3 VO,,.. was found to exceed the level of heat production required to maintain normothermia at either 10°C or 5°C. However, according to HELDMAIER’s (1971) allometric equation relating VO,., to body mass, the VO, „ax recorded in the present study ıs approximately 90 % of that predicted for mammals of similar body sıze following cold (5 °C) exposure (VO,max predicted: 6.37 mlO,xg"'xh’'). Discussion The results of the present study clearly indicate that G. murinus from southern Africa is capable of spontaneous bouts of deep torpor when confronted with cold stress under sımulated winter photoperiod. Furthermore, these bouts of torpor exceeded 24h in duration, which suggests that G. murinus entered hibernation under these conditions. Hibernating dormice rarely arouse when challenged with declining temperatures and instead increase metabolic thermogenesis to maintain T, at non-lethal levels even as T, falls to O0 °C or below (WarHovD 1976; PAJunEn 1986). In the present study torpıd dormice displayed an increase in metabolism when T, fell from 10 °C to 5 °C, which indicates that torpor in G. murinus, in common with other dormice, ıs facultative and under strict thermoregulatory control. However, a consequence of this increase in metabolic rate is that an optimal T, exists for maximum energy savings during hibernation, as described for the garden dormouse, Eliomys quercinus (PAJunEN 1986). For G. murinus the optimal T, appears to fall between 10°C and 5°C which corresponds to the mean night-time temperature of around 5 °C prevailing during winter (June/July) at Vaalkop Dam (Pretoria Weather Bureau: Brits Met. Stn. 25°35’ S 27°49’ E 1939-1984). 44 G. T. H. Ellison and J. D. Skinner 7.0 DL VW, en 3.0) Y2:36.lgTpran des. 30€ 0825 2gn pr end 3 9 Time 12h00 18h00 00h00 06h00 12h00 18h00 00h00 06h00 12h00 (:[D) TR FZZZASLFTZZZZZAISIINEIISN Fig. 2. Oxygen consumption (VO,) of 3 Graphiurus murinus measured over 24 h at 10°C and 5°C Non-shivering thermogenesis is the dominant pathway for heat production in small mammals (BÖcKLER et al. 1988) although hibernators exhibit higher levels of NST than euthermic species of similar body sıze even in the absence of cold adaptation (JANsKY 1973). This enhanced capacity for NST amongst hibernators is believed to be associated with the heat required for arousal from hypothermia (ABBoTTs and Wang 1980). If we assume that the specific heat capacity of animal tissue is 3.4k]Jxg”' (CHArPELL and BARTHOLOMEW 1981) and that the calorific equivalent of oxygen is 20.1 JXmlO," (SCHMIDT-NIELSEN 1983) then rewarming a 31.9 g dormouse from T, = 6.1 °C to 34.4 °C (28.3 °C) in the present study would require 3.07 kJ or 152.74 mlO;. This compares with the total heat produced during NST of 5.75 mlO,xg"'xh! or 183.43 mlO,;xh, which exceeds that required for rewarming by 20 %. For this reason NST in G. murinus may be important not only for arousal from hypothermia, but also for the maintenance of homeothermy in the cold. In this context the lowest T, at which G. murinus can thermoregulate, using NST alone, can be calculated assuming that Newton’s law of cooling holds for this species as proposed by SCHOLANDER et al. (1950): (Thermal conductance below themoneutrality [C„] = VO>x[T,-T,]'). Based upon RMR and T}, measured at 5°C and 10°C, the mean Cr of dormicer in therpresent study, was 0145220078 Thermoregulation and torpor in Graphiurus murinus 45 7.0 150% 6.0 5.0 100% 40 100% 3.0 2.0 7.3% VO,(mlo,.g°1:h"1) ADMR RMR | 'ADMR RMR! | NA 102€ ea N Fig. 3. The relationship between average daily metabolic rates (ADMR) during torpor and resting metabolic rate (RMR) during normothermia measured at 10°C and 5°C in 3 Graphiurus murinus. As an indication of non-shivering thermogenic ability, the metabolic response to noradrenaline injection (NA: VO) of 2 G. murinus ıs displayed to the rıght. Vertical lines indicate + one standard deviation of the mean mlO,xg""xh"!x °C! with a mean normothermic T}), of 34.4 # 0.4 °C. Using NST to produce a VOymax of 5.75 mlO,x g!xh! these dormice could feasibly maintain a similar T; down to T,'s as low as -5 °C, which corresponds to the mean daily minimum temperature (-3 °C) recorded at Vaalkop Dam during winter (June/July: Pretoria Weather Bureau). The body mass cycle displayed by G. murinus in the present study ıs sımilar to that reported for Glis glis and E. quercinus during winter and following cold acclımation (Mrosovsky 1986; PAJunEn 1986). These dormice undergo a distinct increase in body mass prior to hibernation, during which up to 25 % of body mass ıs subsequently lost (JALLAGEAS and AsSSENMACHER 1986). In order to assess whether or not G. murinus displays similar body mass cycles ın nature the mass of 77 specimens collected throughout the southern African subregion were grouped by month of capture and presented in the Table. These specimens did not display a clear circannual cycle of body mass, which sugests that G. murinus does not naturally enter hibernation throughout southern Africa. Indeed, Mean body mass of 77 G. murinus collected throughout the southern African subregion (From the archives of the Transvaal Museum, Pretoria) Mar/Apr Sep/Oct 46 G. T. H. Ellıson and J. D. Skinner the cycle of body mass observed in the present study may have been an artefact of laboratory conditions as both short photoperiod (Könıs 1960) and abundant food (ROTHwELL and Stock 1986) have been reported to elicit increased body mass and subsequent hibernation in other species of dormice. However, under similar conditions, EISENTRAUT (1962) could not elicit torpor or hibernation from G. murinus haedulus from tropical West Africa which suggests that different populations of this species differ in their ability to display torpor and/or hibernate. Similar geographic differences have been reported for forest dormice (Dryomys nitedula) which hibernate for 6 months in Eurasia yet remain active throughout the year in Israel (NEVO and Amır 1964). It therefore appears that G. murinus from southern Africa differ from tropical populations in their ability to enter deep torpor/hibernation, although, like garden dormice in Morocco, thıs ability is probably only expressed when local climatic conditions demand (MORENO and DeLiıBes 1964). Acknowledgements The authors would like to thank BassıE POTGIETER for providing excellent technical assistance throughout the study and Amy and Nick BEZUIDENHOUT for their generous hospitality at Vaalkop Dam. InGo BUCHERT, GUsSTAV PETERS, WOLFGANG RAMDOHR and BRIGITTE WENHOLD kindly helped with german translation and assisted in collating the final manuscript. Collection data was provided by the Mammal Department at the Transvaal Museum, and clımatic information was supplied by the Weather Bureau in Pretoria. G.T.H.E. was supported by grants from the Foundation for Research Development, Department of National Education, and University of Pretoria. Zusammenfassung Thermoregulation und Torpor bei afrıkanischen Baumschläfern, Graphiurus murinus, nach Kaltebelastung Zahlreiche Arten der Gliridae aus den gemäßigten Klimazonen sind Winterschläfer. Über einen saisonbedingten Torpor bei Baumschläfern (Graphiurus murinus) aus den tropischen und subtropi- schen Gegenden Afrikas liegen nur Vermutungen vor. Drei erwachsene G. murinus wurden ım Herbst im südlichen Afrıka gefangen und ım Labor 50 Tage lang bei 15°C und 10°C sowie unter kurzer Lichtperiode gehalten. Sie gerieten dabei in Torpor. Die Körpertemperaturen sanken bis auf den Wert der Umgebungstemperatur. Die durchschnittliche Stotfwechselintensität fiel bei 10 °C und 5°C bis auf 73% bzw. 13,0% des Ruhewertes im normothermen Zustand. Die durch Noradrenalin induzierte zitterfreie Thermogenese übertraf die für das Erwachen aus dem Torpor benötigte Wärme. Die Bilche reagierten auf die Kältebelastung mit einer Gewichtszunahme und mit nachfolgendem Gewichtsver- lust. Tiere, die im Verlaufe des Jahres gefangen wurden, zeigten diesen Zyklus nicht. Vermutlich gibt es bei G. murinus nur unter sehr ungünstigten klimatischen Bedingungen Torpor. References ABBOTTS, B.; Wang, L. C. H. (1980): Seasonal thermogenic capacity in a hibernator, Spermophilus richardsonii. J. Comp. Physiol. B. 140, 235-240. BÖCKLER, H.; STEINLECHNER, $.; HELDMAIER, G. (1982): Complete cold subsitution of noradrenaline induced thermogenesis in the Djungarıan hamster, Phodopus sungorus. Experientia 38, 261-262. CHAPPELL, M. A.; BARTHOLOMEW, G. A. (1981): Activity and thermoregulation in the antelope ground squirrel Ammospermophilus leucurus in winter and summer. Physiol. Zool. 54, 215-223. Derocas, F.; HART, J. S. (1957): Use of the Pauling oxygen analyser for measurements of oxygen consumption of anımals in open-circuit system and in short-lag, closed-circuit apparatus. J. Appl. Physiol. 10, 388-392. EISENTRAUT, M. (1962): Wie verhalten sich verwandte Vertreter von heimischen Winterschläfern aus wärmeren Gebieten unter veränderten Temperaturbedingungen? Zool. Anz. 169, 429432. HELDMAIER, G. (1971): Zitterfreie Wärmebildung und Körpergröße bei Säugetieren. Z. vergl. Physiol. 73, 222-248. Hırı, R. W. (1972): Determination of oxygen consumption by use of the paramagnetic oxygen analyser. J. Appl. Physiol. 33, 261-263. JALLAGEAS, M.; ASSENMACHER, 1. (1986): External factors controlloing hibernation and related endocrine cycles in the edible dormouse, Glis glis. In: Endocrine regulations as adaptıve Thermoregulation and torpor in Graphiurus murinus 47 mechanisms to the environment. Ed. by I. AssEnMACHER and J. Boıssin. Paris: Editions du Centre National de la Recherche Scientifique. Pp. 363-369. Janskv, L. (1973): Non-shivering thermogenesis and its thermoregulatory significance. Biol. Rev. 48, 85-132. Könts, C. (1960): Einflüsse von Licht und Temperatur auf den Winterschlaf des Siebenschläters Glis g. glıs (Linnaeus 1766). Z. Morph. Okol. Tiere 48, 545-575. MORENOo, S.; DELIBES, M. (1982): Notes on the garden dormouse (Eliomys; Rodentia, Gliridae) of northern Morocco. Säugetierkdl. Mitt. 30, 212-215. Mrosovsky, N. (1986): Mirrors of the environment: the enigmatic dormouse and other hibernators. In: Endocrine regulations as adaptive mechanisms to the environment. Ed. by I. AssEnNMACHER and J. Boıssın. Paris: Editions du Centre National de la Recherche Scientifique. Pp. 331-341. NeEvo, E.; Amir, E. (1964): Geographic variation in reproduction and hibernation patterns of the forest dormouse. J. Mammalogy 45, 69-87. Pajunen, I. (1986): Ambient temperature dependance of physiological parameters during long-term hibernation in the garden dormouse, Ehomys quercinus. In: Endocrine regulations as adaptive mechanisms to the environment. Ed. by I. AssenMAcHER and J. Boıssın. Paris: Editions du Centre National de la Recherche Scientifique. Pp. 379-383. ROTHWELL, N. J.; STOCK, M. J. (1986): Spontaneous and experimental varıiations in body weight, food intake and metabolic rate in captive dormice (Glis glis). Comp. Biochem. Physiol. 84A, 141-147. SCHMIDT-NIELSEN, K. (1983): Anımal physiology: Adaptation and environment. 3rd ed. Cambridge: Cambridge University Press. Pp. 178. SCHOLANDER, P. F.; WALTERS, V.; Hock, R.; Irving, L. (1950): Body insulation of some arctic and tropical mammals and birds. Biol. Bull. 99, 237-258. SMITHERS, R. H. N. (1983): The mammals of the southern African subregion. Pretoria: Univ. Pretoria. WALHOVD, H. (1976): Partial arousals from hibernation in a pair of common dormice, Muscardinus avellanarius, (Rodentia, Gliridae), in their natural hibernaculum. Oecologia (Berl.) 25, 321-330. Authors’ address: G. T.H. ErLison, and J. D. SKınnEr, Mammal Research Institute, University of Pretoria, Pretoria 0002, Republic of South Afrıca Z. Säugetierkunde 56 (1991) 48-58 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Time budgets of Waterbuck (Kobus ellipsiprymnus) of different age, sex and social status By P. WıRrTZ and G. OLDEKOP Institut für Biologie I, Albert-Ludwigs-Universität, Freiburg, FRG, and Max-Planck-Institut für Verhaltensphysiologie, Seewiesen, FRG Receipt of Ms. 26. 3. 1990 Acceptance of Ms. 10. 8. 1990 Abstract Daytime budgets of waterbuck (Kobus elhpsiprymnus) of different age, sex, and social status were measured at Lake Nakuru National Park, Kenya. Adult and subadult females spent more time feeding than adult and subadult males. Inside territories, females spent more time feeding than outside of territories. Time spent for agonistic behaviour was highest ın young males. Satellite males spent less time for sexual behaviour than territory holders, but more time than bachelor males. Territory holders and satellite males spent the same amounts of time feeding - much more than bachelor males. Because the amount of forage ingested by ruminants is positively correlated with forage quality, this indicates that bachelor males were relegated to nutritionally inferior areas. Time budgets of males and females, site preferences of females, and faecal crude protein content of animals feeding at different sites (TomLınson 1979) all suggest that territories are high quality feeding areas and that the social system of waterbuck is a resource defence polygyny. Sex differences in time spent feeding at the same site probably reflect a fundamental difference in the behavioural programming of males and females: females of polygynous species are more likely to be “energy maximisers” than males. Recent evidence points to differences in levels of sexual hormones as a proximate cause of sex differences not only of sexual and agonistic behaviour but also of maintenance behaviour. Introduction Regardless of age, sex, or social status, all members of a species have the same amount of time available to spend every day. They can, of course, be expected to spend i it on different activities. In polygynous species, adult males will spend time to acquire females, whereas adult females will spend time to produce viable offspring (SCHOENER 1971; TRIVERS 1972; CLUTTON-BROocK et al. 1982; HoFFMANn 1983; OwEN SMITH 1984). As the amount of time per day is a finite value, spending more time for one activity must mean spending less time for one or several other activities. This study shows how the age classes, sexes, and holders of different social status of a polygynous mammal differ in allotting time to their activities during the day. Waterbuck (Kobus ellipsiprymnus) are antelopes similar in size to red deer; they occur throughout Africa south of the Sahara (Dorst and DAnDELOT 1970; HALTENORTH and Dirter 1977). The social structure of waterbuck living in Lake Nakuru National Park, Kenya, has been described previously (WırTz 1981, 1982). With an average of 30 waterbuck/km?, Lake Nakuru NP has by far the highest population density recorded for the species. Probably as a consequence of the high population density, only about 7 % of the adult males held a territory during the study period and half of the territory holders tolerated one or several additional adult males, “satellite males”, in the territory. Within the territory, satellite males were subdominant to the territory holder and participated in the defence of the territory. About 9% of the adult males acted as satellite males and the remaining 84% were bachelor males spending most of their time outside territories. Females moved in herds of changeable size and composition over home ranges encompas- U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0048 9.025070 Time budgets of Waterbuck 49 sing several territories and non-territorial areas. Most territories were located along the lake shore and along rıvers. During the day most females were inside territories (WIRTZ 1982). Material and methods Study area and recording technique The study area has been described previously (WırTrz 1982) in a paper that also gives the definitions of the age classes used: adult male, young male, juvenile male, adult female, subadult female, and calf. Both study years (1978 and 1979) were exceptionally wet years compared to long-term means (KurıLek 1975); see figure 2 in Wırrz (1982) for the monthly rainfall values. Data on time budgets were collected throughout the course of the two study years. Time budgets of the different age classes, sexes, and social classes (territory holder, satellite male, bachelor male) were measured between 7am to 7 pm. Groups of anımals were observed with binoculars (Leitz Trinovid 10x 40) from a Land-Rover or from tree hides. Observations were recorded on a tape recorder. The anımals were accustomed to cars and during observations would graze as close as 5 m from the Landrover; average observation distance is estimated to be about 50 m. Uninterrupted observations at the same site lasted from one to twelve hours, on average three hours. Data were collected using the „scan sampling“ technique (RoıLinson et al. 1956; ALTMAN 1974; Marrın and BATEsoN 1986). The observer would scan the group at regular intervals and record the momentary activity of each anımal seen. The proportion of records of a behaviour pattern is an approximation of the proportion of time spent performing this behaviour pattern. Inter-scan interval was five minutes. Large groups sometimes took longer to scan and the inter-scan interval was then set at ten minutes. All activities were classified into the eleven types described below (see WALTHER [1958] for a description of behaviour patterns such as Flehmen, and see Tomrınson [1980] for a description of the expressive behaviour of waterbuck in particular): l. Browsing Feeding on dicotyledons, such as shrubs or Acacıa leaves 2. Grazing Feeding on grasses 3. Standing Standing on all four legs; part of the time in this position is spent ruminating 4. Lying head up Lying on the ground with head raised above the ground; part of the time in this position ıs spent rumınating 5. Lying head down Lying with the head resting on the ground 6. Walking Moving forward at slow to moderate speed 7. Running Trotting or gallopıng 8. Agonistic Female butting her head into the flank of another female; male confronting another male with head raised high and horns tipped forward; male approach- ing another male in submissive low stretch posture wıth horns tilted back- wards; males fighting with interlocked horns; male galloping after another male 9. Sexual Male sniffing female, rubbing a female with its head, performing Flehmen or Laufschlag, running after a female, attempting to mount or mounting a female, copulating 10. Grooming Scratching the own body with hoof or horn, licking own body (allogrooming was only observed in mothers grooming their calves; this was recorded as “others”) 11. Others Drinking, defaecating and any other behaviour not mentioned above; also behaviour recorded as “unidentified” when an anımal was partially hidden by the vegetation (0.1% to 1.0 % of all records in the different age classes). The observations are biased in at least the following ways. Only groups of more than five animals were used for activity records. Among other things, this means that the data for territory holders apply only to territory holders that have females with them and not to territory holders without females. Observations were made only on groups in open grassland, open shrub, and open forest but not on groups in dense shrub and dense forest where anımals were difficult to see. Less than 6% of the waterbuck were recorded in these two types of habitat during counts of habitat utilization (WIrTZ and Kaiser 1988). Nevertheless, a slightly higher proportion of “browsing” would probably have been recorded if these two types of habitat had not been ignored. Recording the activity of each animal at the moment it is seen through the binoculars meant that an animal walking a few steps whilst grazing was recorded as “walking” rather than “grazing”. Bouts of ruminating are frequently interrupted and anımals seen at such moments were then recorded as “standing” or “lying head up” instead of “standing ruminating” or “lying ruminating”. We therefore decided not to report values for the two 50 P. Wirtz and G. Oldekop incomplete measures of ruminating activity and have included them in “standing” and “lying head up”, respectively. The data presented here describe daytime activity of waterbuck at Lake Nakuru National Park. Waterbuck are also active at night. For technical reasons, only few quantitative observations on nighttime activity could be taken. During several clear nights, approximately 20 hours of observations were made and there was no indication that the major patterns might be different from those recorded during daytime. Number of observations A total of 957643 observations were taken between 7:00 h and 19:00 h (observations per hour: 5513, 8630, 8480, 8516, 8305, 7855, 6845, 9048, 9906, 8812, 11422, 2411). The numbers of observations for the different waterbuck classes are given in Tables 1-4. The data for territorial males are from 39 individuals, those for satellite males from 36 individuals. More than 100 different individuals contributed to the observations for adult females and for bachelor males. Treatment of data and statistical procedure Numbes of observations were unequally distributed through the course of the day. For instance, relatively more observations were taken during hours when the anımals spent most of their time grazing; a daily mean from unweighted data would then result in an overestimate of this activity. For each hour, the number of observations of each activity was expressed as the percentage of all observations during this particular hour; the daytime mean was then calculated from these percentages (cf Figure as an example). When comparing two different waterbuck classes, their daytime means were reconverted into frequencies of observations on the basıs of the total numbers of observations. The proportions of an activity in the total time budget (e.g. number of observations “grazing” versus number of observations of all other activities) were then compared by chi-square test. When there were fewer than 100 observations for one of the classes compared during one of the hours of the day, this hour was exluded from the comparison. Daytıme budget of the “average waterbuck” was calculated by combining the data for all age classes in the proportions at which they were collected, i.e. by simply summing all observations per hour of all anımals, regardless of age, sex, and social status. The shorter the inter-scan interval, the better the estimate of the true amount of time spent for various activities (HARKER et al. 1954). On the other hand, very short observations intervals can result in measuring the same phenomenon repeatedly and producing “dependent data”, ı.e. an inflated sample size unsuitable for statistical analysis. Scanning a group every five minutes may lead to dependent data for some activities wıth very long bout lengths. A statistical comparison would then lead to unrealistic levels of significance. To be on the safe sıde, we here call a difference “significant” only if p < 0.001 (chi-square value > 10.83, 1 degree of freedom). Results The “average waterbuck” The “average waterbuck” spent 37 % of its daytime feeding (35.9 % grazing plus 1.1 % browsing), 15.5 % standing, 37.8 % lying, 6.7 % walking and less than 3 % for all other activities (Fig.). The two social behaviour patterns “agonistic” and “sexual” took up only 0.3 % each of the daytime activity of the “average waterbuck”. Lying head down also was a comparatively rare behaviour, taking only a few minutes at a time, as ıs typical for large bovids (BALcH 1955). The Figure shows the distribution of the major (most time consuming) activities throughout the day. There was amorning and an evening peak of the activities grazing and walking and a corresponding midday peak of lying. Solar radiation at Nakuru reaches a maximum at about 13-14 h (VArEscH1 1982, Fig. 3a), i.e. the time when the anımals spent the highest proportions lying head up (48 %) and lying head down (2.4 %). The distribution pattern of the activities throughout the course of a day as shown in the Figure was essentially the same for all classes of waterbuck. In the following, we therefore compare only the daily means of animals of different age, sex, and socıal status. Time budgets of Waterbuck 51 2.8% others 6.7% walking 1.5% Iying head down Q, 3 36.3% Iying ead up RN : RN RI % of time & 15.5% standing «0 | »0- 20 35.9% grazing 10 0) 1.1 % browsing TO SO 121312 1572168.172210019 time of day mean Fig. 1. Distribution of activities of the “average waterbuck” through the course of a day and daily means of the different activities. See text for numbers of observations per hour Sex differences Adult male - adult female Adult males and adult females differed markedly in their time budgets (Table 1). Whereas adult males spent 26.8 percent of their time feeding, adult females spent 39.7 percent - almost 1.5 times as long as males (chi-square 877, p < 0.00001). In contrast, adult males spent more time |yıng than adult females (chi-square 464, p < 0.00001). Note that the majority of these adult males are bachelor males and that the time budget of adult males ın general ıs therefore largely determined by the time budget of bachelor males. A compari- son of the time budgets of bachelor males, terrıtory holders, and satellite males ıs given below. Table 1. Comparison of time budgets (% of observations) of adult males, adult females, and subadult females (7-19 h) Adult male Adult female Subadult female Browsing Graszing Standing Lying head up Lying head down Walking Running Agonistie Sexual Grooming Others n observations 32 P. Wirtz and G. Oldekop Juvenile male - subadult female For juvenile males, we have insufficient data for the time 18 to 19h. A direct comparison with subadult females is therefore impossible. After truncating the data for subadult females at 18 h, acomparison of juvenile males and subadult females (Table 2) also shows a Table 2. Comparison of time budgets (% of observations) of subadult females, juvenile males, young males, and adult males (7-18 h) Subadult female Juvenile male Young male Adult male Browsing Grazing Standing Lying head up Lying head down Walking Running Agonistic Sexual Grooming Others n observations large difference in the time spent feeding: whereas juvenile males spent 28.3 % of their time feeding, subadult females spent 37 % (chi-square 89.6, p < 0.00001). In contrast, juvenile males spent more time standing than did subadult females (chi-square 122, p < 0.00001). Age differences Adult female - subadult female Adult females and subadult females closely resembled each other in the times spent for the different activities (Table 1). There were no obvious age differences in their time budgets. Those activities restricted to sexually mature females (e.g. suckling a calf or licking a calf) were recorded in the category “others” and took up less than one percent of the adult females’ time. Adult male - young male - juvenile male Young males spent less time lyıng (38.5 %) than adult males (47.7 %) (Table 2, chi-square 262, p < 0.00001). Instead, they stood longer, groomed longer and spent more time in agonistic interactions (chi-square > 36, p < 0.00001 in each case). Feeding time of young males (26.9 %) was slightly higher than that of adult males (24.7 %) (chi-square 19.1, p < 0.0001). Juvenile males spent even less time lyıng (32.3 %) and more time standing than young males (chi-square > 22, p < 0.00001 ın both cases). They also spent slightly more time feeding than young males, but this difference does not reach the significance level. Juvenile males spent approximately the same amount of time for agonistic behaviour as did adult males (chi-square 0.6), ı.e. much less than young males (chi-square 14.1, p < 0.0002). Thus, with increasing age, males tended to spend less time standing and feeding and more time |ying; time spent for agonistic interactions was highest for young males. Time budgets of Waterbuck 53 Social status Territory holder - Satellite male — Bachelor male Table 3 compares the time budgets of territory holders, satellite males and bachelor males. As there were insufficient data for bachelor males from 7 to 8 h and from 18 to 19 h, the data for the other two classes were also reduced to the time span 8 to 18h. Table 3. Comparison of time budgets (% of observations) of territory holders, satellite males, bachelor males, and adult females (8-18 h) Territory holder Satellite Bachelor Adult female inside territory Browsing 0.4 Grazing 2821 Standing 14.4 Lying head up 339 Lying head down Walking Running Agonistic Sexual Grooming Others n observations Territory holders and satellite males spent the same amounts of time feeding (28.5 % and 28.6 %); in contrast, bachelor males spent much less of their time feeding (19.7 %; chi-square > 88, p < 0.00001 for both comparisons). Bachelor males spent more than half (51.7 %) of their daytime hours lyıng head up or head down. Satellite males and territory holders spent much less time for this type of activity (38 % and 41.9 %; chi-square > 89, p < 0.00001 in both cases). Compared to territory holders, satellite males stood longer (chi-square 12.4, p < 0.0004) but spent less time lying; however, the latter difference does not reach the predetermined significance level (chi-square 7.7, p < 0.005). Bachelor males spent more time grooming themselves than did satellite males (chıi- square 32, p < 0.00001), which in turn spent more time grooming themselves than did territory holders (chi-square 43.5, p < 0.00001). Whereas territory holders spent 5.1 % of their time for sexual behaviour, bachelor males spent only a tenth of this time on sexual behaviour (chi-square 354.8, p < 0.00001). Satellite males spent less time on sexual behaviour than territory holders, but spent considerably more time on sexual behaviour than did bachelor males (chi-square 13.9, p < 0.0002 for comparison with territory holders, chi-square 130.1, p < 0.00001 for compari- son with bachelor males). Territory holders with and without Satellite males Table 4 compares the time budgets of territory holders when a satellite male was present and of terrıtory holders in the absence of a satellite male. Because of insufficient data for the time period of 18 to 19h in one of the classes, both data sets are truncated at 18 h. Territory holders without satellite males spent more time grazing, less time lying and more time for sexual activities but none of these differences reached the predetermined level of significance (6.95 > chi-square > 6.30, 0.02 > p > 0.005). 54 P. Wirtz and G. Oldekop Table 4. Comparison of time budgets (% of observations) of territory holders with satellites and territory holders without satellites and of adult females inside territories and adult females outside territories Territory holder Territory holder Adult female Adult female with satellite without satellite inside territory outside territory (7-18 h) (7-18 h) (8-12, 14-16, 17-18 h) (8-12, 14-16, 17-18 h) Browsing Grazing Standing Lying head up Lying head down Walking Running Agonistic Sexual Grooming Others n observations Adult females inside territories and outside territories During the day, most females were inside territories. Even though adult females were the largest class in the population (31 % of the waterbuck seen during monthly road strip counts were adult females, Wırrz and KAıser 1988), there were insufficient data for the time budgets of adult females outside territories for five of the twelve daytime hours. A comparison between adult females inside territories and adult females outside territories (Table 4) can therefore only be made for the following fragments of a day: 8-12, 14-16, and 17-18 h. When inside territories, adult females spent much more time grazing (41.1 %) than when outside territories (27.6 %) chi-square = 112, p < 0.00001). Outside the territories, adult females spent more time browsing (chi-square = 13.8, p < 0.0002) and grooming themselves (chi-square 323, p < 0.00001) than inside territories. Sex differences revisited : adult females - adult males, inside territories The time budgets of adult females and of adult males were compared in a previous section. However, the two data sets differ not only in the sex of the animals but also in the location where they were taken: the majority of the data for adult males are of bachelor males (i.e. from outside territories), and the majority of the data for adult females are from females inside territories. To eliminate the bias caused by differences in site of observation, the time budget of adult females inside territories has to be compared with that of adult males inside territories. Table 3 shows the time budget of adult females (between 8 and 18 h) ın comparison with that of territory holders and satellites. Inside territories, females still spent much more time feeding than dıd males (chi-square > 85 in both comparisons, p < 0.00001). The sex difference in time spent lyıng, however, is no longer significant when considering only anımals inside territories (chi-square 4.6 ın the comparison with territory holders, p < 0.05; chi-square 2.6 in the comparison with satellite males, p < 0.15). Adult females outside territories and inside territories differed conspicuously from adult males in spending almost no time on agonistic and sexual interactions (p < 0.00001 for all comparisons). Time budgets of Waterbuck 93 Discussion An understanding of the nutritional physiology of ruminants is essential for the interpreta- tion of their time budgets. The food intake of ruminants ıs limited by the time required to process the food in the rumen. They can ingest food only as fast as they can digest it. Better digestible forage moves faster through the digestive system and thus permits the anımal to take up more of it. Low digestibility of food cannot be compensated for by a greater food intake - on the contrary, food intake is reduced because the time required for processing is longer. Except for highly digestible food (Ammann et al. 1973), the amount of forage ingested and the time spent feeding are positively correlated with the quality of the forage (BLAXTER 1962; THORNTON and Mınson 1972; ARNOLD 1985). Waterbuck are “grass and roughage eaters” taking food of comparatively low digestibility requiring relatively long processing times (HoFMAnnN 1973). The social structure of Lake Nakuru waterbuck Inside territories, females spent more time feeding than outside territories. This indicates better quality of the forage conditions inside territories. Females are free to move between these areas and the observation that female density is higher inside than outside territories (WIRTZ 1982) suggests that territories are superior feeding sites. However, there could also be alternative and additional reasons for the female preference for territories (e.g. more frequent harassment by bachelor males outside territories). Adult males show sımilar site differences in feeding times: bachelor males spent less time feeding than territory holders and their satellite males. Waterbuck territories usually border on water and, to the human observer, the grass outside territories often appeared to be higher and dryer. Dry grass usually has a higher proportion of lignin and thus a lower digestibility (cf SIncLAIR 1975), which would cause longer processing times and lower rates of uptake. Protein content ıs generally acknowledged as a major determinant of nutritive value of forage for ruminants (SINCLAIR 1975; FEsTA-BIANCHET 1988 and references therein). Faecal crude protein content is closely correlated with dietary protein and has been used to assess forage quality in studies of domestic cattle (BREDoN et al. 1963) and wild ungulates (see references in FESTA-BIANCHET 1988). In a Rhodesian population of waterbuck, the faeces of territory holders and of adult females had a higher crude protein content than the faeces of bachelor males (TomLınson 1979). This Rhodesian population of waterbuck had the same social structure as Nakuru waterbuck suggesting that TomLinson’s (1979) results would also apply to Nakuru waterbuck. Site-dependent differences ın faecal crude protein of males, site preference of females, site-dependent foraging times of females, and site- dependent foraging times of males all suggest that territories are high quality feeding areas. Bachelor males, being excluded from territories, are probably relegated to inferior feeding areas. The spatial distribution of resources determines the distribution of receptive females, which in turn determines the distribution of males and hence the nature of the mating system (see EMLEn and OrınG 1977 for a general discussion and classification of social systems, and GEısT 1974 for the relationship of ecology and social evolution of ungulates in particular). Territory holders defend high quality feeding sites preferred by females and mate with the females coming to these areas. In the terminology of EmLEn and ORING (1977), the mating system of waterbuck is a “resource defence polygyny”. (Unfortunately, in their influential paper on the evolution of mating systems, EMLEN and Orıng [1977], erroneously refer to waterbuck as an example of “female defence polygyny”.) In contrast to females (see below), the fitness of males of a polygynous species, such as waterbuck, is probably determined by non-foraging activities, in particular efforts to acquire females. Because comparatively little time is spent in aggressive encounters, the sex 56 P. Wirtz and G. Oldekop difference in aggressive behaviour, likely to be very important in terms of energy and mortality risk, is only weakly expressed in the time budgets. Territory ownership did not cause a reduction in feeding time. On the contrary, territory owners spent more time feeding, on more nutritious food, than did bachelor males. The costs of territoriality are only weekly expressed in the time budgets by slightly higher percentages of walking and running. The differences ın energy budget and risk are probably more important. Sex differences Even when feeding at the same site and presumably on forage of the same nutritional value, females spent much more time on energy intake than males - a difference already apparent in subadult anımals. With increasing age of males (from juvenile male via young male to adult male), this sex difference in maintenance behaviour became more and more pro- nounced. SPINAGE (1968) observed the activity of three individually known adult female waterbuck and two individually known adult males (one of them a territory holder) in the Queen Elizabeth Park, Uganda, for continuous periods of up to three consecutive days. The absolute values of his data are not directly comparable with those of the present study, because an anımal walking a few steps from one grazing site to another was recorded as “walking“ at Nakuru but recorded as “grazing” by SPinAGE. Nevertheless, SPInAGE’s data sımilarly show sex-specific differences in maintenance behaviour: the three females spent more time feeding than the two males. Female waterbuck are probably “energy maximisers” in the terminology of SCHOENER (1971), 1.e. their fitness is determined mainly by the amount of energy acquired. That females spend more time feeding than males is a general finding for all ungulates studied (cf review by BunnELL and GILLINGHAM 1985) and also for several other anımal species (e.g. Horrman 1983). This difference is probably a fundamental behavioural and physiological difference between the sexes, rather than an epiphenomenon of other factors such as differences in bite sıze. JEWELL (1986) showed that in a feral population of sheep at St. Kilda Island, Scotland, rams spent much less time grazing than ewes. Interestingly, castrated males spent almost as much time grazing (71 %) as females. The likely basıs for these differences lies in low testosterone levels of castrated males and the accompanying shift to a female type of maintenance behaviour. The proximate cause for the sex difference not only in aggressive and sexual behaviour but also in maintenance behaviour of ruminants could thus be a difference in levels of sexual hormones. Acknowledgements During most of the activity recordings, ANDREW BARTON worked with P. WIRTZ as a voluntary field assistant at Lake Nakuru National Park; he collected part of the data. The Government of the Republic of Kenya granted research permit OP 13/7C245/4. Prof. Dr. W. WICKLER and the Max- Planck-Gesellschaft financed the study. BARBARA KNAUER converted several hundred data sheets into computer files. Profs. V. GEisT and R. R. HOFMANN commented on an early draft of the manuscript. Many thanks to all of them. Zusammenfassung Zeitbudgets bei Wasserböcken (Kobus ellipsiprymnus) von unterschiedlichem Alter, Geschlecht und Sozialstatus Im Nakuru Nationalpark, Kenia, wurde gemessen, wie sich Wasserböcke von unterschiedlichem Alter, Geschlecht und Sozialstatus die Tageszeit einteilten. Weibchen verbrachten mehr Zeit mit Fressen als Männchen und innerhalb von Territorien mehr als außerhalb. Bei jungen Männchen war der Anteil Zeit für kämpferische Interaktionen am höchsten. Satellitenmännchen verbrachten zwar weniger Zeit mit sexuellen Aktivitäten als Territoriumsbesitzer, aber mehr als Junggesellenmännchen. Territoriumsbesitzer und Satellitenmännchen wendeten gleich viel Zeit für Fressen auf — wesentlich mehr als Junggesellenmännchen. Da bei Widerkäuern Nahrungsqualität und Nahrungsaufnahme negativ korrelieren, deutet dies an, daß die Junggesellenmännchen in Gebiete schlechterer Furterquali- Time budgets of Waterbuck >, tät abgedrängt wurden. Sowohl die Zeitbudgets von Männchen und Weibchen, als auch die Ortsbe- vorzugung der fressenden Weibchen und der Proteingehalt von Kotproben (Tomrınson 1979) deuten an, daß Wasserbock-Territorien Orte hoher Futterqualität sind, und daf das Sozialsystem von Wasserböcken als Fortpflanzungsterritorialität bezeichnet werden kann. Geschlechtsunterschiede im Zeitaufwand für Nahrungserwerb reflektieren wahrscheinlich funda- mentale Unterschiede im Verhaltensprogramm von Männchen und Weibchen: bei polygynen Arten sind Weibchen wahrscheinlicher “Energiemaximierer” als Männchen. Neuere Untersuchungen deuten an, daß unterschiedliche Niveaus von Sexualhormonen der proximale Grund nicht nur von Geschlechtsunterschieden im Sexualverhalten und im kämpferischen Verhalten sind, sondern auch der proximale Grund von Geschlechtsunterschieden im Ernährungsverhalten. Literature ALTMan, J. (1974): Observational study of behavior: Sampling methods. Behaviour 49, 227-267. AMMAann, A. P.; Cowan, R. L.; MOTHERSHEAD, C. L.; BAUMGARDT, B. R. (1973): Dry matter and energy intake in relation to digestibility in white-tailed deer. J. Wildl. Manage. 37, 195-201. ARNOLD, G. W. (1985): Regulation of forage intake. In: Bioenergetics of wild hervibores. Ed. by R.]. Hupson and R. G. WHITE, Boca Raton: CRC Press. pp. 81-101. BAccH, C. C. (1955): Slerep ın ruminants. Nature 175, 940-941. BLAXTER, K. L. (1962): The energy metabolism of ruminants. London: Hutchinson. BREDoON, R. M.; HARKER, K. W.; MarRsHALL, B. (1963): The nutritive value of grasses grown ın Uganda when fed to Zebu cattle. J. Agr. Sci. 61, 101-104. BunnELL, F. L.; GILLINGHAM, M. P. (1985): Foraging behaviour: the dynamics of dining out. In: Bioenergetics of wild herbivores. Ed. by R. J. Hupson and R. G. WHrtE. Boca Raton: CRC Press. pp. 53-79. CLUTTON-BRocK, T. H.; Guingss, F. E.; ALson, $. D. (1982): Red Deer. Behavior and ecology of two sexes. Edinburgh: University Press. Dossr, J.; DANDELOT, P. (1970): A field guide to the larger mammals of Africa. London: Collins. EmLen, S. T.; Orınc, L. W. (1977): Ecology, sexual selection and the evolution of mating systems. Science 197, 215-223. Festa-BIancHET, M. (1988): Seasonal range selection in bighorn sheep: conflicts between forage quality, forage quantity, and predator avoidance. Oecologıa 75, 580-586. Geist, V. (1974): On the relationship of social evolution and ecology in ungulates. Amer. Zool. 14, 205-220. HALTENORTH, T.; DILLER, H. (1977): Säugetiere Afrıkas und Madagaskars. München: BLV Ver- lagsges. en K. W.; TavyLor, J. L.; RorLınson, D. H. L. (1954): Studies on the habits of zebu cattle. 1. Preliminary observations on grazing habits. J. Agr. Sci. 44, 193-198. Horrman, $. G. (1983): Sex-related foraging behavior in sequentially hermaphroditic hogfishes (Bodianus spp.). Ecology 64, 798-808. HOFMANN, R. R. (1973): The Ruminant Stomach. Nairobi: East African Literature Bureau. JEwELL, P. (1986): Survival in a feral population of primitive sheep on St. Kilda, Outer Hebrides, Scotland. Nat. Geogr. Research 2, 402-406. KUTILEK, M. J. (1975): Feeding strategies and foraging impact of non-migratory African antelope. Ph. D. thesis, Michigan State University. MARTIN, P.; BATEson, P. (1986): Measuring behaviour. Cambridge: University Press. OWEN-SMITH, N. (1984): Spatial and temporal components of the mating systems of kudu bulls and red deer stags. Anim. Behav. 32, 321-332. Rorınson, D. H. I.; HARKER, K. W.; TAYLoR, ]. J.; LEECH, F. B. (1956): Studies on the habits of zebu cattle. IV. Errors associated with the recording technique. J. Agr. Sci. 47, 1-5. SCHOENER, T. W. (1971): Theory of feeding strategies. Ann. Rev. Ecol. Syst. 11, 369-404. SINCLAIR, A.R. E. (1975): The resource limitation of trophic levels in tropical grassland ecosystems. J. Anım. Ecol. 44, 497-520. SPINAGE, C. A. (1968): A quantitative study of the daily activity of the Uganda Defassa Waterbuck. E. Afr. Wildl. J. 6, 89-93. THOoRNTonN, R. F.; Mınson, D. J. (1972): The relationship between voluntary intake and mean apparent retention time ın the rumen. Aust. ]J. agric. Res. 23, 871-877. Tomuinson, D. N. $S. (1979): The feeding behaviour of waterbuck in the Lake McIlwaine Game Enclosure. Rhod. Scı. News 13, 11-14. — (1980): Aspects of the expressive behaviour of the waterbuck Kobus ellipsiprymnus ellipsiprymnus in a Rhodesian game park. $. Afr. J. Zool. 15, 138-145. TRIVERS, R. L. (1972): Parental investment and sexual selection. In: Sexual selection and the descent of man. Ed. by B. Camper. Chicago: Aldine. pp. 136-179. 58 P. Wirtz and G. Oldekop VARESCHI, E. (1982): The ecology of Lake Nakuru (Kenya). III. Abiotic factors and primary production. Oecologia 55, 81-101. WALTHER, F. (1958): Zum Kampf- und Paarungsverhalten einiger Antilopen. Z. Tierpsychol. 15, 340-380. WırTz, P. (1981): Territorial defence and territory take-over by satellite males in the waterbuck Kobus ellipsiprymnus (Bovidae). Behav. Ecol. Sociobiol. 8, 161-162. — (1982): Territory holders, satellite males, and bachelor males in a high density population of waterbuck (Kobus ellipsiprymnus) and their associations with conspecifics. Z. Tierpsychol. 58, 277-300. WırTz, P.; KAISER, P. (1988): Sex differences and seasonal variation in habitat choice in a high density population of Waterbuck (Kobus ellipsiprymnus). Z. Säugetierkunde 53, 162-169. Authors’ address: Dr. PETER WIRTZ, Zoologisches Institut, Albertstr. 21a, W-7800 Freiburg, FRG Z. Säugetierkunde 56 (1991) 59-63 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 ZISSENSICTENIETPLETIEFKUNRZINDIEEEEIEIN.G Some remarks on size differences of northwest German game populations from Nebolithic to modern times By D. HEINRICH Institut für Haustierkunde, Universität Kiel, FRG Receipt of Ms. 20.7. 1990 Acceptance of Ms. 20.9. 1990 Starting-point of this consideration is the bone material from medieval Schleswig, excava- tion Schild (Vocer 1983). For a certain period during the Middle Ages Schleswig, sıtuated in the northern part of Schleswig-Holstein (Fig. 1), was one of the most important trade centres in Europe, i.e., from the 11°" century after the decline of nearby Haithabu until the middle of the 13'® century when Lübeck, situated 100 km southeastwards and finally the capıtal of the Hanseatic League, succeeded in its place (HoFFMmann 1981). At the excavation sıte nearly 112000 bones of mammals were found. Only 1350 of these bones, which ıs a small fraction of 1.2 %, were from wild mammals, the rest from domestic ones (HÜSTER 1990). This is typical for medieval sites (REICHSTEIN et al. 1980). According to the relative frequencies of the remains the most important game were the artiodactyl species red deer (n = 397), roe deer (n = 323) and wıld boar (n = 82), and further the brown hare (n = 312) and red fox (n = 97). Although the amount of bone material of Denmark No JO Schleswig RN (} Haithabu Scharstorf® Bischofswarder ® Lübeck® en Hitzacker Fig. 1. Schleswig-Holstein. Location of the medieval trade centres Haithabu, Schleswig, and Lübeck as well as some other medieval settlements U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5601-0059 $ 02.50/0 60 D. Heinrich these species is hardly satistactory, ıt was the basıs for size analyses, and was compared with corresponding bones not only from other early historic and prehistoric sites, but also with recent skeletal material (collection of Institut für Haustierkunde, University of Kiel). In roe deer and in wild boar no sıze differences are apparent between the Schleswig material and that of other medieval or neolithic sites. However, compared with recent skeletons from Schleswig-Holstein it is obvious that roe deer and wild boar today are significantly smaller than the individuals of these species identified for medieval Schleswig and for the other sites. In roe deer this is shown by the measurements ‘maximum length’ of radıus and metacarpus and ‘distal width’ of metacarpus (Fig. 2). This tendency - evident differences in size between pre- or early historic findings and recent skeletal material - ıs also true for red deer (Fig. 3). In any case the recent material is relatively small-sızed and corresponding differences in body size can be deduced. But the comparison of the Schleswig material with other subfossil bones has shown that red deer of the early medieval strongholds Scharstorf and Bischofswarder, situated in eastern Hol- stein, partly must have been even larger. This is shown by the measurement ‘maximum length’ of calcaneus, whereas similar differences cannot be established in phalanx 1, the second skeletal element analysed. RADIUS, maximum length (GL) MIDDLE AGES Schleswig —— 10 RECENT Northern Germany + —— 35 150 158 166 174 182 190 mm METACARPUS, maximum length (GL) MIDDLE AGES Schleswig —— 16 RECENT Northern Germany + + — 38 140 148 156 164 172 180 mm METACARPUS, distal width (Bd) MIDDLE AGES Schleswig — 18 3 settlements —— 11 NEOLITHIC 2 settlements — En — 6 RECENT Northern Germany + + —— 38 17 19 21 23 25 mm kin — u — 11x -SE +SE x Fig.2. Roe deer, Capreolus capreolus. Comparison of size between finds from Schleswig, varıous northwest German sites (Middle Ages: Haithabu — REICHSTEIN 1990; Lübeck Königstraße — PauL 1980; Hitzacker - WALCHER 1978; Neolithic: Hüde I- HüBner et al. 1988; Rosenhof — Nogıs 1975) and recent material; + 24 22,11 8&d; ++25 292,13 dd Size differences of game populations from Neolithic to modern times 61 PHALANX 1, maximum length of the peripheral half (GLpe) MIDDLE AGES Schleswig — 19 Hitzacker — 46 Scharstorf — ln 37 Bischofswarder —— 29 NEOLITHIC 5 settlements ern og 15 RECENT Schleswig- Holstein + —— 88 48 52 56 60 64 68 mm CALCANEUS, maximum length (GL) MIDDLE AGES Schleswig — a — 14 Hitzacker ——— en — — 14 Scharstorf — — 19 Bischofswarder ze 18 NEOLITHIC 5 settlements —— arm 11 RECENT Schleswig- Holstein** ——l —— 21 ——— nd nn nn ann Insben Iuper Er a en en [er FE Era] Tram ST ER ln RN TEE Er NN) TREE Fler TERRA EEN 104 110 116 122 128 134 mm Fig.3. Red deer, Cervus elaphus. Comparison of size between finds from Schleswig, various northwest German sites (Hitzacker - WALCHER 1978; Scharstorf -— HEınRıcH 1985; Bischofswarder — REICHSTEIN et al. 1980; Neolithic: Hüde I- HÜBnER et al. 1988; Bistoft - JoHmansson 1979; Süssau — Nosıs 1971; Rosenhof - Nosıs 1975, Stinthorst - GEHL 1976) and recent material; +7 292,4 dd; ++6 2 2,4 8d,1 sex? Corresponding analyses of brown hare and red fox dıd not yıeld such results. In the case of hare only very few measurements showed significant sıze differences, but always without a clear tendency. In the case of the fox sıze differences were found only in the mandibula, which is larger in recent specimens compared with those from Schleswig. In the analysıs of the large amount of material of red fox from Haithabu REIcHsTEIN (1990) obtained similar results. The clear dissimilarities that could be shown especially in red deer, roe deer and wild boar, concern only size but apparently not shape. Shape differences between single bones would point to dissimilarıties of the stature of animals. But when correlating two measurements of phalanx 1 anterior of the red deer by regression methods these bones of all populations converged to one straight line, which describes the allometrical relationship of the two measurements taken (RöHrs 1959). Looking at roe deer, however, no correlations at all could be found. This points to a large varıability ın this species. Such methods could not be employed in the analysıs of wild boar, because the quality of the material was too poor. This size reduction ın the artiodactyl species must be a modern phenomenon for one cannot see similar differences between the samples of the various early historic or prehistoric sites. This means that from neolithic to medieval times there were no changes 62 D. Heinrich which could bring about diminution or enlargement. Thus, it can be noted that the degree ot climatıc influences in the course of this long period was too low to alter the size of the species in the sense of Bergmann’s rule (climate-size-rule). Corresponding results espe- cıially for red deer and roe deer have been repeatedly shown, for instance by BoEssNEcK (1956), REQUATE (1956), REICHSTEIN (1974), and KRATOCHViL (1988). This phenomenon - no alterations in size over a long period followed by a sudden diminution in the recent past - can be explained by human influence on landscape and vegetation. Although man has modified the landscape by agrıcultural activities since centuries — e.g. especially the medieval clearıng of woodland for cultivation — the alterations caused by human activities are becoming more extensive and frequent especially in the last few decades. Here, one must not only see the modern cultivation, but also the development of settlements and the progress of traffic. Further, one must note that modern forestry is not suitable tor game. The same is true for modern hunting with its demand for trophies. Whereas these factors are essentially responsible for the diminution in red deer and probably also in wıld boar, the relatively small size of roe deer, however, presumably ıs only indirectly connected with them: According to KLEIN and STRANDGAARD (1972) in this species size ıs correlated with population density to a high degree, which in return is dependent on landscape and vegetation, cover, food value etc. The above-mentioned varıability of roe deer in shape must also be seen in this context (SÄGESSER 1966). All in all the transformation of the natural landscape to a cultural one with all of its implications can be considered the main cause for dıminution of these species, which are vegetarian or partly so (Sus scrofa) and which are more or less confined to the woods. It cannot be decided, whether this recent sıze reduction is a result of mutations or — until now — only expression of a modification caused by the bad conditions of life tor these species. However, it is obvious that a phylogenetic tendency of dıminution like that supposed by VON LEHMANN (1960) ın the case of roe deer after considering a far longer period cannot be the explanation for the present results, as the described size reduction ıs a modern event. Hare and fox are not affected in this way. These species require quite different living conditions. Originally the hare is a mammal of the steppe. Thus, the development of the cultural landscape with fields and pastures instead of woodland improved its ecological basıs, although some other living conditions were worsened. Therefore ıt ıs understandable that one cannot observe a clear trend of diminution. But one should also bear in mind, for instance, that the recent population from Schleswig-Holstein is not a truly autochthonous one: In the beginning of this century hares from many areas, especially from southeast Europe, were released here (MoHR 1931). The fox ıs a carnıvore and for this reason ıs only indirectly dependent on vegetation or landscape like the other anımals. Moreover it has a good ecological adaptabilıry. Acknowledgements I am much indebted to Mr. CHR. FREITAG for checking the English manuscript. The studies were supported by the Deutsche Forschungsgemeinschaft. References BoESsSNECK, J. (1956): Zur Größe des mitteleuropäischen Rehes (Capreolus capreolus L.) ın alluvial- vorgeschichtlicher und früher historischer Zeit. Z. Säugetierkunde 21, 121-131. GEHL, O. (1976): Die steinzeitliche Siedlung Stinthorst bei Waren/Müritz ım Spiegel des Säugetier- fundgutes. Bodendenkmalpflege in Mecklenburg, Jahrbuch 1975, 39-53. HEINRICH, D. (1985): Scharstorf, eine slawische Burg in Ostholstein. Haustierhaltung und Jagd. Offa-Bücher 59. Neumünster: Wachholtz. Horrmann, E. (1981): Schleswig und Lübeck im 12. und 13. Jahrhundert. Beitr. Schleswiger Stadtgeschichte 26, 26-38. HÜBNER, K.-D.; SauR, R.; REICHSTEIN, H. (1988): Die Säugetierknochen der neolithischen Seeufer- siedlung Hüde I am Dümmer, Landkreis Diepholz, Niedersachsen. Göttinger Schr. z. Vor- u. Frühgeschichte 23, 35-142. Size differences of game populations from Neolithic to modern times 63 Hüster, H. (1990): Untersuchungen an Skelettresten von Rindern, Schafen, Ziegen und Schweinen aus dem mittelalterlichen Schleswig (Ausgrabung Schild 1971-1975). Ausgrabungen in Schleswig. Berichte und Studien 8. Neumünster: Wachholtz. JoHansson, F. (1979): Die Knochenfunde von Säugetieren und Vögeln von Bistoft LA 11. In: Socio- ekonomiska strukturer ı tidigt neolitikum och deras förutsättningar. Studier över Bistoft LA 11 - ett boplatsfynd frän Schleswig-Holstein. Hrsg. von L. Jomansson. 98-111. Göteborg. Kreın, D. R.; STRANDGAARD, H. (1972): Factors affecting growth and body size of roe deer. J. Wildl. Manage. 36, 64-79. KrarTochviL, Z. (1988): Körpergröße des Rehwildes (Capreolus capreolus L.) ın der Gegenwart und Vergangenheit (tschechisch mit deutscher Zusammenfassung). Folia Venatoria 18, 115-133. LEHMANN, E. von (1960): Das Problem der Größenabnahme (Deminuations-Tendenz) beim Reh. Z. Jagdwiss. 6, 41-51. Mon, E. (1931): Die Säugetiere Schleswig-Holsteins. Altona. Nosıs, G. (1971): Die Tierreste der prähistorischen Siedlung Süssau Kr. Oldenburg (Schleswig- Holstein). Schr. Naturw. Ver. Schlesw.-Holst. 41, 89-99. — (1975): Zur Fauna des ellerbekzeitlichen Wohnplatzes Rosenhof in Ostholstein I. Schr. Naturw. Ver. Schlesw.-Holst. 45, 5-30. Paur, A. (1980): Untersuchungen an Tierknochen aus dem mnittelalterlichen Lübeck (Grabung Königstraße 59-63). Lübecker Schr. z. Archäologie u. Kulturgeschichte 2, 5-104. REICHSTEIN, H. (1974): Ergebnisse und Probleme von Untersuchungen an Wildtieren aus Haithabu (Ausgrabung 1963-1964). Berichte über die Ausgrabungen in Haithabu 7, 103-144. Neumünster: Wachholtz. — (1990): Die wildlebenden Säugetiere von Haithabu (Ausgrabungen 1966-1969 und 1979-1980). Berichte über die Ausgrabungen in Haithabu 29, Neumünster: Wachholtz (in press). REICHSTEIN, H.; TAEGE, K.-C.; Vocer, H.-P. (1980): Untersuchungen an Tierknochen von der frühslawischen Wehranlage Bischofswarder am Großen Plöner See (Schleswig-Holstein). In: Bosau. Untersuchungen einer Siedlungskammer in Ostholstein 4. Hrsg. von H. Hınz. Offa- Bücher 42, 9-75. Neumünster: Wachholtz. REQUATE, H. (1956): Die Jagdtiere in den Nahrungsresten einiger frühgeschichtlicher Siedlungen in Schleswig-Holstein. Schr. Naturw. Ver. Schlesw.-Holst. 28, 21-41. Rönrs, M. (1959): Neue Ergebnisse und Probleme der Allometrieforschung. Z. wiss. Zool. 162, 1-95. SÄGESSER, H. (1966): Über den Einfluß des Standortes auf das Gewicht des Rehwildes (Capreolus c. capreolus [Linne 1758]). Z. Jagdwiss. 12, 54-62. VoceL, V. (1983): Archäologische Stadtkernforschung in Schleswig 1969-1982. Ausgrabungen in Schleswig. Berichte und Studien 1, 9-54. Neumünster: Wachholtz. WALCHER, H. F. (1978): Die Tierknochenfunde aus den Burgen auf dem Weinberg in Hitzacker/Elbe und in Dannenberg (Mittelalter). II. Die Wiederkäuer. Diss. Univ. München. Author’s address: Dr. Dirk Heinrich, Institut für Haustierkunde, Universität Kiel, Olshausen- str. 40, W-2300 Kiel, FRG BEKANNTMACHUNG Einladung Die 65. Hauptversammlung der Deutschen Gesellschaft für Säugetierkunde e. V. findet von Sonntag, den 22. September, bis Donnerstag, den 26. September 1991, in Hamburg im Zoologischen Institut und Zoologischen Museum statt. Vorläufiges Programm Sonntag, 22. September: Anreise 17.00 Uhr: Vorstandssitzung 19.00 Uhr: Zwangloser Begrüßungsabend im Hotel Baseler Hof in der Esplanade Montag, 23. September: 9.00 Uhr: Grußworte und Eröffnung der Tagung durch den 1. Vorsitzenden im Hörsaal des Zoologi- schen Instituts und Zoologischen Museums 9.30 Uhr: Hauptvorträge und Vorträge zum Themen- schwerpunkt „Systematik und Stammesge- schichte der Säuger“ 14.30 Uhr: Vorträge 17.00 Uhr: Mitgliederversammlung Dienstag, 24. September: 9.00 Uhr: Hauptvortrag und Vorträge zum Themen- schwerpunkt „Anpassung an aquatische Lebens- räume“ 11.00 Uhr: Posterdemonstration 14.30 Uhr: Vorträge 16.30 Uhr: Posterdemonstration 20.00 Uhr: Filmabend Mittwoch, 25. September: 9.00 Uhr: Hauptvortrag und Vorträge zum Themen- schwerpunkt „Domestikation“ 14.00 Uhr: Vorträge 16.00 Uhr: Führung durch Hagenbecks Tierpark 19.00 Uhr: Geselliger Abend im Restaurant des Tierparks Donnerstag, 26. September: Hagenbeck Exkursion ın das östliche Schleswig-Holstein (Naturpark Lauenburger Seen) Alle Interessenten sind herzlich eingeladen. Neben den angekündigten Themenschwer- punkten werden auch dieses Mal andere Fachrichtungen mit Vorträgen und Posterdemon- strationen berücksichtigt. Das Programm mit der Vortragsabfolge wird allen Mitgliedern — auf Anfrage auch Nichtmitgliedern - rechtzeitig vor der Tagung zugesandt. Falls persön- liche Einladungen gewünscht werden, erbitten Sie diese vom 1. Vorsitzenden, Prot. Dr. E. KuULZER, Institut für Biologie III, Auf der Morgenstelle 28, W-7400 Tübingen. Bitte melden Sie Vorträge, die nicht länger als 15 min dauern sollen, sowie Posterde- monstrationen beim Geschäftsführer unserer Gesellschaft, Prof. Dr. U. SchmiDT, Zoolo- gisches Institut, Poppelsdorfer Schloß, W-5300 Bonn 1, Tel. (02 28) 73 54 68, möglichst frühzeitig, jedoch spätestens bis zum 30. Aprıl 1991 (Ausschlußfrist) an. Falls Sie Fragen zum Tagungsort und zur Organisation haben, wenden Sie sich bitte an Prof. Dr. H. SCHLIEMAnN, Zoologisches Institut und Zoologisches Museum, Martın- Luther-King-Platz 3, W-2000 Hamburg 13, Tel. (0 40) 41 23 39 17. Deutsche Gesellschaft für Säugetierkunde: Referate, Vorträge und Posterdemonstrationen der 64. Hauptversammlung 1990 in Hauptziel der Deutschen Gesellschaft für Säugetierkunde ist, auf ihren Jahrestagungen über Säugetiere arbeitende Wissenschaftler verschiedenster Fachrichtungen zusammenzuführen, den Gedanken- und Erfahrungsaustausch anzuregen, um so Erkenntnisse aus den einzelnen Forschungsgebieten zu integrieren. Die Kurzfassungen der Vorträge und Posterdemonstrationen von der 64. Haupt- versammlung, die in diesem Jahr in Osnabrück stattfand, liegen wiederum als Sonderheft der »Zeitschrift für Säugetierkunde« vor. Die Tagung war durch drei Themenschwerpunkte gekennzeichnet, die ökologische, ontogenetische und ernährungs- physiologische Fragestellungen behandeln. Zusätzlich wurde eine große Zahl freier Beiträge angeboten. Diese auch in der Vergangenheit geübte Art der Themen- darbietung soll die Breite der aktuellen Säugetierforschung widerspiegeln. Die säugetierbiologische Arbeit war in der Ethologie von Anfang an ein Forschungsschwer- punkt. Im Vordergrund stehen Projekte mit öko-ethologischer Fragestellung. Da sich dafür stenöke Säugetiere besonders eignen, werden bevorzugt silvicole und ripicole Arten in Gehegen und im Freiland beobachtet. Eine intensive Freilandarbeit, auch auf Exkursionen, förderte die Untersuchung von Kleinsäugetiergemeinschaften. Die Kurzfassungen der Vorträge und Posterdemonstrationen der Deutschen Gesellschaft für Säugetierkunde sind ab der 58. Hauptversammlung 1984 in Göttingen noch lieferbar. Zu beziehen durch jede Buchhandlung. % Deutsche Gesellschaft für Säugetierkunde, 64. Hauptversammlung in Osnabrück, 23. bis 27. September 1990. Kurzfassungen der Vorträge und Posterdemonstrationen. Heraus- egeben von Dr. Christel Schmidt, Bonn, und Prof. Dr. Rüdiger PAJL Schröpfer, Osnabrück. 1990. 55 Seiten. Kartoniert 24,- DM Verlag Paul Parey - Spitalerstraße 12 - 2000 Hamburg 1 P/REV Subscription information for USA, Canada and Mexiko obtainable from Paul Parey Scientific Publishers/IR Unltd., 35 West 38th Street, Suite 3 W, New York, NY 10018, USA. Erscheinungsweise und Bezugspreis 1991: 6 Hefte bilden einen Band. Jahresabonnement Inland: 338,— DM zuzüglich 13,80 DM Versandkosten, Gesamtpreis 351,80 DM einschließlich 7% Mehrwert- steuer. Jahresabonnement Ausland: 338,- DM zuzüglich 18,-— DM Versandkosten. Das Abonne- ment wird zum Jahresanfang berechnet und zur Zahlung fällig. Es verlängert sich stillschweigend, wenn nicht spätestens am 15. November eine Abbestellung im Verlag vorliegt. Die Zeitschrift kann bei jeder Buchhandlung oder bei der Verlagsbuchhandlung Paul Parey, Spitalerstraße 12, W-2000 Hamburg 1, Bundesrepublik Deutschland, bestellt werden. Die Mitglieder der „Deut- schen Gesellschaft für Säugetierkunde“ erhalten die Zeitschrift unberechnet im Rahmen des Mitgliedsbeitrages. Z. Säugetierkunde 56 (1991) 1, 1-64 Mit einer Beilage des Verlages Paul Parey Edgar Schallenberger Charakterisierung von Sekretionsrhythmen der Gonadotropine und Ovarsteroide während des Brunstzyklus, der Gravidität und post partum beim Rind (Advances in Veterinary Medicine - Fortschritte der Veterinärmedizin, No. 40) Von Dr. Dr. E. Schallenberger, München. 1990. 117 Seiten mit 60 Abbildungen und 21 Tabellen. 25x 17 cm. Kartoniert DM 68,— ISBN 3-489-51316-9 Ein aktuelles Arbeitsgebiet von Biotechnik und Medizin ist die willkürliche Steuerung und Manipulation von Fortpflanzungsfunktionen. Obwohl bei den landwirtschaftlichen Nutztieren häufig und oft durchaus erfolgreich Fortpflanzungssteuerung durchgeführt wird, sind die endokrinen Abläufe, in die eingegriffen wird, teilweise erst in Ansätzen bekannt. Deshalb wer- den in dieser Arbeit einige Zusammenhänge von Hormonsekretionsrhyth- men beim gesunden adulten Rind während Brunstzyklus, Gravidität und Post-Partum-Phase aufgezeigt, um einen Einblick in wesentliche Grund- prinzipien hierarchisch gegliederter neuroendokriner Regelsysteme zu geben. Viele der exemplarisch beim Rind aufgezeigten Zusammenhänge können zur vergleichenden Interpretation von Befunden bei anderen Spe- zies herangezogen werden. Die Daten sind für Veterinärmediziner, Tierpro- duzenten, Biologen und Endokrinologen gleichermaßen von Interesse. Aus dem Inhalt: Allgemeiner Überblick über die Steuerung der Reproduktion beim weiblichen Rind — Das Hypothalamus-Hypophysensystem — Endokrine Signalübertragung — Entwicklung von Versuchstechniken und Meßverfahren zur Erfassung diskonti- nuierlicher Hormonsekretion — Brunst- und Zyklusfeststellungen — Probenent- nahmen und -konservierung — Darstellung diskontinuierlicher Hormonsekretion — Radioimmunologische Hormonbestimmung — Eigene Ergebnisse — Sekretions- muster von Gonadotropinen und Ovarsteroiden während des physiologischen Brunstzyklus — Messungen der Gonadotropine in der V. jugularis bei 5 Minuten Probenintervall — Vergleichende Messungen in der V. cava caudalis und der V. jugularis während der Lutealphase - Vergleichende Messungen in der V. cava caudalis und der V. jugularis während der Vorbrunst- und Brunstphase — Hormon- sekretionsmuster während der Gravidität — Diskussion der eigenen Befunde — Hypophysen- und Gelbkörperfunktion p. p. — Regelkreis wichtiger Fortpflan- zungshormone -— Allgemeine Schlußfolgerung bezüglich der Relevanz der unter- suchten Hormone in Blutproben — Summary. PAUL Berlin und Hamburg | P/REV /ol. 56 (2), 65-128, April 1991 ISSN 0044-3468 C21274F ZEITSCHRIFT FÜR SAUGETIERKUNDE INTERNATIONAL JOURNAL OF MAMMALIAN BIOLOGY Organ der Deutschen Gesellschaft für Säugetierkunde Bruorton, M. R.; Perrin, M. R.: Comparative gut morphometrics of Vervet (Cercopithecus aethiops) and Samango _ (C. mitis erythrarchus) monkeys. — Vergleichend-morphometrische Untersuchungen am Verdauungskanal von Grünaffen (Cercopithecus aethiops) und Diademaffen (C. mitis) 65 Poglayen-Neuwall, I.; Shively, J. N.: Testicular cycles of the Ringtail, Bassariscus astutus (Carnivora: Procyonidae). I Testicularzyklen des Katzenfretts, Bassariscus astutus (Carnivora: Procyonidae) 72 lade, J. van der: Sexual bimodality in some recent pig populations and application of the findings to the study of _fossils. - Geschlechtsbimodalität bei einigen rezenten Schweinepopulationen und Anwendung der Befunde für das Studium an Fossilien 81 Ingold, P.; Marbacher, H.: Dominance relationships and competition for resources among chamois Rupicapra u pra rupicapra in female social groups. — Dominanzbeziehungen und Konkurrenz um Ressourcen in Gruppen von weiblichen Gemsen (Rupicapra rupicapra rupicapra) 88 Jacobsen, N. H. G.; Newbery, R. E.; de Wet, M. J.; Viljoen, P. C.; Pietersen, E.: A contribution of the ecology of the Steppe pangolin Manis temminckii in the Transvaal. — Ein Beitrag zur Ökologie des Steppenschuppentieres _ Manis temminckii in Transvaal 94 \eal, B. R.: Seasonal changes in reproduction and diet of the Bushveld gerbil, Tatera leucogaster (Muridae: _Rodentia), in Zimbabwe. — Saisonale Schwankungen in der Fortpflanzung und in der Nahrungszusammenset- "zung beim Bushveld Gerbil Tatera leucogaster (Muridae: Rodentia) in Zimbabwe 101 Jomparatore, Viviana M.; Maceira, N. O.; Busch, Cristina: Habitat relations in Cienomys talarum (Caviomorpha, "Octodontidae) in a natural grassland. — Habitat-Beziehungen von Ctenomys talarum (Caviomorpha, Octodonti- ‚dae) auf natürlichem Grasland 112 Nissenschaftliche Kurzmitteilungen "alomares, F.: Use of an active badger sett by Egyptian mongooses, Herpestes ichneumon, in Southwest Spain. — _ Benutzung eines bewohnten Dachsbaus durch Ichneumone, Herpestes ichneumon, in Südwest-Spanien 119 arrette, Cyrille: Fully formed first lower premolars in a White-tailed deer (Odocoileus virginianus Zimmermann, - 1780). - Vollständig ausgebildete erste untere Prämolaren bei einem Weißwedelhirsch (Odocoileus virginianus Zimmermann, 1780) I pinosa, Maria Beatriz; Vitullo, A. D.; Merani, Maria S.: Chromosomes of the Argentine Andean mouse, Akodon andinus (Cricetidae: Sigmodontinae). - Chromosorgen. der argentinischen Andenmaus Akodon andinus (Criceti- dae: Sigmodontinae) HSONIa 124 kanntmachung SM Se 126 Ichbesprechungen 127 JUN 2 7 1991 | LIBRARIES Verlag Paul Parey Hamburg und Berlin HERAUSGEBERJEDTIEORS P. J. H. van BrEE, Amsterdam - W. FIEDLER, Wien — H. Frıck, München - W. HERRE, Kiel - H.-G. Krös, Berlin - H.-J. Kunn, Göttingen - E. Kurzer, Tübingen - B. Lanza, Florenz - W. MATER, Tübingen — J. NIETHAMMER, Bonn — Anne E. Rasa, Pretoria — H. REICHSTEIN, Kiel - M. Röhrs, Hannover - H. SCHLIEMAnN, Hamburg — D. STARcK, Frankfurt a. M. - E. THENnIUs, Wien - P. VoGeEL, Lausanne SCHRIEITEITUNGZEDTLORIEIZOFEINER D. Kruska, Kiel - P. LANGER, Gießen This journal is covered by Biosciences Information Service of Biological Abstracts, and by Current Con- tents (Series Agriculture, Biology, and Environmental Sciences) of Institute for Scientific Information Die Zeitschrift für Säugetierkunde veröffentlicht Originalarbeiten und wissenschaftliche Kurzmittei- lungen aus dem Gesamtgebiet der Säugetierkunde, Besprechungen der wichtigsten internationalen Literatur sowie die Bekanntmachungen der Deutschen Gesellschaft für Säugetierkunde. Verantwort- licher Schriftleiter im Sinne des Hamburgischen Pressegesetzes ist Prof. Dr. Dieter Kruska. Zusätzlich erscheint einmal ım Jahr ein Heft mit den Abstracts der Vorträge, die auf der jeweiligen Hauptversammlung der Deutschen Gesellschaft für Säugetierkunde gehalten werden. Sie werden als Supplement dem betreffenden Jahrgang der Zeitschrift zugeordnet. Verantwortlich für ihren Inhalt sind ausschließlich die Autoren der Abstracts. 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For copying from back volumes of this journal see “Permissions to Photo-Copy: Publisher’s Fee List” of the Fortsetzung 3. Umschlagseite © 1991 Paul Parey. Verlag: Paul Parey, Hamburg und Berlin. Anschriften: Spitalerstr. 12, W-2000 Hamburg 1: Lindenstr. 44-47, W-1000 Berlin 61, Bundesrepublik Deutschland. — Printed in Germany by Westholsteinische Verlagsdruckerei Boyens & Co., Heide/Holst. Z. Säugetierkunde 56 (1991) 65-71 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Comparative gut morphometrics of Vervet (Cercopithecus aethiops) and Samango (C. mitis erythrarchus) monkeys By M.R. Bruorton and M.R. PERRIN Department of Zoology and Entomology, University of Natal, Pietermaritzburg, South Africa Receipt of Ms. 2. 5. 1990 Acceptance of Ms. 18. 9. 1990 Abstract A comparative morphometric examination of the gastrointestinal tracts of vervet (Cercopithecus aethiops and samango C. mitis erythrarchus monkeys revealed that samangos possess significantly larger volumes in the hindgut (caecum and colon). The surface area of the main absorptive region of the gastrointestinal tract (the small intestine) was also larger in samangos than in vervets. Samangos include larger amounts of fiıbrous leaf material in their diets than vervets, and the importance of the larger volumes are discussed with reference to the fermentative process required to digest and utilise this foliar component. Introduction The structure of the gastrointestinal tract (GIT) ıs faırly homogeneous among different orders of mammals (CHiveErs and Hrapık 1980), and development of different parts of the GIT generally retlect adaptations to different foods. Plant food with a high content of structural carbohydrates can be digested with the help of microbes in either the stomach or the hindgut, where the enzymes produced by the microbes degrade the food and render its chemical constitutents absorbable (LANGER 1988). Among the African primates are examples of both foregut (stomach) and hindgut (caecum and colon) fermenters. The folivorous colobines possess plurilocular, haustrated stomachs, while most of the frugivor- ous/omnivorous cercopithecines possess simple unilocular, glandular stomachs (LANGER 1988), a well-developed caecum and a large and haustrated colon. The gross morphology and ultrastructure of the GIT of vervet monkeys (Cercopithecus aethiops) and samango monkeys (C. mitis) ıs similar (BRUORTON 1989). Samangos are frugivorous, but include a relatively high proportion of leaves in their diets (Rupran 1978; Corps 1987; Lawes 1990), while vervets are generally regarded as frugivorous/omnivor- ous and consume roughly equal proportions of fruit and anımal (mostly insect) material (Dunsar and Dunsar 1974; KavanacH 1978; Watson 1985). GIT morphology may show considerable varıation even between frugivorous primate species, depending on the amount of insects or leaves included in the primarily fruit diet (CHIvErs and HLanık 1980), and this may include expansion within the tract. This is usually associated with the fermentation process, since the larger the volume, the more fermentation can occur (CHIVERS and Hrapiık 1980). A comparative examination was initiated to determine whether the volumes of varıous regions of the gastrointestinal tract (and surface area of small intestine) differ significantly between the two primate species. Volumes are indicators of physiological processes, such as microbial digestion, that are important in herbivore nutrition. This comparative examination might therefore assist in providing an explanation of the digestive strategies of the two species, and in determining how samangos digest the foliar components of their diet. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0065 $ 02.50/0 66 M. R. Bruorton and M. R. Perrin Materials and methods Nine vervet monkeys (4 adult males, 5 adult females) and eight samangos (4 adult males, 4 adult females) were shot on the farm Braco in the Karkloof area of Natal and used in this examination. Animals were weighed in the field and body lengths were measured from tip of nose to tip of tail. Gastrointestinal tracts were removed immediately. These were examined in the laboratory 2-3 hours later, which allowed for complete relaxation of the musculature in the GIT wall. No specimens were fixed prior to measuring, as this can cause distortion of the tract wall. Measurements of the stomach, small intestine, caecum and colon, completely cleared of all mesenteric tissue, were made in a water-filled 2 mx0.5 mx0.1 m basin. Measuring the length of the gut under water minimises stretching, and is especially important when measuring small intestinal length. Small and large intestines were treated as cylinders and volumes were calculated from mean measurements of length and circumference. Width varıes along the length of intestine, and at least 5 measurements of width (of intact tissue) were taken at regular points along the length. At a calculated mean width the tissue was opened and the circumference was measured. The surface area of the small intestine was also calculated from length and circumference measurements. To calculate the volume of the caecum, an incision was made dorsally from the ıleocaecal junction to the apex of the caecum. The contents were then flushed out and the lateral walls of the caecum were flattened into an approximate cylindrical shape. The length measurement and average circumference could then be taken and the volume calculated as for the colume of a cylinder. Accurate measurements of stomach volume were difficult to obtain, largely as a result of its irregular shape and size in different anımals. In most anımals the length of the intact stomach was measured. It was then opened up along the lesser curvature from the gastro-oesophageal opening to the pyloric sphincter, and the incision was extended dorsocranially along the greater curvature for approximately 3 cm (Fig.). The stomach could then be opened out (flattened), the circumference measured and the volume calculated as for the volume of a cylınder. Two other methods used, a. estimation of stomach volume by filling it with water and b. calculation of stomach volume from the greater curvature measurement (CHIVERS and HrLADIK 1980), were found to be unacceptable and were discontinued. Previous comparative work on primate GIT morphometrics involved use of length and width measurements of intact organs (Jones 1970; MILTON 1981). Preliminary work in this study included this method, but these measurements were found to be unacceptable. Firstly, the varyıng thickness of gut musculature means that simple width measure- ments are mostly inaccurate (often by more than 1 cm). Secondly, the degree of stomach (or intestinal) fill, or the time elapsed since feeding, mean that stomachs of essentially similar size can provide very different measurements depending on when (time of day) they were collected. It is essential that measurements be taken only of dissected sections of the GIT with the contents removed. This negates the effects of varying muscle thickness (because measurements are taken on the mucosal surface of the gut), and is likely to provide more consistent results independent of whether the animal has recently fed or not. Diagrammatic representation of a stomach, showing the incision made before stomach is opened out (flattened) and measured. P = pyloric sphincter, L = lesser curvature (gastro-oesophageal opening to pyloric sphincter), G = gastro-oesophageal opening, GC = greater curvature (gastro-oesophageal opening to pyloric sphincter) Comparative gut morphometrics of Vervet and Samango monkeys 67 To compensate for the effect of the larger body size of samangos, volumes of regions of the GIT in each animal were divided by the metabolic live mass (mass°°”) of that animal. This provides a ratio that enables direct statistical comparison between species of the volumes of specific regions of the gut. Data were analysed using the Mann-Whitney two sample rank testing procedure (ZAr 1974). Results Table 1 compares the measurements of volumes of stomach, small intestine, caecum and colon of adult (male and female) samangos and vervets. The mean body mass of samangos significantly (P<0.05; Mann-Whitney U) exceeds that of vervets (males 8.33 kg : 5.58 kg; females 4.93 kg : 3.82 kg). Table 2 presents gut volumes corrected for differences in body mass of the two species, and includes significant differences in GIT volumes. Table 3 shows small intestinal surface areas of males and females ın each of the two species, and includes significant differences in the surface areas, corrected for different body mass, between the two species. Table 1. Measurements of body length and mass, and of volumes of stomach, small intestine, caecum and colon in samango and vervet monkeys (Mean + standard deviation) Length Mass Volume (cm’) (cm) (kg) Stomach SI Caecum Colon Samango Adult males [432EE215788: 4 265-9165 366== 45, 2106==715557285=3762 (m = 4) Adult females PIE: f 110919207375 22273 199,4 8 0425=12,99 (n=4) Vervet Adult males 109 #& 1.4 i ; 194 #44 179 #54 2 8), Sl (n= 4) Adult females ONE ROSE EAU 168 +24 148 +41 SE I) ze 1l3) (n = 5) SI = Small intestine. Table 2. GIT volumes corrected for differences in body mass between samangos and vervets Volume of section of tract/metabolic mass of the anımal. (Mean + standard deviation) Volume (cm?) Stomach SI Caecum Adult males Samango (n=4) 64.3 + 3.3 88.5 + 9.9 ee er Vervet (n=4) 61.0 #10 56.0 + 13.5° 08205. .10052 13.9.7 Adult females Samango (n=4) 58.3 + 7.9 (sea ED SE 29 Vervet (ns=5) 68.4 # 8.0 9022 == 1. GO 3Rl. VE Values with common superscripts in columns differ significantly using Mann-Whitney U (P <.0.05); SI = Small intestine. 68 M. R. Bruorton and M. R. Perrin Table 3. The mean surface area of the small intestine in samango and vervet monkeys, including correction for the different body mass of the species (Mean + standard deviation) Species Surface area (cm?) actual corrected Samango male (n=4) 11163 =e 12% al 28277 female (n=4) 1111 #48 DEE 2ER Vervet male (n=4) 692 # 87 DIESEN, female (n=5) 595 + 92 DADEBS Values with common superscripts differ significantly using Mann-Whitney U (P < 0.05). Discussion Vervets and samangos occur syntopically in the Karkloof, and the same food resources were therefore available to both species. This was considered important because of the propensity of the gastrointestinal tract to undergo adaptive changes ın relation to forage quality in different areas. These adaptive changes related to dietary varıation are well documented (PoksEy and SCHNEEMAN 1983; Gross et al. 1985; PERRIN 1987). Similarly, in this study ıt was noted that culled vervets from marginal habitats had stomachs which were sıgnificantly smaller than those from the Karkloot area. It therefore seems important that comparative GIT measurements be taken on species occurring in sympatry, where the etfects of food quality in different habitats would not affect gut morphology or capacity, and where different species are able to specialise on their preferred diets. Only adult anımals were used to obtain the comparative morphometric data. (There were large variations in intraspecific measurements of juvenile and subadult age-classes.) The importance of different organs might vary according to anımal age and organ function (Jornson and McBee 1970; BRUORTON and PERRIN 1988), and ıt was concluded that only GIT measurements from adult anımals would provide consistent and valıd results. There were no significant differences in stomach volumes between the two species. Both have a simple unilocular glandular stomach (BRUORTON and PERRIN 1988), which is characteristic of Old World cercopithecines (Hırr 1958, 1966). On the basıs of their similar morphology and function, it ıs perhaps to be expected that minimal size varıations should occur between the two. MıLTon (1981) showed that stomach size of two sympatric primates in Panama, one highly folivorous (Alounatta palliata) and one highly frugivorous (Ateles geoffroyi), were approximately equivalent. This is supported (for Alonatta and Ateles) by Hıapık (1967) in his work on the relative surface area of sections of the digestive tracts of 24 primate species. SCHIECK and MILLAR (1985) showed that it is the distal parts of the digestive tract (caecum and colon) that correctly indicate diet type within rodent families. It appears, therefore, that stomach volumes remain fairly constant relative to body weight within primates possessing simple stomachs, and that differences in GIT volumes related to dietary adaptations would be found in other sections of the digestive tract. Significant differences in small intestinal surface area existed between both males and females of the two species. This finding is not entirely consistent with previous work on other species. SCHIECK and MırrAr (1985) stated that small intestine lengths did not accurately reflect the amount of fibre in the diet of the 35 small mammal species they examined. Similarly, Mırron (1981) and Hıavık (1967) showed that small intestinal surface areas of the sympatric primates Alouatta palliata and Ateles geoffroyi (even with their extremely different diets) were approximately equivalent. Jones (1970) also found that relative lengths of the small intestine were fairly constant among the eight African Comparative gut morphometrics of Vervet and Samango monkeys 69 cercopithecids he studied. However, CHIVERS and HraDik (1980) showed, by regressing surface area of small intestine against body size, that absorptive surface area increases proportionately with metabolic body mass. The generally larger areas of small intestine ın samangos might therefore simply be a factor of their larger sıze. Also, the consistently larger (relative) surface area ın adult females (than males) might be related to the greater energy demands of the female during pregnancy and lactation, which are known to cause an increase in both small intestinal length and water content (Crıpps and WILLıaMms 1975; Gross et al. 1985). The caecum and colon of samangos were highly significantly more voluminous than those of vervets. This finding of greater caecal and colonic lengths and volumes ın the more herbivorous species is well supported by previous studies. SCHIECK and MILLAR (1985) reported that colon lengths and weights represented diet types well, with herbivores having relatively larger large intestines than granivores and omnivores. JOnEs (1970) found that in eight cercopithecids the more folivorous species possessed relatively greater colon lengths, and in some cases greater caecum length. Sımilarly, MırrTon (1981) showed that the folivorous Alonatta palliata possessed a longer colon (double the surface area) than the frugivorous Ateles geoffroyi. The caecum of herbivores ıs also generally larger than that of granivores or omnivores (SCHIECK and MıLLAr 1985), and functions in the microbial conversion of fibrous material into compounds that can be absorbed by the colon (SHARKEY 1971). Evidence for the importance of the caecum of the samango in fermentation has already been presented (BRUORTON and PERRIN 1988). A further study has also shown high concentrations of organic acıds, as well as numerous bacteria, within the caecum and colon of both samangos and vervets. It might be expected that the larger volumes of caecum and colon in the more folivorous samangos confers some selective advantage over vervets with respect to microbial digestion of leaves and other fibrous material. This would occur ıf the greater volumes were found to be important ın food retention (or decelerating the passage rate of fibrous material), or if different species of fibre-digesting bacteria were present in one of the species. However, CLEMENS and Maroıy (1981) and CLEMENS and PHırLıps (1980) have shown that passage rates of food in vervets and in Sykes monkey C. mitis kolbi are extremely similar, with vervets retainıng some particulate markers for longer time periods than did Sykes. They also suggest that food retention ıs not necessarıly an important factor for the accumulation of organic acıds, as high concentrations were observed in the colon of the bushbaby Galago crassicandatus (CLEMENS and Marory 1981), which ıs smooth and non-complex and permits rapıd passage of digesta. It ıs also suggested that herbivorous primates do not necessarıly hold a fermentative advantage over omnivorous primates (CLEMENS and PhiırLıps 1980), and that GIT structure does not appear to influence the concentrations of organıc acıds found in the mammalian hindgut. Any advantages conterred by larger hindgut volumes are probably related to fermenta- tion capacity ın the two species. The larger the volume, the greater the amount of organic acıds which may be produced by the microbial populations. Thus, even though the concentrations of organic acıds ın the hindgut of the two species are sımilar, the larger caecum and colon of the samango allows for greater production of these important microbial degradation compounds. Volatile fatty acıds are rapıdly absorbed by the large- intestinal mucosa of a wide range of mammalıan species, usually at a rate similar to that determined for rumen epithelium (RERAT 1978; STEvENS et al. 1980). However, the quantity of organıc acıds absorbed by the large intestine of a given species depends primarily upon the surface area available (STEvEnSs et al. 1980). With the high component of leat material in their diets, the extensive hindgut regions of samangos are, therefore, not only important in allowing a greater production of organic acıds, but also for the greater absorption and assımilation of these important fermentation end-products. 70 M. R. Bruorton and M. R. Perrin Acknowledgements We would like to thank the Natal Parks Board for issuing us with permits to collect 10 samangos. MRB thanks the Foundation for Research Development (C.S.I.R.) of South Africa for a bursary during the course of his study, and both authors thank the University Research Committee of the University of Natal for funds granted during the course of the study. Zusammenfassung Vergleichend-morphometrische Untersuchungen am Verdauungskanal von Grünaffen (Cercopithecus aethiops) und Diademaffen (C. mitıs) Vergleichend-morphometrische Untersuchungen am Verdauungskanal von Grünaffen und Diademaf- fen zeigten, daß Diademaffen wesentlich größere Raumanteile im hinteren Darm (Caecum und Colon) besitzen. Die Hauptabsorptionsoberflächen des Verdauungskanals (das Ileum) zeigte bei Diademaften ebenfalls größere Ausdehnung als bei Grünaffen. Diademaffen fressen grundsätzlich mehr faseriges Laubmaterial als Grünaffen. Die Bedeutung der größeren Darmräume wird ım Hinblick auf den Gärungsprozeß, der zu Aufbereitung und Verdauung dieses Laubbestandteils nötig ist, diskutiert. References BRrUORTON, M. R. (1989): A comparative morphological and physiological examination of the gastrointestinal tracts of vervet and samango monkeys. M. Sc. Diss., Univ. Natal, Pietermaritz- burg, South Africa. BRUORTON, M.R.; PERRIN, M. R. (1988): The anatomy of the stomach and caecum of the Samango monkey, Cercopithecus mitis erythrarchus Peters, 1852. Z. Säugetierkunde 53, 210-224. CHivErs, D. M.; Hrapik, C. M. (1980): Morphology of the gastrointestinal tract in primates: comparisons with other mammals in relation to diet. J. Morphol. 116, 337-386. CLEMENS, E. T.; PHırLıps, B. (1980): Organic acıd production and digesta movement in the gastrointestinal tract of the baboon and sykes monkey. Comp. Biochem. Physiol. 66, 529-532. CLEMENS, E. T.; Marory, G. M. ©. (1981): Organic acıd concentrations and digesta movement in the gastrointestinal tract of the bushbaby (Galago crassicandatus) and vervet monkey (Cercopithecus pygerythrus). J. Zool., Lond. 193, 487-497. Corps, M. (1987): Mixed-species associations of Cercopithecus monkeys in the Kakamega forest, Kenya. Berkeley: University of California Press. Crıpps, A. W.; Wırrıams, V. J. (1975): The effect of pregnancy and lactation on food intake, gastrointestinal anatomy and the absorptive capacıty of the small intestine in the albino rat. Br. ]. Nutr. 33, 17-32. Dunsar, R. I. M.; Dunsar, E. P. (1974): Ecological relations and niche separation between sympatric terrestrial primates in Ethiopia. Folia. primatol. 21, 36-61. Gross, ]J. E.; Wang, Z.; WUNDER, B. A. (1985): Effects of food quality and energy needs: changes in gut morphology and capacity of Microtus ochrogaster. J. Mammalogy 66, 661-667. Hırr, W.C. ©. (1958): Pharynx, oesophagus, stomach, small intestine and large intestine. Form and position. Primatol. 3, 139-207. — (1966): Primates. Comparative anatomy and taxonomy. VI. Catarrhini, Cercopithecoidea, Cer- copithecinae. New York: Interscience Publ. Hıapık, C. M. (1967): Surface relative du tractus digestif de quelques primates, morphologie des villosites intestinales et correlations avec le regime alımentaire. Mammalıa 31, 120-147. Jo#nson, J. L.; McBer, R. H. (1967): The porcupine caecal fermentation. J. Nutrition 91, 540-546. Jonss, C. (1970): Stomach contents and gastrointestinal relationships of monkeys collected at Rio Muni, West Africa. Mammalia 34, 107-117. KavanacH, M. (1978): The diet and feeding behaviour of Cercopithecus aethiops tantalus. Folia primatol. 30, 30-63. LANGER, P. (1988): The mammalian herbivore stomach. Comparative anatomy, function and evolu- tion. Stuttgart, New York: Gustav Fischer Verlag. Lawes, M. J. (1990): Socio-ecology of the samango monkey and conservation in Natal. Ph. D. Diss., Univ. Natal, Pietermaritzburg, South Africa. Mırton, K. (1981): Food choice and digestive strategies of two sympatric primate species. Amer. Nat. 117, 496-505. PErRRINn, M. R. (1987): Effect of diet on the gastric papillae and microflora of the rodents Mystromys albicandatus and Cricetomys gambianus. S. Afr. J. Zool. 22, 67-76. Poxsey, K. $.; SCHNEEMAN, B. ©. (1983): Pancreatic and intestinal response to dietary guar gum in rats. J. Nutr. 113, 1544-1549. Comparative gut morphometrics of Vervet and Samango monkeys 71 RErAT, A. (1978): Digestion and absorption of carbohydrates and nitrogenous matters in the hindgut of the omnivorous nonruminant anımal. J. Anım. Scı. 46, 1808-1837. Rupran, R. (1978): Intergroup dietary differences and folıvorous tendencies of two groups of blue monkeys, Cercopithecus mitis. In: The ecology of arboreal folivores. Ed. by G. G. MonTGoMm- ERY. Washington D. C.: Smithsonıan Inst. Press. SCHIECK, J. ©.; MILLAR, J. S. (1985): Alımentary tract measurements as indicators of diets of small mammals. Mammalıa 49, 93-104. SHARKEY, M. J. (1971): Some aspects of coprophagy in rabbits and guinea pigs fed fresh lucerne. Mammalıa 35, 162-168. STEVENS, C. E.; ARGENZIO, R. A.; CLEMENS, E. T. (1980): Microbial digestion: rumen versus large intestine. In: Digestive physiology and metabolism ın ruminants. Ed. by. Y. RuckeEguscH and P. THIvEnD. Westport, Connecticut: Avı Publishing Comp. Watson, L. H. (1985): The feeding ecology of chacma baboons and vervet monkeys in the Hluhluwe-Umfolozi Reserve. M. Sc. Diss., Univ. Natal, Pietermaritzburg, South Africa. ZAR, J. H. (1974): Biostatistical analysıs. London: Prentice-Hall International. Authors’ address: MICHAEL R. BRUORTON and Prof. Dr. MıcHAEL R. PERRIN (for correspondence), Department of Zoology and Entomology, University of Natal, P. ©. Box 375, Pietermaritzburg 3200, Natal, South Afrıca Z. Säugetierkunde 56 (1991) 72-80 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Testicular cycles of the Ringtail, Bassariscus astutus (Carnivora: Procyonidae) By I. PoGLAvYEN-NEUWALL and ]J. N. SHIVELY Department of Ecology and Evolutionary Biology and Department of Veterinary Science, University of Arızona, Tucson Receipt of Ms. 22. 5. 1990 Acceptance of Ms. 13. 7. 1990 Abstract Annual varıation of testes in Bassariscus astutus was studied by palpation and morphological technique. Seasonal changes of spermatogenesis and testicular weight/size were compared. It could thus be confirmed that in Arizona the mating season extends from late winter into spring and that testes become aspermic ın summer and reach their smallest size in autumn. Introduction The purpose of this study ıs to ascertain the annual cycle (seasonal development and regression) of the testes in Bassariscus astutus, and the determination of the male’s ability (or inability) to produce active spermatozoa throughout the year. Ringtails appear to be seasonally monestrous, with the mating season extending from about mid February to May as evidenced by the majority of litters occeurring during May and June (GrRINNELL et al. 1937; POGLAYEN-NEUWALL and POGLAYEN-NEUWALL 1980; TAyLor 1954; TOWEILL 1976). There exists, to date, no histological study supporting this assumption. Material and methods Subjects and context Thirteen wild-caught males were palpated at the time of capture and released while 23 captive males, ages ranging from 1 to 7 years, were palpated bi-weekly throughout the year to determine the size of the testes. In addition, 21 pairs of testes were taken from sacrificed anımals, trap casualties, and fresh road kills. These were acquired for each month of the year. Ages of the anımals, if not known, were estimated by tooth wear and by the morphology of the baculum (after WoonD 1952). Testes were excised, and after removal of the tunıca vaginalıs, weighed (including epididymis) to the nearest 0.01 g. The greatest length and width (exclusive epididymis) were recorded to the nearest 0.01 mm. Each testis was hemisected longitudinally. Of each pair one testis was fixed in 10% phosphate- buffered formalın of pH 7-0, for light microscopy. The other was fixed in a combination of 4% commercial formaldehyde and 1% glutaraldehyde in a buffer of 176 m/O/sm/liter (McDoweELL and Trump 1976) for electron and/or light microscopy. Specimens for electron microscopy were not available from the months of January, September und October. Histological methods For light microscopy longitudinal slices of the formalın-fixed testicle were cut, dehydrated and embedded in paraffın. Sections were cut a 5 um and stained with hematoxylin and eosın. For transmission electron microscopy blocks no larger than 1 um in any dımension were cut from the most superficial tissue, postfixed in 2% OsO, in phosphate buffer, pH 7.4 for 1 hour, dehydrated ın an ascending series of alcohol and propylene oxıde and embedded ın an epon-araldite mixture (MOLLENHAUER 1963). These sections were cut with glass knives, mounted on naked copper grids and stained with uranyl acetate and lead citrate. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0072 $ 02.50/0 Testicular cycles of the Ringtail, Bassariscus astutus 73 Seminiferous epithelial area (SEA) (BAsURTO-Kußpa et al. 1984) was estimated by photographing randomly selected seminiferous tubules at a magnification of x50. Photographic prints were made to give a final magnification of x425. Photographs were taken wıth a Nikon camera with Nikon attachment AMF on a Nikon labophot microscope. The basal membrane and the luminal margin were delineated to include all epithelial cells in 15 to 40 approximately round tubules in each animal. Coordinates of the seminiferous tubules were then digitized using a BQ CAM microcomputer system (R. and M. Biometrics, Inc., 5611 Ohio Ave., Nashville, TN 37209). The epithelial area of each tubule cross section was determined as the area inclosed by the basal membrane minus the area of the lumen. The SEA of each anımal was expressed in square mm as the mean of the tubules measured. Student’s t-test was used to test significance of differences between anımals. Five degrees of activity using a scale of O to 4+ were used to characterize sections of each testicle. For spermatogenesis in seminiferous tubules activity corresponded to the following: O0 = absence of sperm; I+ = at least 1 sperm/sperm head; 2+ = several sperm/sperm heads; 3+ = moderate number of sperm/sperm heads; 4+ = many sperm/sperm heads. A similar scale was used to evaluate the number of sperm in the epididymis. Results Summary of the morphologic evaluation, morphometry (SEA) and statistical sıgnificance testing ıs indicated ın Table 1. In late winter and early spring testes were most active. SEA was statistically low in December, January and February. A significant increase occurred in March, April and May and a marked decrease in June. From July to November testes were inactive. A rapıd increasee in SEA between December and January was apparent. The quantitiy of sperm in seminiferous tubules lagged 1 to 2 months behind changes in SEA (Table 1). Sımilarly changes in sperm concentration ın epididymis usually was recorded a month later than in seminiferous tubules. Table 1. Spermatogenetic stages Month Sperm in sem. tub. Sperm in epididymis jan: Beb: Mar. Apr. May June July Aug. Sept. Orc: Nov. Dec ESISISIOLSET Et t ! Calculated from the first day of each month. Histopathological characteristics of the testis and epididymis are related to the stage of activity which correlated with the time of year (Figs. 1-5) and resembled those of other species in similar state of activity, e.g. Norway rat, rabbit, European boar and domestic cat (Moens et al. 1975; MoRTOoN et al. 1986; BasurTo-Kußa et al. 1984; ELcock et al. 1984). Degenerative changes occurred in the seminiferous epithelium during the period of declining activity, although there were a few stem cells present. Weights of testes (Table 2) were lowest in September-October with steep increase toward March. Testes in March show maximum weight. From April to September the weight declines steadily. Similarly, linear measurements of testes show maximum length and width in March, and minimum length and width in September-October (Fig. 6). Fig. 2. Seminiferous tubules of inactive phase. Sertoli cells predominate (September). (X200) Y a x \ “_» vn eo 6%. a En ' Pr % gun wm a , eh 6, Pt ur Be NE Fig. 4. Epididymis from inactive phase (August). (X200) 76 1. Poglayen-Neuwall and J. N. Shively Fig. 5. Spermatids (D) at lumen (L) of active seminiferous tubule (February). Bar = 1 um. The acrosomal vesicle (V) is at the nuclear membrane in 3 spermatids; it has not developed or is not in the plane of section in another spermatid (double arrow). The acrosomal granule (A) has not redistri- buted. Sacs of the Golgi complex (G) are in cytoplasm. Tangential sections of the principle piece of spermatozoa (arrowhead) occur. Part of a Sertoli cell (C) is included Testicular cycles of the Ringtail, Bassariscus astutus 7 | t mn en tn \ornssasenninnneihahenen LUD 3 10 11 12 13 Fig. 6. Testes of Bassariscus astutus. Left: average size non-breeding season; right: average size breeding season Table 2. Weights (g) and linear measurements (mm) of testes Month Weight Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. Bold: maxima; italics: minima. Measurements are means of left and right testes of a pair. Differences between left and right testis vary from 0 to 0.2 g and O to 1.7 mm. Where more than 1 individual per month was available data were averaged. Discussion The physiological capacity to breed is characterized by the mass of the testis and the presence of sperm in the epididymis. The material described in this study clearly indicates an annual developmental cycle of redevelopment and regression of the testes of Bassariscus astutus. At age 16 weeks of the young, testes are tiny, ca. 4 mm diameter when descended and palpable, and from then on remain scrotal (ToweıLL and Toweırr 1978). Although 78 I. Poglayen-Neuwall and J. N. Shively there is no quantitative study, an unknown percentage of young males reach maturity at 10 months of age (PoOGLAYEn-NEuUwALL and POGLAYEn-NEUWALL 1980; POGLAYEN- NeEuwaALL 1987) and thus are able to mate at or near the peak of the mating season. Of 5 yearlıng males, held in the senior author’s colony, 2 have successfully bred; also 2 wild- caught young captured ın March had large testes. Only 20 % of yearlıng male raccoons, according to WooD (1955), possess motile sperm in the Texas post oak region. Among Michigan raccoons many yearling males are capable of reproducing, but females enter estrus about 2 months earlier, and by the time the young males are sexually mature most females are already bred by older males (STUEwER 1943). Active spermatogenesis of B. astutus ıs maximal during the late winter-spring mating season, while it rather abruptly diminishes in June and ceases in July. Our findings show that it takes 1 month plus for spermatogonia to mature to spermatozoa in ringtails, as compared with 20 days in rats (BLooMm and FAwcETT 1962), and 64 days ın humans (Dym 1977). The number of animals we examined resulted in maximum testis measurements for March, with most active spermatogenesis in April. A broader sample most likely would show a closer correlation between the two. We found the degree of testicular regression ın B. astutus to be similar to that ın Procyon lotor (SANDERSON and NALBANDOV 1973; WooD 1955), and not nearly as stıking as in Mustela frenata, whose testes are 1/8 of their maximal sıze during the peak of their non-breeding season (WRIGHT 1947). Variation of testis size/ mass correlated wıth the season is known in many mammalıan species (Amann 1970), e.g. Martes americana (MARKLEY and Bassett 1942), Mustela erminea and Vulpes fulva (AspELL 1964), Dasypus novemcinctus (MCCuskeEr 1985), many cervids (AMAnN 1970), several lemurs (BoRART et al. 1977; PETTER-RoussEAuUx 1972), Sammıri sciureus (Du MonD and HurcHınson 1967), Macaca mulatta (SADE 1964) and Cercopithecus aethiops on St. Kitts (Conaway and SaDE 1969). The latter shows a distinct breeding season (different from that ın Kenya), wıth a testis regression ın the non-breeding season, which ıs much less pronounced than in the aforelisted primates. Sperm were present in the epididymides ot all C. aethiops examıned throughout the year (Conaway and SADE 1969). The tropical Bassariscus sumichrasti, apparently a seasonal breeder, namely from January to May (GAumer 1917; Harr 1981), does not show palpable testicular regression, as observed on 4 captıve adults held by the senior author. No histological study of testes of B. sumichrasti has as yet been undertaken. Likewise Azlurus fulgens, a seasonal breeder, does not show cyclic varıatıon of the testes’ size (]. GITTLEMAN, comm. via g. CONOVER). Electroegjaculation on an adult 2. astutus and an adult B. sumichrasti (conducted by Dr. B. DurrRANT and the senior author ın early May at the San Diego Zoo’s Research Department) produced only minimal volume and very few, non-motile and mutilated spermatozoa. Likewise, electroejaculation on Procyon lotor was unsuccesstul (SANDERSON 1951). There is a correlation between daily sperm production and testicular weight for continuous breeders and for seasonal breeders during the breeding season (ORTAVANT, cit. by Amann 1970). On the other hand MARTINET’S (1966) quantitative, histological studies of Microtus arvalıs revealed seasonality of the testes weight but there ıs no seasonal change in the efficiency of sperm production. Sudden regression of testes during the mating season over a period of only very few weeks and concomitant decrease of active sperm has been noted by us in one newly captured B. astutus, perhaps as a direct result of the trauma suffered, and ın 2 other ringtails probably as a consequence of constant harassment by a dominant female, which likely caused drastic hormonal changes conducive to the aforementioned reproductive condition. These 3 anımals have not been considered ın thıs study. Testicnlar cycles of the Ringtail, Bassariscus astutus 79 Acknowledgements D. Soxor who helped with morphometric and statistical analyses, and H. Russer and J. MaAcMıLLan who made preparations for histopathological and transmission electron microscopic study, deserve thanks for their contributions. We are grateful to Dr. B. DuRRANT for her help with the electroejacu- lation technique. Dr. I. POGLAYEN-NEUWALL gave much of her time, experience and skill toward capture and care of the animals as well as technical assistance. The Arızona Game and Fish Department provided collecting and holding permits, which made this study possible. Zusammenfassung Testicularzyklen des Katzenfretts, Bassariscus astutus (Carnivora: Procyonidae) Mittels Palpierung und morphologisch-histologischen Techniken wurden die Testes von Bassariscus astutus im Jahresablauf untersucht. Jahreszeitlicher Wechsel der Spermiogenese wurde mit Gewicht und Größe der Testes verglichen. Es konnte bestätigt werden, daß die Fortpflanzungsperiode in Arızona von Spätwinter bis ın den Frühling dauert, und daß die Testes im Sommer und Herbst inaktiv (aspermisch) sind. Sie erreichen ihre kleinsten Maße im Herbst. References AMAnn, R. P. (1970): Sperm production rates. In: The testis. Vol. I. Ed. by A. D. Jonnson, W.R. GoMmEs and N. L. VANDEMARK. New York: Academix Press. Pp. 433-482. AsDELL, S. A. (1964): Patterns of Mammalian reproduction. Ithaca, New York: University Press. BAsurTo-Kuga, V. M.; Heath, E.; Wagner, W. A. (1984): Spermatozoa and testes in the boar: correlative analysis of sperm morphologic features, seminiferous epithelial area, and testes weight. Am. J. Vet. Res. 45, 1328-1332. BLoom, W.; FAwcETT, D. M. (1962): Textbook of histology. Philadelphia: Saunders. Pp. 550-583. BORART, M. H.; CooPER, R. W.; BENIRSCHKE, K. (1977): Reproductive studies of black and ruffed lemurs, Lemur m. macaco and L. varıegatus ssp. Internat. Zoo Yb. 17, 77-82. Conaway, C. H.; SaDe, D. $. (1969): Annual testis cycle of the green monkey (Cercopithecus aethiops) on St. Kitts, West Indies. J. Mammalogy 50, 833-835. Du Mono, F. V.; Hurcainson, T. C. (1967): Squirrel monkey reproduction: the “fatted” male phenomenon and seasonal spermatogenesis. Science 158, 1067-1070. Dvm, M. (1977): The male reproductive system. In: Histology. Ed. by L. Weiss; R. OÖ. GrREEP. New York: McGraw-Hill. Pp. 979-1038. Ercock, L. H.; SCHONING, P. (1984): Age related changes in the cat testis and epididymis. Am. ]. Vet. Res. 45, 2380-2384. GAUMER, G. F. (1917): Monografia de los mamiferos de Yucatän. Secret. de Fomento, Dept. Talleres Gräficos. Mexico. GRINNELL, J.; Dixon, ]J. S.; LiNSDALE, J. M. (1937): Fur-bearing mammals of California. Berkeley: University of California Press, Pp. 166-183. Hart, E. R. (1981): The mammals of North America. New York: John Wiley and Sons. Vol. 2, 965-966. Jounson, A. S. (1970): Biology of the raccoon (Procyon lotor varıns Nelson and Goldman) in Alabama. Agrıc. Exp. Stat. Auburn Univ. Bull. 402, 1-148. MARKLEY, M. H.; BassETT, C. F. (1942): Habits of captıve marten. Am. Midl. Nat. 28, 604-616. MARTINET, L. (1966): Modification de la spermatogenese chez le campagnol des champs (Microtus arvalıs) en fonction de la dur&e quotidienne d’Eclariment. Ann. Biol. Annale, Biochim., Biophys. 6, 301. MCCuskER, J. S. (1985): Testicular cycles of the common long-nosed armadillo, Dasypus novemcinc- tus, ın north-central Texas. In: The evolution and ecology of armadillos, sloths, and vermilinguas. Ed. by G. G. MoNnTGoMERY. 255-261. McDoweıt, E. M.; Trump, F. B. (1976): Histologic fixatives suitable for diagnostic light and electron microscopy. Arch. Path. lab. Med. 100, 405-414. Moens, P.; Hu1gEicHoLz, A. (1975): The arrangement of sperm cells in the rat seminiferous tubule: an electron microscopie study. J. Cell. Sci. 19, 487-507. MOLLENHAUER, H. H. (1963): Plastic embedding mixture for use in electron microscopy. J. Stain Tech. 39, 111-114. MoRrTOoN, D.; WEISBRODE, $. E.; WYDER, W. E.; MAURER, ]J. K.; Capen, C. C. (1986): Histologic alterations in the testes of laboratory rabbits. Vet. Path. 23, 214-217. ORTAVANT, R. (1958): Le cycle spermatogenetique chez le belier. D. Sc. thesis, Univ. Paris. PETTER-RousseAux, A. (1972): Application d’un systeme semestriel de varıation de la photoperiode chez Microcebus murinus (Miller, 1777). Ann. Biol. Anim. Bioch. Biophys. 12, 367-375. 80 I. Poglayen-Neuwall and J. N. Shively PoOGLAYEN-NEUWALL, I. (1987): Management and breeding of the ringtail or cacomistle, Bassariscus astutus, ın captivity. Internat. Zoo Yb. 26, 276-280. POGLAYEN-NEUWALL, 1.; POGLAYEN-NEUWALL, I. (1980): Gestation period and parturition of the ringtail, Bassariscus astutus (Lichtenstein, 1830). Z. Säugetierkunde 45, 73-81. SaDE, D. S. (1964): Seasonal cycle in size of testes in free-ranging Macaca mulatta. Folia primat. 2, 171-180. SANDERSON, G. C. (1951): Breeding habits and a history of the Missouri raccoon population from 1941 to 1948. Trans. Sixteenth N. A. Wildlife Conf. 445461. SANDERSON, G. C.; NALBANDOV, A. V. (1973): The reproductive cycles of the raccoon in Illinois. Il. Nat. Hist. Surv. Bull. 31, 25-85. STUEWER, F. W. (1943): Reproduction of raccoons in Michigan. J. Wildlife Mgt. 7, 60-73. TavyLor, W. P. (1954): Food habits and notes on the life history of the ring-tailed cat in Texas. ]. Mammalogy 33, 55-63. Toweıuı, D. E. (1976): Movements of ringtails in Texas’ Edwards Plateau region. Unpubl. MS thesis. College Station: Texas A&XM Univ. Toweırı, D. E.; Toweırr, D. B. (1978): Growth and development of captive ringtails (Bassariscus astutus flavus). Carnıvore 1, 46-53. Woop, J. E. (1952): Ecology of furbearers ın the upland post oak region of eastern Texas. Unpubl. Ph. D. dissert. College Station: Texas AXM. Univ. — (1955): Notes on reproduction and rate of increase of raccoons in the post oak region of Texas. ]. Wildlife Mgt. 19, 409-410. WRIGHT, P. L. (1947): The sexual cycle of the male long-tailed weasel (Mustela frenata). ]. Mammalogy 28, 343-352. Authors’ addresses: Dr. Ivo POGLAYEN-NEUWALL, Department of Ecology and Evolutionary Biol- ogy; Dr. James N. SHIvELY, Department of Veterinary Science, University of Arızona, Tucson, AZ 85721, USA Z. Säugetierkunde 56 (1991) 81-87 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Sexual bimodality in some recent pig populations and application of the findings to the study of fossils By J. VAN DER MADE Institunt voor Aardwetenschappen, Rijksuniversiteit Utrecht, Nederland Receipt of Ms. 9.4. 1990 Acceptance of Ms. 21.9. 1990 Abstract Studied the size differences in some skull and tooth measurements between males and females in recent Suidae. In three populations of Sus the linear measurements of the cheek teeth of the females (0 < n <= 25) are on average 97 % those of the males (O<n= 28). There ıs much overlap between the sexes. In Babyrousa such size differences are of the same order of magnitude. In an earlier study, using skull length, greater differences between the sexes were found. The value of skull length and some other skull measurements is discussed. Male Suidae have larger diastemas and longer skulls because of their larger canınes. The results of this study are applied to a sample of fossil Bunolistriodon from Cörcoles (Guadalajara, Spain). The material is morphologically homogeneous, but there are two size groups differing on average by 15 %. There is no overlap between the groups. It ıs concluded that it is more likely that the material from Cörcoles represents two species rather than males and females of the same species. Introduction A morphologically homogeneous collection of Bunolistriodon tossıls from the Lower Miocene locality of Cörcoles (Guadalajara, Spain) consists of two sıze groups, dıffering by 15 %. The question arose whether these groups represent two species or males and females of the same species. Usually sexual dimorphism in Suidae is apparent in the canınes, but this crıterion could not be used due to the state of preservation of the fossils. The present dilemma is not an ısolated one in the study of fossıl pıgs (PICKFORD and WILKInson 1975). In some families sympatric morphologically sımilar species differ in size by 15 % (recent peccaries: Ruscon1 1929; Dorcatherium: MoTTL 1961; FAHLBUSCH 1985), whereas in other families males and females of one species ditfer by up to 13 % (Ursidae: KuURTEN 1969; TORRES 1984). This paper deals with the size differences between males and females in recent pigs. The cheek teeth were measured as it is the intention to apply the findings to the study of fossils. In addition some skull measurements were taken. Material and methods Three collections of recent species, in which the canines indicate the sex of the specimen are studied. These collections represent approximately three populations, because they were made in a small area during a relatively short time. Not all measurements could be taken in all skulls, because of teeth that are not yet erupted or because of damage. The number of measurements taken are given in the tables. The largest population measured is one of Sus scrofa vittatus from the former residence Delı, Sumatra, now called Sumatera Utara. The collection is stored in the Zoological Museum of Amsterdam (ZMA). The lower cheek teeth were measured and several skull measurements were taken. Males and females are about equally represented. The males are on average younger, in many the M; is not yet erupted or the talonid is still partly covered by bone (such individuals are considered to be subadults). U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0081 $ 02.50/0 82 J. van der Made Fig.1. The way of measuring. Diastemas and distances measured as the shortest distance between alveoli: 1. diastema C-P;; 2. dıastema P,-P>; 3. distance C-P>; 4. distance P> - front of canıne; 5. distance I;-P>; 6. DT, the greatest width (either measured at the anterior or at the posterior lobe) and 7. DAP, the greatest length of a tooth. (Measurements are in mm) A smaller sample of Sus barbatus from east Bulongan, Borneo in the ZMA consists mainly of skulls without mandibles. Sus verrucosus was measured in the ZMA and at Leyden in the Rijksmuseum voor Natuurlijke Historie (RMNH). Only specimens with a label indicating the provenance of Java were used. The females are underrepresented. No attempt has been made to measure all male material, which is very abundant. Most of the females of this species in the RMNH, also those which do not have a label specifying the provenance, do not have a fully erupted M;. But most of the males are fully adult. In addition data on recent Babyrousa from Bumbulan (females 3<n=5, males 12 <n< 13) and Buru (females 2<n= 3, males 13 <n < 16) were taken from GrovEs (1980). The fossil Bunolistriodon material (0 <n = 5) will be compared to the recent material. It is stored in the geology department of the Universidad Complutense in Madrid and it is under study by V. D. MADE and ALFEREZ. Measurements of diastemas and the length of the skull are given for Sus scrofa only, as it ıs the largest sample. Length and width of the cheek teeth and length of diastemas were measured as indicated ın figure 1. Skull length was measured in the median plane from the tip of the nasalia to the posterior edge of the occiput. All measurements are in mm. The value Q is introduced to express the mean size of a character of the females relative to the mean size of the males. It is given as a percentage. Q = (mean females/mean males) x 100 %. Results Average lengths and widths of the cheek teeth of Sus are given in table 1. On average females appear to be only slightly smaller than the males. Q values show little varıation in the large Sus scrofa sample (92 < Q = 101), the average value is 97. Q values in the Ss verrucosus sample show more variation (88 < Q= 123), the average value is 98. The average Q value for Sus barbatus is 97 (87 <Q=< 105), for Babyrousa from Buru 95 (93 <Q = 99) and for Babyrousa from Bumbulan 101 (94 < Q = 106). Ihe average Q value for all populations is 98. Although the only sample in which females are well represented is that of S. scrofa, it is striking that in none of the other samples the average Q ” 83 Sexual bimodalıty in some recent pig populations umu ur SJUOWSANSTON "UONeI SJUSWMDANSEIW Fo Jaquınu = U !yIpım 15998913 = LA ‘y23ua] 15918913 = IVA snS JO 4393) Y99y9 9yJ Jo suorsuswip SIe.I9AY ‘I aJgPL sojeut LA soJruls} A2L4 sojeul dvd sopeu} AVA snivqavq SNS soJeul AlL@l sofeuls} Lq4 sopeut IVA sofeuls! dvd SNSOINAAIR SNS sofeuu Ldq4 soJeuls] LA sopeu VA soTruls} dvd snivq4vq sns sapeu LA sopewsl LA sopew IJVA AAN SNSOINAAOR SNS sopew LA sopews} LA A RN sapeus] IJVA snvıma vfo4s snS J. van der Made 54 ee) (sm) £6 29 sıynpe & 76 6L sınpegns Ö GG - (way ıpnpe/'woJ Ipnpegns) x 00T c6 +8 (sapeur ıpQnpe/safeur ıpnpeqns) x 001 sueaW Ne) N [oo] Ne) — UWINUIIXPUI uwsu (Isuss3) UINUITUTLU sojeuu ı7npV che Orr 0743 SaLE 66C 6°C Ice RE 86C ES ILILG CSc I SITE L6C 0’Sc OLC 0’Eel Sec SIT GLC GET LSC EE UINWUIIXTUI uwsu (I su=j7]) UINUITUTUI ıpnpeans UIMUITXPUI uesw („[SUusH#]) UINUITUTUI sofeuls] ı7npV UINUITXPUI Ur9WI UINUITUTUU ı[npeqns OT EN HEIDEN oOoliwn SüN “no un ya3uay [nyS <4 - D zuorg 9IUPISICT SnJEJJIA EJOIIS SNS JO (uruı ur) syJ3ua] ny7s pure seursIseip IyJ JO SUOISUSUNCT 'Z 2JqYL Sexnal bimodalıty in some recent pig populations 85 value differs much from the value of this sample. The smaller size group in the Bunolıis- triodon material averages 85 % of the larger sıze group. Size overlap between the sexes of Sus scrofa vittatus is great, in none of the check teeth two concentrations are formed (figure 2). This ıs also the case in Sus verrucosus and Sus barbatus. The size groups are better separated in sımilar diagrams for Bunolistriodon. The size of the dıastemas ıs about twice as large in the males as ın the females (table 2). Also the total distance between the last incisor and the P, ıs much larger. In a sımilar way subadults of each sex have lower values than the adults, although there are some exceptions. The Q value for skull length is 94 % (table 2). DAP 7 SIE ENEDT DES AAES1ST 14 15 16 17 18 DT Fıg. 2. Scatter dıagrams for Sus scrofa vittatus from Delıi. In the diagram for the M; a particularly large molar of which the complete width could not be measured is also given. (Measurements are in mm) 86 J. van der Made Discussion The cheek teeth in $Sus scrofa females are 97 % the size of the cheek teeth in the males. As the smaller samples do not differ in a substantial way (their average Q values vary from 95 to 101) the value of 97 % seems to be representative for the check teeth. Average size ditferences between the sexes of 3% only, seem to contradict the findings of Grovss (1981). He found that females averaged 85.4-95.3 of the males, 80.8-83.3 in Sus verrucosus. GROVES’ values are based on the condylobasal lengths of the skulls. These measurements are roughly comparable to the skull length as measured in this study. The skull length includes the part with the canines; these zones are highly sexually dimorphic. The large Sus scrofa sample will be used to show how sexual dimorphism in the canine area causes a bimodality in skull lengths. In the Deli sample the mean distance between I; and P; is 8.4 mm greater in the adult males than in the adult females; the difference for the skull length is 23 mm. If the males had no enlarged canınes and diastemas their skulls would have an average length of 312 mm. The average female skull measures 297, which ıs 95 % of 312 mm. The dentition indicates a value of 97 %. So a large part of the sıze difference in female and male skulls is accounted for by the large canine zone in the males. In $. verrncosus diastemas tend to be much longer than in S. scrofa and in the small $. verrucosus sample size differences of the P>-I; distance is 22.8 mm, which is much greater than in $. scrofa. So, a greater sexual bıimodality in skull length is to be expected in S. verrucosus, although sıze differences in cheek teeth are not greater than ın S. scrofa. Part of the size difference for Sus verrncosus found by Grovzs (1981) might be explained in this way, but the difference in length of the M3 for S. verrucosus verrucosus, as listed by hım, is large: temales average 86 % of the males. My measurements indicate 96 % for the length of the M;. There is no explanation for this discrepaney. It should be noted that for $. verrucosus blouchi the same value is 95 %. Also the standard deviation for the M, of the females of S. v. verrucosus, as given by GroveEs (1981), is large. In any case, ıf an indication of size ıs needed for taxonomical purposes it is advisable to use a measurement that ıs less variable than skull length. Sexually dimorphic canines influence both skull length and diastema length. Diastema length ıs used by some authors for taxonomical purposes (THENIUS 1972). The difference in size of the diastemas in the females and males is related to the size of the canınes. The larger the upper canines the larger are the diastemas behind the lower canınes, and the same reasoning applies to the lower canines and the diastemas in front of the upper canınes. As the canınes of the males keep on growing (also their antero-posterior and transverse diameters increase) the diastemas have to grow too: the subadult/adult values for the distances C-P>, front of canıne - P, and ];-P> are lower in the males than in the females (table 2). The continued sıze increase of the canınes in the males seems to cause a decrease of the C-P| and C-J]; distances, indicating that bone of the mandible is resorbed. The subadult/adult ratıo for the P-P; (too high) and P,-C (low) distances in the females is caused by a large number P} in contact with the P; in the adult females. The position of the P, seems to be varıable. The value of the size of the diastemas for taxonomy is restricted, as this character ıs related to sex and age and is influenced by the variable position of the P,. If it ıs necessary to indicate the size of diastemas, it seems better to give the distance I3-P2 or the distance from the front of the canine to the P; in the mandible or the distance from the P! to the I° or I” in the upper jaw. In the studied species the size difference between the males and females is small; ın their direct ancestors this is probably also the case. Species of other subfamilies than Suinae are only known as fossils, so a study of this kind can be done for the Suinae only. The fact that size variation in large fossil samples is not greater than in recent species is an indirect Sexnal bimodalıty in some recent pig populations 87 indication that large sıze differences between the sexes in fossil Suidae do not occur. Size varıation in Suoidea will be subject to a separate study. Scanty fossil material of which the sex is known also indicates that there are no great sıze differences between females and males. The presence of two species of Bunolistriodon ın Cörcoles instead of merely two sexes of one species, is likely because ot: 1. The average size difference between the two size groups in Bunolistriodon from Cörcoles is 15 %, which is much more than the 3 % of average size difference found in recent male and female Suidae. 2. The size groups are separated and do not have a large overlap as males and females of extant species. Acknowledgements Ithank Drs. F. ALFEREZ, P. J. H. van BREE and CH.R. SmEENK for allowing me to study material and Dr. P. J. H. van Bree, Dr. P. Y. SONDAAR, Dr. J. DE Vos and Mr. T. LELIEvELD for reading the manuscript critically and discussing the matter. Zusammenfassung Geschlechtsbimodahtät bei einigen rezenten Schweinepopulationen und Anwendung der Befunde für das Studium an Fossilien Diese Studie beschäftigt sich mit den Größenunterschieden zwischen weiblichen und männlichen Schweinen. In drei rezenten Populationen von Sus betragen die linearen Maße der Backenzähne der Weibchen (O<n= 25) durchschnittlich 97% derjenigen von Männchen ((<n=28). Bei Baby- rousa liegen die Unterschiede in gleicher Gröfsenordnung. In einer früheren Untersuchung wurden bei Bezug auf die Schädellänge größere Unterschiede zwischen den Geschlechtern gefunden. Diese Studie stellt den Wert von Schädellängenmessungen und anderen Maßen zur Diskussion. Männliche Suidae haben längere Diastemen und längere Schädel wegen ihrer größeren Canınen. Die Ergebnisse dieser Studie wurden auf eine Probe von Fossilen Bunolistriodon von Cörcoles (Guadalajara, Spanien) angewandt. Dieses Material ist morphologisch homogen, aber in bezug auf die Größe können zwei Gruppen unterschieden werden. Der Unterschied beträgt 15 %, und es gibt keine Überschneidungen. Daraus wird geschlossen, daß das Material von Cörcoles eher zwei Arten repräsentiert als die Geschlechter derselben Art. Literature FAHLBUSCH, V. (1985): Säugetierreste (Dorcatherium, Stenofiber) aus der miozänen Braunkohle von Wackersdorf/Oberpfalz. Mitt. Bayer. Staatsslg. Paläont. hist. Geol. 25, 81-94. Grovss, C. (1980): Notes on the systematics of Babyrousa (Artiodactyla, Suidae). Zool. Med. 55, 2946. — (1981): Ancestors for the pigs: taxonomy and phylogeny of the genus $us. Technical Bull. 3, Dept. Prehistory, Res. School of Pacific Studies, Australian National Univ., Canberra. 3. printing 1985, 1-96. KurteEn, B. (1969): Sexual Dimorphism in Fossil Mammals. In: Sexual Dimorphism in Fossil Metazoa and Taxonomic Implications. Ed. by G. WESTERMAnNn. Stuttgart. 226-233. MorTL, M. (1961): Die Dorcatherien der Steiermark. Mitt. Mus. Bergb. Geol. Techn. 22, 21-71. PıckForD, M.; WıLkınson, A. (1975): Stratigraphic and phylogenetic implications of new Lis- triodontinae from Kenya. Neth. ]J. Zool. 25 (1), 132-141. Rusconiı, C. (1929): Anatomıa craneo-dental de los tayassuinos vivientes (pecaries). An. Soc. Dient. Argentina. 107, 1-75. THEnIUs, E. (1972): Microstonyx antiguus aus dem Alt-Pliozän Mittel-Europas. Zur Taxonomie und Evolution der Suidae (Mammalia). Ann. Naturhistor. Mus. Wien 76, 539-586. TORRES P£EREZ-HIDALGOo, T. DE (1984): Ursidos del Pleistoceno-Holoceno de la Peninsula Iberica. Thesis doctoral, Madrid. Author’s address: JAN VAN DER MADE, Instituut voor Aardwetenschappen, Faculteit Geologie en Geofysica, Rijksuniversiteit Utrecht, Postbus 80.021, NL-3508 TA Utrecht, Nederland Z. Säugetierkunde 56 (1991) 88-93 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Dominance relationships and competition for resources among chamois Rupicapra rupicapra rupicapra in female social groups By P. IncoLp and H. MARBACHER University of Berne, Zoology Department, Ethological Station Haslı, Switzerland Receipt of Ms. 10. 4. 1990 Acceptance of Ms. 30. 10. 1990 Abstract Studied in a region ın the Bernese Oberland (Switzerland) which anımals in social groups of female chamois Rupicapra rupicapra rupicapra are able to assert themselves in competition for resources (licks, resting and feeding sites), and how losers behave after an interaction. The older animals regularly asserted themselves against younger individuals. Within the class of adult females (with or without kids) certain anımals regularly asserted themselves against others. The most striking behaviour on the part of the loser, after having been forced to give up a resource to which it had previous access, was urınation directly following the interaction, as a result of the latter and as a possible sign of stress. Introduction Outside the rutting season, which occurs in November/December, chamois live separately with regard to sex. The female chamoıs form social groups consisting of adult and subadult females, yearlings (including males) and kids (KRÄMER 1969; Knaus and SCHRÖDER 1983). Even in nonhunted, local populations (e.g. in game reserves) the number of anımals remains faırly constant over a longer period of time, migration occurring only very rarely (KRÄMER 1969; Knaus and SCHRÖDER 1983). These findings aroused the suspicion that social processes could play an important role in limiting population density (KRÄMER 1969). Later on, some ıindications supported this suspicıon (BERDUCOU and Bousses 1985; Lovarı and Rosto 1985). LovAarRI and RosTo (1985) showed, that younger female chamois had a lower bite rate ın the presence ot older (dominant) females, suggesting, that in the long run that could be a reduced food intake of young females, influencing e.g. winter survival and reproductive processes. The aim of the present study was to investigate one aspect of this problem, ın particular: 1. whether certain individuals assert themselves in the competition among licks, resting and grazing sites; 2. how the behaviour of losers is influenced by the winners of an interaction. Study area and anımals The study was carried out in the Swiss Federal Game Reserve Augstmatthorn in the Bernese Oberland (Switzerland) in summer 1987. The study area lies on the north-west slope of the Augstmatthorn at a height between 1500-2100 m altitude. It covers a surface area of approximately 3 km. It was possible to observe a maximum of 126 predominantly female chamois; approxımately 61 adult and 26 subadult females and yearlings as well as 39 kids. Anımals were allocated according to age classes, using as criterion the length of their horns in relation to ear length (Lovarı 1985; PATTERSON 1988). Males (except for male yearlings) were seldomly observed ın this area. Every day many animals visited the apparently very attractive licks including 3 optimal licks, the remainder being less attractive (MARBACHER 1989), which were spread over a small area. The fact char, as a rule, there were considerably more anımals than licks led to very strong competition at these sites. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0088 $ 02.50/0 Dominance relationships and competition for resources among chamois 89 Methods The chamois were observed from May to October at a total of 90 days. Observations were made each day from morning to evening using the “behaviour sampling” method (Ar rmann 1974). Aggressive interactions (e.g. advancing with lowered horns towards conspecifics) in connection with competition for resources, occurred at the few available licks, at resting sites and during grazing. In order to classify which individuals were able to assert themselves, the numbers of anımals remaining compared to those leaving were recorded, during situations when an older anımal approached a younger one within three body lengths or vice versa at the licks, at resting and grazing sites. The Fisher exact test was used for statistical analyses for a total of 250 events. It was possible on two occasions to observe two groups of four adults with kids (individually identifiable by morphological features such as colour of coat) at the licks during a period of several hours and to determine which anımal drove the other away. The winner of an interaction took over the resource or continued to use it and the loser retreated a few meters, stopped and often began to urinate. In order to determine whether urination was indeed a result of having been driven away, we then proceeded as follows: We recorded the time until any urination occurred over a period of 5 minutes, a. after an anımal had been forced to leave a site, b. after an anımal had left a corresponding site on ıts own accord. The comparison was made with the Mann/Whitney U-Test. Results Winner and loser in the competition for resources At licks: In all combinations of the 4 age-groups there were significant differences in the number of anımals leaving to those staying at the approach of an other anımal (Table 1). Almost without exception the younger anımals retreated when an older one approached, while in the opposite case the older ones stayed. Among the ındıvidually recognizable adult females with kids, the same anımals always asserted themselves in the course of a series of interactions (except ın the case of anımals E and F; Fig. 1), ı.e. the subdominant anımals had to “ınvoluntarily” leave the lick together with their kids (Fig. 1). At resting sıtes and when grazing: In the combinations studied the result was the same as at the licks. In all combinations studied, younger anımals retreated at the approach by older individuals but not vice versa (Table 1). In some cases certain adult females with their kids retreated from other adult females. Urination by the loser as a consequence of the interaction with a dominant animal The most striking behaviour of an anımal that had been driven away was urination after it had retreated a few meters. Anımals driven away from licks and when grazing urinated in about 95 % of the cases, those driven away from resting places in slightly less than 90 %. Those not driven away, if they had left the corresponding place “voluntarily”, urinated in 79 %, 52 % and 56 % of the cases. The difference between anımals driven away and those not driven away when grazing and at resting sites was significant (Table 2). The lack of a difference at licks was clearly due to the fact that animals that had been licking for some time as a rule urinate upon leaving the lick. In all three situations, however, urination occurred significantly earlier in cases where anımals had been driven away than in voluntary departures, ı.e. within a few seconds (Fig. 2). Discussion Competition does not only occur at the few attractive licks, but also at resting sites and during grazing. Older animals in general tend to assert themselves in certain situations against younger ones, as has already been mentioned elsewhere (KRÄMER 1969; LOvARı 90 P. Ingold and H. Marbacher Table 1. Frequency of animals (belonging to different age classes) winning or losing in competi- tion for licks, resting and feeding sites OF = old female; YF = young female; YA = Yearling; sK = strange kid; K = kid; a = anımal at the site; b = anımal approaching; ns = not significant (Fisher exact test) anima ieaestchesshite remains retreats at licks (0) 12 at resting sites OF YF 3 YF OF (6) OF YA 6 YA OF (6) OF sK 7 sK OF (6) (6) 2 (6) 7 0) U at feeding sites OF YF 1 YF OF ) OF YA 6 YA OF ) OF sK 7 sK OF 0) YF YA 2 YA YF 0) YA K 4 K YA 0 OMooN! nS OMOOnl! OXoonl! nS ORoo! OR on! "ns and Rosto 1985). However the same evidently also occurs among older females, and in particular among anımals whose age difference cannot be readly determined (on the basıs of the length of their horns). The kids of the respective mothers shared the advantage or disadvantage. For example, kids of dominant mothers could use a lick unhindered during the period that their mother used the resource and conversely kids of subdominant mothers were forced to leave a lick together with their mother. Older animals and those dominant within their class of adults were thus privileged in the utilization of resources. At licks: The best licks (with respect to humidity, access, see Dominance relationships and competition for resources among chamois 91 2 3 4 AL a & 6 ns © 10 1 & Fig.1. Sociograms showing the number of successful approaches of adult females to other adult females staying at licks. A-H: Individually identifiable females with kids Table 2. Number of animals urinating after leaving a lick, a resting or a feeding site after (a): it had been driven away or (b): it had left the site on its own accord; (+) = urination occurred; (-) = urination did not occur; ns = not significant atmulieks at resting at feeding sites sites (+) (3) ) 30 chi-square P SEC 300 xx XXX XXX 250 200 x*** =p< 0.004 = Kaas 020 100 50 ! n D 1 2 i D ae lieks at resting at feeding sıtes sıtes NE aa 20 19 13 16 18 Fıg. 2. Time until urination after an animal had been driven away or after it had left on its own accord at the licks, resting and feeding sites. Median, first and third quartil, minimum and maximum; 1 = driven away, 2 = left on its own accord (Mann/Whitney U-Test) 92 P. Ingold and H. Marbacher MARBACHER 1989) were mainly occupied by adult females with kids and as far as could be determined also by certain adult individuals without kids. At resting sites: Dominant animals occupy and retain desirable resting sites (e.g. shady sites during summer or at the very limited areas remaining snow in late spring), in contrast to subdominant animals. When grazıng: Chamoıis eat selectively and have a particular preference for certain plants (Knaus and SCHRÖDER 1983; FERRARI and Rossı 1985; Leon 1985). The fact that anımals, after having driven away another one, continued eating at exactly the same place suggests that they fed on plants discovered by the previous anımal. In individual cases this could be clearly ascertained. The distances between the chamois were small (a few body lengths on average; MARBACHER 1989) so that it can be assumed that they could see what kind of food their neighbour had been eating. It may be a question of special tactics on the part of the dominant anımal, ı.e. to keep an eye on what subdominant anımals have found and to appropriate this at the expense of the others, as ıs said to be the case with Brent geese (DRENT, pers. comm.). It is quite possible that when grazing, subdominant animals are at a disadvantage, particularly when large areas are covered with snow or when snow remains for a long time. It ıs striking that at the licks but also in a herd of grazing anımals, short agonistic interactions occurred almost continuously. In such cases mostly younger, but also certain adult anımals in the social group gave up resources, some of which they had discovered themselves and others that were well known resources (licks). Therefore they were probably almost always under a certain stress. The regularly occurring urination of an anımal driven from a resource could be, in our opinion, the sign of such stress (some farm anımals show enhanced defecation and/or urination under the influence of certain stres- sors, e.g. WooD-GusH etal. 1975). In any case it could be of importance as an indication of _ ıinferiority, as described in mountain sheep (Geist 1971). Nevertheless, ın the latter species it ıs thought to be a kind of submissive behaviour which prevents attacks from dominant animals. We have no indication for this theory in the present study. The dominant anımal seems to be fixed on the resource, and pays apparently very little attention to the subdominant anımal that has been driven away. There were virtually no observations of behaviour towards the urinating anımal or to the urine itself (e.g. snitfing) wıthin the herd of largely female anımals studied (the only males were yearlıngs ın this group). More or less strongly pronounced stress leads to physiological responses which may change the function of the immune system, increasing susceptibility to disease (explored ın some laboratory and farm animals, e.g. FRASER and BRooM 1990). Within social groups of female chamois ıt appeared that the social processes of competition for resources could have such an effect, especially under the circumstances of high population density. In order to examine this supposition, long-term studies on individually marked anımals would be necessary. Acknowledgements We thank the Game Department of the Canton of Berne, especially Dr. H. BRÜLLHARDT and M. ZUBER, for the cooperation and for providing accommodation on the Lombachalp. Preparation of the manuscript was assisted by OLıvIa LAssIERE and REINHARD SCHNIDRIG. Zusammenfassung Dominanzbeziehungen und Konkurrenz um Ressourcen in Gruppen von weiblichen Gemsen (Rupicapra rupicapra rupicapra) In einem Gebiet im Berner Oberland, Schweiz, wurde im Sommer 1987 untersucht, welche Indivi- duen in Gruppen von vorwiegend weiblichen Gemsen (Rupicapra rup. rup.) sıch in der Konkurrenz um Ressourcen (Leckstellen, Liegeplätze, Nahrungsorte) durchzusetzen vermögen, und wie sich die Verlierer im Anschluß an eine Auseinandersetzung verhalten. Generell setzten sich ältere Tiere gegenüber jüngeren Tieren durch und innerhalb der Klasse der adulten Geissen mit Kitzen regelmäßig bestimmte Tiere gegenüber andern. Dominance relationships and competition for resources among chamois 93 Auffälligstes Verhalten der Verlierer, welche eine durch sie gehaltene Ressource aufgeben mußten, war Harnen unmittelbar nach der Auseinandersetzung als Folge davon und als möglicher Ausdruck von Streß. References ALTMANN, J. (1974): Observational study of behavior: sampling methods. Behaviour 49, 227-267. BERDUcCOU, C.; Bousses, P. (1985): Social grouping patterns of a dense population of chamois in the western Pyrenees National Park, France. In: The biology and management of mountain ungulates. Ed by S. Lovarı. London: Croom Helm. 166-175. FERRARI, C.; Rossı, G. (1985): Preliminary observations on the summer diet of the Abruzzo chamois (Rupicapra rup. ornata Neum.). In: The biology and management of mountain ungulates. Ed. by S. Lovarı. London: Croom Helm. 85-92. FRASER, A. F.; BRoom, D. M. (1990): Farm anımal behaviour and welfare. 3. ed. London: Bailliere Tindall. Geist, V. (1971): Mountain sheep. A study in behaviour and evolution. Chicago: Univ. Chicago Biress: Knaus, W.; SCHRÖDER, W. (1983): Das Gamswild. 3. Aufl. Berlin, Hamburg: Verlag Paul Parey. KRÄMER, A. (1969): Soziale Organisation und Sozialverhalten einer Gemspopulation (Rupicapra rupicapra L.) der Alpen Z. Tierpsychol. 26, 889-964. LEonı, G. (1985): Nahrungswahl des Steinbockes auf alpinen Silikatrasen bei Davos mit Vergleich zur Gemse. Dipl.-Arbeit, Geobot. Institut. ETH Zürich. Lovarı, $.; RosTo, G. (1985): Feeding rate and social stress of female chamoıis foraging in groups. In: The biology and management of mountain ungulates. Ed. by S. Lovarı. London: Croom Helm, 102-105. MARBACHER, H. (1989): Soziale Organisation und Konkurrenz in einer Herde jungeführender Gemsgeissen. Dipl.-Arbeit, Ethol. Station Haslı, Universität Bern. PATTERSON, 1. J. (1988): Responses of Appenine chamoıs to human disturbance. Z. Säugetierkunde 53, 245252. Woop-GusH, D. G.M. et al. (1975): Social stress and welfare problems in agricultural anımals. In: The behaviour of domestic anımals. Ed. by E. $. E. Harez. London: Bailliere Tindall. 182-200. Authors’ address: Prof. Dr. P. IncoLD and lic. phil. nat. H. MARBACHER, Zoology Department, Ethological Station Haslı, University of Berne, Wohlenstr. 50a, CH-3032 Hinter- kappelen, Switzerland Z. Säugetierkunde 56 (1991) 94-100 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 A contribution of the ecology of the Steppe pangolin Manis temmincki in the Transvaal By N. H. G. JacoBseEn, R. E. NEWBERY, M. ]. DE WET, P. C. VıLJoEn and E. PIETERSEN Directorate Nature and Environmental Conservation, Pretoria and National Parks Board, Skukuza, South Africa Receipt of Ms. 9.4. 1990 Acceptance of Ms. 27.9. 1990 Abstract Various aspects of the ecology of the Steppe pangolin in the Transvaal are discussed. These include distribution outside the Kruger National Park, activity times and movement. Food and feeding are discussed as well as aspects of reproduction and growth. Mortality factors and Management problems, indicate the necessity for greater in-depth studies of these enigmatic anımals. Introduction The Steppe pangolın Manis temmincki is probably one of the least known mammals in Africa in spite of its faırly extensive distribution from Sudan in the north to the northern Cape Province and Orange Free State in the south. Various authors (SMITHERS 1971, 1983; Dorst and DAnDELOT 1970; Kınapon 1971; STUART 1979; RAUTENBACH 1982; COULSON 1989) have recorded the distribution and aspects of habits and feeding. Van EE (1966) recorded the first successful breeding of this anımal in captıvity. An extensive account concerning the biology of pangolins including Manis temminckü was written by E. MOHR (1961). Despite these accounts the Steppe pangolın largely remains an enigma. This paper contributes towards the biology and ecology of the species in the Transvaal. Materials and methods During the period 1977 to 1983, 32 records of pangolins seen by farmers and Nature Conservators were examined by the senior author. In many cases the anımals were confiscated from members of the public and returned to the sites of origin. In two cases the pangolins were rehabilitated at sites far from their original haunts with no success. These rehabilitations were monitored using transmitters. A transmitter was also used to follow a single female on several farms in the Thabazimbı district for a period of two weeks. The transmitters used in all cases were SM1 type (AVM-Instrument Co.), and LA12 receivers (AVM - Instrument Co.) were used in tracking. The radio transmitters were attached using fibreglass resin and steel (0,008 gg) wire. Four small holes were drilled through one of the large posterior dorsal scales (near the root of the taıl). Two small loops were made with the wire which then held the transmitter to the scale. Liberal application of resin attached the transmitter firmly to the scale and gave excellent waterproofing. The trailing antenna was made from steel guitar wire (0,008 gg) and 45 cm long (Y4 wave length, frequency 148 mHz). Tracking was done on foot using hand-held yagı antennae. Monitoring of the rehabilıtation animals started at dusk while that of the free ranging female continued as the anımal became active, both day and night. During the tracking of this female, sites visited and fed at, were examined and the prey collected. These were then identified. The pangolins did not react to torch light and carried on foraging unconcernedly, allowing a relatively close monitoring of movement. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0094 $ 02.50/0 Contribution of the ecology of the Steppe pangolın ın the Transvaal 35 Results and discussion Distribution The Steppe pangolin is widely distributed in the Transvaal being found in the bushveld areas of the western, northwestern, northern, north-eastern and eastern Transvaal. It ıs obvious from the frequecy of sightings that the western and north-western Transvaal are the stronghold of the species outside of the Kruger National Park, (Figure 1). Most sightings (69 %) were made between May and October possibly because it is easıer to find them when the vegetation cover ıs least. Fig. 1. Distribution of the Steppe pangolin Manis temminckü in the Transvaal, outside of the Kruger National Park during the period 1977 to 1983 Size and mass Steppe pangolins in the Transvaal, are relatively small mammals reaching 1,3 m in length (nose tıp to taıl tip) but are most often between 0,7-1,0 m in length (Table 1). Males appear to be heavier and larger than females and there is a linear correlation between mass and length (Figure 2). 96 N. H. G. Jacobsen, R. E. Newbery, M. ]J. de Wet, P. C. Viljoen and E. Pietersen 14_ Mass (kg) 550 650 750 850 950 1050 1150 1250 1350 Length (mm) Fig. 2. Correlation between mass and length in the Steppe pangolin Table 1. Mass and size of Transvaal pangolins IL = Total length Mass (kg) Females 10.09 Mean = 6.69 Males 9.15 9.60 1345 Mean = 10.63 Juveniles Habits Steppe pangolıns are generally recorded as being nocturnal and partıially diurnal anımals (SMITHERS 1983; DorsT and DAanDELoT 1970; KıncDon 1971). According to recorded sightings and while radio tracking these anımals ıt ıs apparent that they are crepuscular but forage into the nıght and even into the early hours of the morning, while also being found in broad daylıght especially during the late afternoon. A summary of 17 pangolın sightings Contribution of the ecology of the Steppe pangolin in the Transvaal IN. by various observers in the Transvaal over a period of sıx years indicate a hıgh frequency of activity (56 %) between 16h00 and 18h00. The remaining anımals were observed at various times between midnight and midday. An adult female translocated from the Vryburg District ın the northern Cape Province and released on the farm Elandsfontein in the Thabazımbı District, Transvaal remained strictly nocturnal and moved about between 18h30 and 24h00, over a period of 10 days. Another female followed over a period of 18 days, inıtially followed a similar pattern of activity but changed its activity pattern repeatedly, becoming active ın the early afternoon and sleeping by early evening (20h30). The Steppe pangolın in the Transvaal spends these resting periods ın available cover which ıs usually under a large rock, pile of stones, litter and vegetation. It has been suggested by BIGALkE (1932) and MoHR (1961) that these anımals dig their own burrows as some of the Asiatic species do but this does not appear to be so. The only burrows used for shelter are those of antbears, Orycteropus afer and springhares, Pedetes capensis and ın some cases hollowed out termitaria. These anımals move over large distances inspite of their apparently clumsy mode of movement. The translocated female moved a mınımum of 18.1 km over the 10 days of tracking. This averages 1.81 km day but as the anımal was not closely followed this figure could be considerably larger. However, it agrees closely with the 50 m/minute recorded by BIGALkE (1932) for the Steppe pangolın in captivity. Similarly the other radio-tracked anımal which was released where found, appeared to travel on average a minımum of 0.7 km per day but as ıts meanderings were not measured, this could conceivably be as much as the first. As the translocated anımal appears to have behaved atypıcally, more emphasıs ıs placed on the anımal released where ıt was found. This particular pangolin, a female travelled from the point of release up the side of a hill to the crest where it spent the next 10 days feeding and meandering about over a very small area (Fig.3). Over the next fıve days ıt moved along the crest of the hill to another site where she spent a mınımum of three days whereafter tracking was stopped. Another 18 N, EB site of capture,tagging 2 and release A pangolin route contour interval = 30,4 m & ee N Fıg. 3. Movements of a resident Steppe pangolin over a period of 18 days. The shaded areas 1 and 2 are where the pangolin foraged over 11 and a minimum of 3 days respectively 98 N. H. G. Jacobsen, R. E. Newbery, M. J. de Wet, P. C. Vihoen and E. Pietersen days later the signal from its transmitter was located in the close vicinity of where it was last seen. It is apparent that the anımal used a system of bases from which it foraged, obviously locating sufficient food on her excursions. In fact she was seen to forage in places where she had previously foraged. The foraging excursions were of relatively short duration ranging from five to six hours. When she went to ground she chose sites under and among boulders and stumps and in holes, on occasions using the same holes as she had on previous occasions and on other occasıons new hiding places. One such site was used for seven days with the anımal returning to it after each foraging excursion. On one hot day (28 March 1981) the female was observed to lie up in the shade of some small trees and rest. It was then observed to lie on its back and expose its bare belly. This was moist, whether from sweat or urine and permitted the anımal to cool off. Such stops occurred on several occasions thereafter, lasting from a couple of minutes to as much as 1% hours. It was noticed that all pangolins kept ın captivity urinated on themselves squirting urine in short bursts. This obviously has a cooling function but also permits the anımal to advertise its presence to other pangolıns as ıt moves about an area. This was confirmed while following the female on the farm Kansberg, where she was seen to leave wet marks on the rocks which she traversed. In addition, the urine has a pungent odour which is recognisable by humans and therefore even more pronounced to the smell sensitive pangolin. Foods and feeding Pangolins forage by walking slowly along sniffing ın likely places. Once a nest of ants or termites has been located the pangolin rips open the nest with the short powerfully armed front feet. The exposed holes are sniffed at extensively and the tongue is only extended down those holes where there are large quantities of these insects, other holes are ignored. At each site the anımal will remain still with ıts tongue extended down the hole and while the supply of ants continues. As soon as there are no more insects the pangolın will scratch at another site and the process repeated. In no case was a nest of ants or termites totally devastated and the raided colonies would quickly be able to rebuild the nests and numbers of the varıous castes consumed. Throughout this sniffling at the exposed holes of the ants nest the nose ıs continuously wet and at times the anımal is seen blowing bubbles. The Steppe pangolin in the Transvaal feeds mostly on ants and termites expecıally the former. Table 2 lists the species of ants and termites fed on. SWEENEY (1973) also records that this pangolin species ın the Sudan and in Malawı fed mainly on ants but would only eat the larvae and eggs of Camponotus maculatus and Crematogaster sp. and not the workers. He also recorded that they would not feed on Table 2. Prey species of a free ranging Steppe pangolin in the Transvaal FORMICIDAE Acantholepis capensis Monomorium albopilosum Anoplolepis custodiens Myrmicaria natalensis Camponotus eugeniae Pheidole megacephala Camponotus thales Polyrhachis schistacea Camponotus sp. Tapenonia luteum Crematogaster amita Technomyrmex albipes Xiphomyrmex weitzaeckeri ISOPTERA Odontotermes badius Termitidae - humus feeders Trinervitermes rhodesiensis (possibly Alyscotermes sp. or Astalotermes sp.) Contribution of the ecology of the Steppe pangolin ın the Transvaal 99 Trinervitermes geminatus. In Malawi they were observed to eat worker driver ants, those of several Camponotus species and Trinervitermes sp. nymphs. The pangolıns were reluctant to feed on Odontotermes smeathmani when the nest was unopened, but would eat these termites when the nest was artıfıcially broken open. SwEENEY (1973) also recorded that the pangolins never broke open nests of Trinervitermes and Macrotermes but ate the termites when these were found. The pangolin foraged for termites under cow pats and wood sometimes picking these up and holding them to the chest. Although this last observation has not been observed in the Transvaal, ıt has also been found that pangolins do not break open Trinervitermes or Macrotermes nests but appear to concentrate on litter feeding termites and on those termites found away from the nest. They do, however, concentrate far more on ants of the family Formicidae as the accompanyıng table shows. A large male Steppe pangolın defaecated, after being ın captivity for approxımately 60h, 160 8 of faecal material, largely soil but also some vegetation, sand, remnants of formicid ants and termites. There were even several stones measuring up to 9x 3 mm. These are no doubt accıdental ingestions. Another day later this anımal voıded a further 107.5 g of faecal material much the same as the last. Therefore this anımal had ingested at least 267.5 g of food and secondary material prior to capture as no food had been consumed since. Reproduction The Steppe pangolin gives birth to a single neonate (Van EE 1966; SMITHERS 1983; MOHR 1961; Courson 1989). This was substantiated during the course of this study with one exception, that of a female, observed by H. PETTIFER (pers. comm.) on the farm Tangala ın the Transvaal lowveld, accompanied by two young of sımilar sizes which indicated that they were from the same litter. This would be the first recorded birth of twins ın the species. During the course of this study an injured female gave birth to what appears to be a premature neonate, ın August. This neonate, born at the Onderstepoort Veterinary School, measured 22.5 cm ın total length and had a mass of only 120 g. During thıs study a 2.4 kg juvenile was seen to ride on the back of her 8 kg mother. The two juveniles observed by H. PETTIFER above, had masses of 392 g and 510 g respectively while their mother had a mass of 4.74 kg. Mortality During the period of study, four pangolins were found dead or died subsequently from wounds inflicted by humans. Three of these were killed for consumption while the fourth was found shot with a shotgun for no apparent reason. The only natural mortality was that of an adult male from the Farm Schietfontein 55 MS in the Messina distriet which was found by a farmer on the 7th August, 1983. Two days later ıt died and on investigation was found covered with mites (Manitherionyssus heterotarsus) and tampans (Ornithodorus moubata). According to Dr. BOOMkER (Para- sıtologist, Onderstepoort, pers comm.) these eyeless tampans were probably responsible for the pangolin’s death by inducing progressive paralysis. Management On account of the number of pangolins brought in, some of which were relocated, it was desirable to determine whether such relocations work. Monitirung using transmitters was only carried out with two individuals one of which subsequently proved to have been injured by a knife or spear in the upper region of the hind limbs. In both cases the animals 100 N. H.G. Jacobsen, R. E. Newbery, M. ]J. de Wet, P. C. Viljoen and E. Pietersen were found dead 10 days after release. Mention has already been made of the female from Vryburg, while the injured anımal came from the Thabazımbı District and was released at the Hans Strydom Dam Nature Reserve on top of the Waterberg. The female from Vryburg was found dead with the head and soft parts of the body consumed. Whether the death was a result of predation or stress could not be deduced as rains had obliterated any tracks. From the behaviour of this anımal it is likely that relocations or translocations may be unsuccessful. Until more knowledge of this species’ habits is acquired it seems wise at this stage to only attempt to relocate confiscated anımals in those vicinities where they were initially found and not try to translocate them to nature reserves where their death could result. Acknowledgements The authors thank all those who participated especially J. GROvE who located many of the specimens. Many colleagues contributed observations and these are gratefully acknowledged. Drs. W. CoATEN and G. PrınsLoo of the Department of Agriculture are thanked for the termite and ant identifications. The co-operation of the landowners especially Mr. J. WHEATER and Dr. E. Young are acknowledged with thanks. The Director of the Transvaal Nature Conservation Division ıs thanked for permission to publish this paper. Zusammenfassung Ein Beitrag zur Ökologie des Steppenschuppentieres Manis temminckü in Transvaal Angaben über das Steppenschuppentier (Manis temmincku) ın Transvaal werden zusammengetragen: Verbreitung außerhalb des Krüger-Nationalparks, Körperlängen und Gewichte, Aktivität und Ernäh- rung. Die Bewegungen eines Weibchens wurden längere Zeit telemetrisch kontrolliert. References BIGALKE, R. (1932): Beobachtungen an Smutsia temmincki (Smuts) in der Gefangenschaft. Zool. Garten 5, 173-178. Couzson, I. (1989): The pangolın (Manis temmincki Smuts, 1835) in Zimbabwe. Afr. ]. Ecol. 27, 149-155. Dossrt, J.; DANDELOT, P. (1970): A field guide to the larger Mammals of Africa. London: Collins. EE, E. A. van (1966): A note on breeding the Cape Pangolın Manis temminckü at Bloemfontein Zoo. Int. Zoo Yearbook 6, 163-164. Kıncoon, ]J. (1971): East African Mammals. Vol. 1. London: Academic Press. MOoRHR, E. (1961): Schuppentiere, Neue Brehm Bücherei. Wittenberg: Ziemsen Verlag. Raum, U. (1955): Beobachtungen an den Schuppentieren Manis tricuspis und Manıs longıcandata der Elfenbeinküste. Rev. Suisse Zool. 62, 361-367. RAUTENBACH, 1. L. (1982): Mammals of the Transvaal. Ecoplan Monograph 1. Pretoria: Ecoplan. SMITHERS, R. H.N. (1971): The Mammals of Botswana. National Museums of Rhodesia. Museum Memoir No.4. Salisbury: The Trustees of the National Museums of Rhodesıa. — (1983): The Mammals of the Southern African Subregion. Pretoria: University of Pretoria. STUART, C. T. (1979): The distribution and status of Manıs temmincki Smuts, 1983 (Pholidota: Manidae). Z. Säugetierkunde 28, 123-129. SWEENEY, R. C. H. (1973): Background of Baobabs. London: Constable and Co. Authors’ addresses: N. H. G. JacoBsen, R. E. NEwBERY and M. ]. DE WET, Directorate: Nature and Environmental Conservation, private bag 209, Pretoria 0001, Rep. South Africa; P. C. VıLJoeEn and E. PiETERSEn, National Parks Board, private bag X402, Skukuza 1350, Rep. South Africa Z. Säugetierkunde 56 (1991) 101-111 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Seasonal changes in reproduction and diet of the Bushveld gerbil, Tatera leucogaster (Muridae: Rodentia), in Zimbabwe By B. R. Near Department of Biology, University of Saskatchewan, Saskatoon, Canada Receipt of Ms. 18.6. 1990 Acceptance of Ms. 14.8. 1990 Abstract The reproductive ecology and diet of Tatera lencogaster were studied in two areas, mixed woodland and Mıombo, in the Sengwa Wildlife Research Area, Zimbabwe. Both populations ceased breeding during the cool dry season but one population started breeding three months before the other. The two populations had similar diets, and rainfall and temperature were also similar in the two areas. The difference in duration of breeding in the two areas was probably related to food abundance. In both study areas the testes mass of adult males made up about 6 % of their body mass. The adult sex ratio was strongly biased in favour of females during the breeding season. In the mixed woodland area, females had an average litter size of 5.6 # 0.29 and an annual reproductive capacity of 28 young per adult female. In Miombo, females had an average litter size of 4.8 # 0.36 and an annual reproductive capacity of 12 young per adult female. The diet mainly consisted of seeds (50-75 %) and insects (10-40 %), with a small amount of stem and leaf material (< 10 %). There was little seasonal varıation in diet. Introduction The bushveld gerbil, Tatera lencogaster, ıs widely distributed throughout southern Africa (SMITHERS 1983) and has a varıable pattern of breeding. It breeds throughout the year ın Botswana (SMITHERS 1971) but mainly breeds during periods of high rainfall in Zambıa (CHIDumAYo 1980) and the Transvaal (PERRIN and SWANEPOEL 1987). The main objective of the present study was to document the breeding biology and diet of T. leucogaster in two sımilar habitats separated from one another by a distance of 8-9 km. A comparison of the results from the two areas gives one an opportunity to separate the effects of clımate (rainfall and temperature), which is similar ın the two areas, from the more local effects of the abundance and availabilıty of food. This approach is particularly useful if, as was the case ın this study, the breeding seasons in the two areas are very different. A second objective was to examine certain aspects of the reproductive biology of the males. Adult male T7. leucogaster have extremely large testes during the breeding season (PERRIN and SwANEPOEL 1987), similar to certain other species of Tatera which have the largest proportion of body mass allocated to testicular tissue known for mammals (KEnaGy and TROMBULAK 1986). I examine the constancy of this feature within the genus and show how the sıze of the testes is correlated with adult sex ratios and regression of the testes during the non-breeding season. Material and methods Study areas A total of 300 T. leucogaster were collected from a 4 km? area of the Samapakwa plateau (18°05’ S; 28°08’ E) during the period July 1987 to June 1988, and during the period November 1987 to June 1988 a further 145 anımals were collected from a 2 km? area of mixed woodland adjacent to the four square mile area (18°10’ S; 28°10’ E) in the Sengwa Wildlife Research Area in Zimbabwe. U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0101 $ 02.50/0 102 B. R. Neal The Sengwa Wildlife Research Area is described by Cumming (1975). The study area on the Samapakwa plateau is Miombo (a Brachystegia-Julbernardia mixed woodland). The commonest trees were Brachystegia boehmü, B. spiciformis, Julbernardia globiflora, Terminalia sericea, Combretum zeyheri, and Baphia massaiensis. The shrub Xeromphis obovata was particularly common. Most of the larger trees had been destroyed by elephant a decade or so prior to my study but an elephant reduction program, which ended in 1982, has allowed a prolific regrowth of trees. To prevent a late dry season fire from destroying this regeneration of trees the area has been subjected to cooler annual burns at the beginning of the dry season. Consequently, herbs and grasses were sparse. Common grasses included Eragrostis viscosa, E. curvula, E. cilianensis, Arıstida adscensionis, A. pilgeri, A. vestida, Dactyloc- tenıum gigantum, Heteropogon contortus, Leptocarydion vulpiastrum, Panicum maximum, Pogonarthria squarrosa, Schizachyrium jeffreysü, Schmidtia pappoporoides, and Setaria pallidifusca. The second study area was mixed woodland and bushland dominated by Colophospermum mopane, Combretum apiculatum, C. zeyheri, and Erythroxylum zambesiacum. The area was not burnt and so herbs and grasses were much more abundant than on Samapakwa. Common grasses included Eragrostis viscosa, E. cılianensis, E. regidior, E. superba, Dactyloctenium gigantum, Chloris virgata, Urochloa trichopus, Digıtaria eriantha, Andropogon gayanus, and Schmidtia pappoporoides. Rainfall and temperature were monitored at the Sengwa Wildlife Research Institute (18° 10’ S; 28° 13’ E). Monthly means were determined from daily records. Collection and processing of animals The two areas were sampled monthly by lines of snap traps placed at irregular intervals adjacent to burrows. Traps were left in place for two days and were then moved. Trapped areas were not retrapped. Anımals were measured, sexed and weighed soon after removal from the traps. A clean mass was determined by removing the gut, from the lower oesophagus to the rectum, and also the uterus and embryos of pregnant females. Seasonal changes in female reproductive activity were assessed as follows. First, females were classed as mature if they were reproductively active (see below) or had been in the past, i.e. had placental scars, otherwise they were classed as immature. Mature females were further categorized as 1. pregnant if they had implanted embryos; 2. reproductively active if corpora lutea were detected in the serially sectioned ovarıes or if the uterus was distended with fluid (i.e. in proestrus); 3. remaining females were considered to be reproductively inactive. In addition, females were classed as lactating if milk could be expressed from their nipples. Note that ıt was possible for a lactating female to be categorized as reproductively inactive. Seasonal changes in male reproductive activity was assessed in two ways. First the reproductive organs were dissected out and a sperm smear taken from each cauda epididymis. The abundance of spermatozoa for each smear was then quantified as 0 ıf none were present, 1 ıf only a few spermatozoa were present, 2 ıt spermatozoa were common but still scattered on the slide, and 3 ıf the smear mainly consisted of spermatozoa. An average sperm abundance could be determined for any group of anımals. Secondly, the paired testes and seminal vesicles were fixed in Bouin’s solution and weighed to the nearest 10 mg. To determine ıf the reproductive organs of adult males regressed during the non- breeding period the means of the log transtormed mass of the testes and seminal vesicles were computed for monthly samples of males in age classes 4-8 (see below). This procedure largely eliminated changes resulting from a seasonal influx of juveniles into the population. The number of litters and reproductive capacity (production of young per adult female) during the breeding season were calculated using the method described by NEAL (1981, 1982). This method considers that the interval between litters is made up of a non-pregnant period (p,), a pre-ımplantation period (pı), and a post-implantation period (p,) when embryos are macroscopically visible. The value of pı was assumed to equal 4d (see Near 1981) and that of p, was estimated as 24 d because the gestation period, which equals pı + p,, is 28 d (CHipumayo 1980). The average value of p, could then be estimated by assuming that the ratio of p, + pı to p, was the same as the ratio of non-visibly- pregnant adults to visibly-pregnant adults. The average interval between litters could then be estimated. The average number of litters and production of young could then be determined knowing the duration of the breeding season and the mean litter size. The relative age of individuals was estimated using the eruption and subsequent wear of the upper molars as criteria. Eight age classes were recognised, ranging from the condition when the third upper molar was not fully erupted (age-class 1) to the condition when all molars were worn to the state where no enamel was visible on the upper surface (age-class 8). Stomachs were preserved in Bouin’s solution before transfer to 70% alcohol. The stomach contents of animals collected in the same month and area were combined and the composite samples were sent to the Composition Analysis Laboratory at Colorado State University for analysıs by microhistological techniques. Details of the analysıs are described by Near (1984a, b). Food fragments were categorized into four classes: anthropod (mainly insect), seed, plant stem and leaf, and fungi or lichen. Seasonal changes in reproduction and diet of Tatera leucogaster 103 Results Climate There were 909 mm of raın during my year of study compared to a yearly average of 670 mm during the previous 22 years. Prior to my study ıt had last rained on March 26, 1987 and until early February 1988 the monthly rainfall was below the long term average (Fig. 1) except for December 1987. Rainfall was considerably higher than the long term average during the period February-March 1988. Thus, the April-November dry season was longer and more severe than usual, and rainfall was heavier and more concentrated towards the end of the rainy season than usual. The temperature also varıcd scasonally (Fig. 1), being at amaxımum in November at the end of the dry season and beginning of the rains (mean monthly maxımum of 37.0°C; mean monthly minimum of 27.7°C) and a minimum in July (mean monthly maxımum of 27.0°C; mean monthly minimum of 8.3°C). Thus, the year may be divided into three main seasons: a cold dry season from mid-May to mid-August when the daily maxima and minima averaged 27-29°C and 8-11°C respectively; a hot dry season from September to November when the daily maxima and minima averaged 34-37°C and 17-22°C respectively; and a warm wet season from December to Aprıl when the daily maxıma and mınıma averaged 31-34°C and 18-21 °C respectively. The precise timing of these seasons varies slıghtly from year to year. 300 200 100 Rainfall (mm) ERS TEON BDA EM AM] Month Fıg.1. A: Seasonal changes in mean monthly minimum and maximum temperatures (°C). B: Histogram of total monthly rainfall (mm) from July 1987 to June 1988. The average total monthly rainfall during the period 1965-1987 is also indicated 104 B. R. Neal Body mass The mean birth mass of 7 young born in the laboratory was 2.8 g (range 2.6-3.0 g) and they were weaned at a mass of 16-22 g at an age of 18-28 d (Near 1990). Individuals of both sexes usually attained sexual maturity at a body mass of 60-70. g. The mean body mass of 65 adult males was 79 g (range 58-113 g) and that of 191 adult females was 72 g (range 44-122 g). This slight sexual dimorphism of body mass is consistent with that reported in other studies of this species and for other species of Tatera (NEAL 1982). Adult body mass varied seasonally, reaching a maximum during the rains and early part of the dry season and declining to a minimum towards the end of the dry season. For example, parous, non-pregnant females in the Miombo habitat had a mean body mass of 61 g during September-November compared to a mass of 74 g during January-June and the body mass of adult males averaged 72 g and 83 g for the same two periods. Seasonal cycles in body mass have also been reported for T. nigricauda (NEAL 1982) and T. afra (ALanson 1958; MEASRocH 1954). Sex ratios The sex ratio of juveniles (1.e. individuals in the youngest two age classes) was not significantly different from parity both in mixed woodland (y’ = 1.1; P> 0.2) and Miombo (x? = 0.4; P > 0.5). Similarly, the sex ratio of adults (i.e. individuals in age classes 3-8) was not significantly different from parity during the non-breeding period, May- -November, in Miombo (x? = 0.75; P > 0.3). In contrast, the sex ratio of adults during the breeding season was strongly biased in favour of females both in mixed woodland (x? = 100 u... MIXED WOODLAND o 10 9 19 22 16 1238 MIOMBO e- 50 Female Reproductive Activity (%) I MAATS 70. ANErDEe] Month Fig.2. Seasonal changes in adult female reproductive activity of T. leucogaster in two habıtats. Percentage of adult females: 1. with corpora lutea or in proestrus (@----@), 2. with implanted embryos (A— A), and 3. lactating (histogram). Numbers represent sample sızes Seasonal changes in reproduction and diet of Tatera leucogaster 105 12.2; P< 0.001) and Miombo (x? = 5.7; P < 0.02). CHıpumayvo (1980) has also reported 1:1 sex ratios for juveniles and subadults, but a significant excess of old females during the breeding season in 7. lencogaster ın Zambia. Breeding season Female breeding activity was seasonal (Fig. 2) and mainly confined to the period of rainfall. The breeding season was longer ın the mixed woodland area than in Miombo. Pregnancies were first observed in Miombo during the latter half of December resulting from conceptions sometime after the first week of December. In contrast, when the area of mixed woodland was trapped for the first time during the latter halt of November most females were pregnant and the presence of lactating females (Fig.2) and young juveniles (Fig.4) indicate that breeding had been ın progress for at least two months. Thus, pregnancies occurred at least three months earlier in mixed woodland than in Miombo. Breeding in both study areas ceased coincidentally with the end of the rains at the end of April. Adult testes mass and seminal vesicles mass were at a maxımum during the rains and MIXED WOODLAND 10 3 3 3 2 1.04 : — ee — te (3:0) (30) (3:0) rn 8 N un un E un od .Q un (6) > = ‚or MIOMBO = A ER an 9 1 ) 5 ® 5 o—— nt © (3:0) D——O 5 (2:7) 30) O0) ”® "I (2:7) S ___ _b (2) 1 (1:6) B a 5 (0:8) 2-20 _ 2, 7 (11) 2 a = nr < un 0-5 R /\ © 4 _— Sa % 9 k 1 -& - (0-1) ji x ’ / AR SE ON Di ers Mine Ars Misi oJ Month Fıg. 3. Seasonal changes in testes mass (e—e®) and seminal vesicles mass (4----A) of 7. leucogaster ın two habitats. Data points represent means and vertical lines # 2 x S.E. Numbers in parentheses are the average sperm rating of the sample, and numbers above each figure represent sample sizes 106 B. R. Neal regressed during the dry season (Fig. 3), but this regression was subject to considerable individual varıation and individuals with large testes and fully mature sperm ratings were present throughout the year. Litter size The mean number of live embryos of 25 pregnancıes ın mixed woodland was 5.6 + 0.29 (range 3-8) and of 10 pregnancies in Miombo was 4.8 + 0.36 (range 3-6). In both areas, litter size increased as the breeding season progressed, from a mean of 4.2 during November and December to 6.5 during March and April in mixed woodland, and from 3 in December to 5 during March and April in Miombo. Both the difference between areas and the difference between months were significant when analyzed by two-way analysis of variance. Litter size was not significantly correlated with body mass or age. Breeding rates and reproductive capacity The potential annual production of young per adult female was calculated independently for each study area assuming a breeding period from September to Aprıl in mixed woodland and from December to April in Miombo. In mixed woodlands females could theoretically have had an average of five litters during the breeding season resulting in a production of 28 offspring, whereas females in Miombo could only have had an average of 2.5 litters resulting in 12 offspring during the breeding season. Counts of placental scars indicated that no female had more than three litters and that it was unusual for females to have more than two litters. Thus, the high reproductive capacity in mixed woodland must have been achieved by a high turnover rate of adult females. Population structure The age structures indicated marked seasonal recruitment of young into the populations (Fig.4). This was particularly evident in Miombo where most young were produced by females that had survived from the previous breeding season, although a few young matured rapidly and began breeding in April at the end of the breeding season. By the beginning of the dry season ın May and June, most of the old anımals had died off and been replaced by their offspring. In mixed woodland, the replacement of old anımals by their otfspring occurred in much the same way, but young began breeding by January and formed the majerity of the breeding population by March. Thus, the Miombo population generally consisted of older individuals than the mixed woodland population except during the last three months of the study, and less of the Miombo young bred during the same breeding season of their birth compared to the mixed woodland population. Presumably these differences are mainly related to the length of the breeding season in the two areas. Similarly, CHhrpumayo (1980) showed that the main recruitment of young occurred at the end of the rains and beginning of the dry season (March-June) ın Zambia. These juveniles advanced in age and formed the majority of the population by the start of the following breeding season, after which they became rare or disappeared from the popula- tion. Diet The diets in the two study areas were superficially similar (Fig. 5). Seeds formed the bulk of the diet (50-75 %) throughout the year and insects were also commonly eaten (10-40 %). Only a small amount of stem and leaf material was consumed (< 10 %), and thıs was at a maxımum during the rains and early part of the dry season. Towards the end of the rains and early part of the dry season (March-June) a small amount of fungi and lichens were also consumed. However, there was surprisingly little seasonal varıatıon in diet (Fig. 5). Seasonal changes in reproduction and diet of Tatera lencogaster 107 MIXED WOODLAND 12 5 24 12 29 18 32 D m > 8 - 5 j I 6 5 i ® 3 Eh En S DO ® MIOMBO < 17 25 44 43 31 24 22 10 20 34 30 g ] ER i 7 & 5 4 3 | 1 Der SZ 03 NED ) EM A, M-) Month Fıg.4. Monthly age structure of T. leucogaster in two habitats. Open blocks represent immature anımals and solid blocks represent mature anımals. Sample sızes are shown above each figure In contrast, PERRIN and SwANEPOEL (1987) observed a marked seasonal varıation in diet in the Transvaal: insects and seeds predominated during the rains and herbage was eaten ın large amounts during the dry season. Overall, the diet consisted of 41.4 % insects, 32.2 % herbage, and 26.4 % seeds. Discussion Testes size and sex ratio Information on adult male body mass, testes mass and adult sex ratios during the breeding season are summarized for different species of Tatera ın the Table. There is a wide range of testes size in the genus, and the relative size of the testes of 7. leucogaster ıs second only to T. afra which has the largest proportion of body mass allocated to testicular tissue that is known for any mammal (Kenacy and TROMBULAR 1986). Testes size in mammals ıs considered to be related to copulatory frequency and consequently to the type of mating system (HARCOURT et al. 1981; Kenacy and TROM- BULAK 1986). There are two broad categories of mammalıan mating systems: 1. Single-male systems, where a female mates with a single male, copulation is infrequent and the testes are small, and 2. multi-male systems, where a female mates with more than one male, copulatory frequency ıs high and the testes are large. Little is known about the mating systems of most myomorph rodents although many are thought to be promiscuous, i.e. multi-male breeders (KEnaGy and TROMBULAK 1986). Based on their study of the relationship between relative testes size and mating system, we 108 100 MIXED WOODLAND 19 IM FunsvLichen SEEDS 50 [__] sTEM & LEAF Bi ArtHRoroD MIOMBO 20 % Diet Composition JA SW IQ: Ne "Dr IHRE Fr MAT EN Month Fig.5. Seasonal changes in diet composition of T. lencogaster in two habitats. Numbers represent sample sızes would predict (Table) that 7. nigricauda has a single-male breeding system and the other species have multi-male breeding systems. In this case we would expect 7. nigricanda to be monogamous and have an even sex ratio (as observed) because the other type of single-male mating system, extreme polygyny in which a single male copulates with all females in a group, Is not very likely in a small mammal with a dispersed distribution. The other species of Tatera probably have promiscuous mating systems with either even or skewed sex ratios. In fact there seems to be a relationship between adult sex ratios during the breeding season and relative testes sıze (Table). Species wıth small testes had even sex ratios and species wıth large testes had skewed sex ratios in favour of females. The juvenile sex ratio of all species, except 7. brantsü, were not sıgnificantly different from parity. Most juveniles matured between breeding seasons and so it is interesting to note that adult sex ratios outside of the breeding seasons were also not significantly different from parity. This implies that the change in sex ratio occurred at the time of breeding and so may be related to the type of mating system. The skewed sex ratios are probably the result of differential mortality rather than different capture rates because one would expect promiscuous males to be more active than females. In conclusion, it would seem that Tatera would make an interesting study anımal for mammalıan mating systems because of the wide range ot testes sıze and adult sex ratios in different species of the genus. The testes of T. indica (Prasan 1956), T. leucogaster (PERRIN and SwANEPOEL 1987; this study) and 7. afra (Arnanson 1958) regress during the non-breeding season, whereas those of T. nigricanda, T. robusta and T. valida (NEAL 1982) show little seasonal varıation in size. T. brantsii breeds throughout the year (ALLanson 1958). Thus, the regression of the testes shows a positive correlation with testes size (Table) in seasonally breeding Tatera. Seasonal changes in reproduction and diet of Tatera leucogaster 109 Adult sex ratios and percentage of testes mass in relation to total body mass of adult males during the breeding season for different species of Tatera Relative testes size is the ratio of observed mass to that predicted by the equation: Testes mass (g) = 0.031 x (Body mass g)°”’ for rodents (KenaGy and TROMBULAK 1986). Sample size is for testes and body mass measurements Sample Testes/Body Relative X values Reference size mass testes (n) (%) sıze Tatera nigricanda ! NEaAL 1982 T. robusta : NEAL 1982 T. valıda i NEAL 1982 T. indica E : £ Prasap 1956, 1961 T. brantsii i ö 130/202 Ko ALLANSoN 1958; MEAsRocH 1954 T. lencogaster' 20/49 2 lhrsstudy, T. lencogaster” 22/41 7 This study T. afra 90/142 7 SE INETANSONDL958; MEAsRocH 1954 ! Mixed woodland area. -* Miombo area. * p < 0.05; *** p < 0.001. Perhaps the regression of the testes ıs related to the energy costs of maintaining mature- sızed gonads during the dry season when, presumably, food supplies are more limiting. The gonads and accessory reproductive glands of male 7. lencogaster and T. afra comprise more than 10 percent of their body mass and so considerable savings of energy could be achıeved by reducing the sıze of the reproductive system during the non-breeding season. There was considerable individual varıatıon in the regression of the testes of 7. leucogaster at Sengwa. During the dry season, the testes and seminal vesicles of anımals provided with supplemental food were considerably larger than those of anımals from non-supplemented areas (see Breeding Season below; NEAL and ArısHar 1991), which suggests that the regression of the testes ıs related, at least in part, to energy consumption. Breeding season The population of 7. lencogaster in mixed woodland appears to have started breeding in September 1987 approximately three months ahead of the Miombo population which began breeding in December 1987. This difference was not related to rainfall or tempera- ture because these were similar in the two areas, which were only 8-9 km apart. The diets of anımals from the two areas were also similar and were virtually identical in terms of the proportions of seeds, stem and leaf, and arthropods consumed in November (Fig. 5) when the mixed woodland population was breeding but the Miombo population was sexually quiescent. There is circumstantial evidence, however, that food may be more abundant in the mixed woodland area compared to Miombo, which would allow a greater consumption of food and provide more energy for reproduction. First, a large proportion of Miombo is burnt annually at the start of the dry season whereas the mixed woodland area was never burnt. Such burns would decrease the abundance of seeds, which form a major component of the diet, and also of arthropods by reducing the plant biomass. Second, the density of small mammals in the mixed woodland (258 animals at 862 trap locations) was almost twice as high as that in Miombo (178 animals at 1029 trap locations) although the densities of T. leucogaster were similar in the two areas (123 Tatera at 862 trap locations in mixed woodland and 171 Tatera at 1029 trap locations in Miombo). Third, the larger body mass, litter size and reproductive rate of mixed woodland animals compared to Miombo anımals 110 B. R. Neal is also more consistent with the proposed availability of food ın the two areas rather than other posıble causes such as differences in population density or age structure. On the basis of age structure one would expect Miombo anımals to be heavier on average than those in mixed woodland but the reverse was true; no relationship was observed between litter size and age; and the density of this species was sımilar in the two study areas. Finally, a reproductive response to extra food has been demonstrated for this species, at least for males (NEAL and ArısHa1 1991). A field experiment in August 1987 provided supplemen- tary seeds for a period of 10 d to a population in Miombo habıtat. In seed supplemented areas both sexes were significantly heavier, the testes of the males were approximately twice as heavy, and the seminal vesicles were three to four times as heavy as those of animals from control (1.e. non-supplemented) areas. Thus, the seasonal breeding cycle may be explained as follows. Animals cease breeding in May at the start of the cool dry season. This is probably a response to the rapid decline in minimum temperature (Fig. 1), when the animals are actıve, although a photoperiod response cannot be discounted. The cessation of breeding does not seem to be linked to the availability of food or moisture because both areas were still lush and green from the previous month’s rain, and seeds and insects were plentiful. Breeding starts again at the beginning of the hot dry season ın September if high quality food is sufficiently available, as would seem to be the case in the mixed woodland area. If food is insufficient, breeding is delayed until the onset of the rains when the availability first of insects, and then seeds, increases. This would explain the breeding delay in Miombo. The general pattern of breeding of T. leucogaster in Zambia (CHipumayo 1980) and the northern Transvaal (PERRIN and SwANEPOEL 1987) ıs similar to that described for the mixed woodland area in Sengwa. Food has been suggested to have an important influence on the reproductive cycle of this and other species of the genus (CHipumayo 1980; NEAL 1982; PERRIN and SwANEPOEL 1987), and ROwE-RoWE and MEESTER (1982) have sug- gested that low temperatures can further influence the timing of reproduction of small mammals ın South Africa. Acknowledgements I am indebted to the Department of National Parks and Wild Life in Zimbabwe for permission to work at the Sengwa Wildlife Research Institute and for Drs. D. Cumming and D. Gisson for help in establishing this study. I am particularly grateful for the field assistance of J. NEAL, X. MBULAwA and J. Murunna. I wish to thank I. Courson for the meteorological data from Sengwa, Z. MAHLANGU for identifying the grasses in the two study areas, E. BUECKERT for sectioning ovaries, and T. FoppE, Colorado State University, for organizing the identification of stomach contents. This research was supported by the Natural Sciences and Engineering Research Council of Canada and a sabbatical leave travel grant from the University of Saskatchewan. Zusammenfassung Saisonale Schwankungen in der Fortpflanzung und in der Nahrungszusammensetzung beim Bushveld Gerbil Tatera lencogaster (Muridae: Rodentia) in Zimbabwe Die Fortpflanzungsverhältnisse und die Nahrungszusammensetzung von Tatera leucogaster wurden in zwei Gebieten (in gemischtem Waldland und in Miombo) in der Sengwa Wildlife Research Region untersucht. In beiden Populationen fand während der kühlen Trockenzeit keine Fortpflanzung statt. Eine Population begann aber drei Monate vor der anderen sich fortzupflanzen. Die zwei Populationen zeigten ein ähnliches Nahrungsspektrum, und Regenfall und Temperatur waren in den untersuchten Gebieten ebenfalls ähnlich. Der Unterschied in der Fortpflanzungsdauer beider Populationen hing wahrscheinlich mit dem Nahrungsangebot zusammen. In beiden Gebieten betrug das Gewicht der Testes der adulten Männchen etwa 6 % ihres Gesamt- Körpergewichts. Das Verhältnis von männlichen zu weiblichen adulten Tieren während der Fort- pflanzungsperiode ergab mehr Weibchen. Im gemischten Waldgebiet hatten die Weibchen eine durchschnittliche Wurfgröße von 5,6 + 0,3. Daraus ergibt sich eine durchschnittliche jährliche Fortpflanzungskapazität von 28 Jungen pro adultem Weibchen. In Miombo hatten die Weibchen eine durchschnittliche Wurfgröße von 4,8 # 0,4 Seasonal changes in reproduction and diet of Tatera leucogaster 16 und entsprechend eine geringere durchschnittliche jährliche Fortpflanzungskapazität von 12 Jungen pro adultem Weibchen. Die Nahrung bestand hauptsächlich aus Samen (50-70%) und Insekten (10-40%) und ın geringerem Maße aus Gräsern und Blättern (< 10%). Es waren nur geringe jahreszeitliche Verände- rungen im Nahrungsspektrum festzustellen. References ALLANSoN, M. (1958): Growth and reproduction in the males of two species of gerbil, Tatera brantsiü (A. Smith) and Tatera afra (Gray). Proc. zool. Soc., Lond. 130, 373-396. CHIDUMAYO, E. N. (1980): Population ecology of Tatera lencogaster (Rodentia) ın southern Zambia. J. Zool., Lond. 190, 325-335. Cumming, D. H. M. (1975): A field study of the ecology and behaviour of warthog. Mus. mem. Natl. Mus. Monum. Rhod. 7, 1-179. HARcoUuRT, A. H.; Harvey, P. H.; Larson, $. G.; SHORT, R. V. (1981): Testis weight, body weight and breeding system ın primates. Nature 293, 55-57. Kenagry, G. ]J.; TROMBULAR, $. C. (1986): Size and function of mammalıan testes in relation to body sıze. J. Mammalogy 67, 1-22. MEASsRocH, V. (1954): Growth and reproduction in the females of two species of gerbil, Tatera brantsüi (A. Smith) and Tatera afra (Gray). Proc. zool. Soc., Lond. 124, 631-658. Near, B. R. (1981): Reproductive biology of the unstriped grass rat, Arvicanthis, in East Africa. Z. Säugetierkunde 46, 174-189. — (1982): Reproductive biology of three species of gerbils (Genus Tatera) ın East Africa. Z. Säugetierkunde 47, 287-296. — (1984a): Relationship between feeding habits, climate and reproduction of small mammals in Meru National Park, Kenya. Afr. J. Ecol. 22, 196-205. — (1984b): Seasonal feeding habits of small mammals in Kenya. Z. Säugetierkunde 46, 226-234. — (1990): Observations on the early post-natal growth and development of Tatera lencogaster, Aethomys chrysophilus and A. namaquensis from Zimbabwe, with a review of the pre- and post- natal growth and development of African Muroid rodents. Mammalıa 54, 245-270. Near, B. R.; ArısHar, $. K. (1991): Reproductive response of Tatera lencogaster (Rodentia) to supplemental food and 6-methoxybenzoxazolinone ın Zimbabwe. J. Zool., Lond. (in press). PERRIN, M. R.; SwANEPOEL, P. (1987): Breeding biology of the bushveld gerbil Tatera lencogaster in relation to diet, rainfall and life history theory. S. Afr. J. Zool. 22, 218-227. PrAasap, M. R. N. (1956): Reproductive cycle of the male Indian gerbille Tatera indica cuvieru (Waterhouse). Acta Zool. 37, 87-122. — (1961): Reproduction in the female Indian gerbille Tatera indica cuvieru (Waterhouse). Acta Zool. 42, 245-256. RowE-RoOWE, D. T.; MEESTER, J. (1982): Population dynamics of small mammals in the Drakensberg of Natal, South Africa. Z. Säugetierkunde 47, 347-356. SMITHERS, R. H.N. (1971): The mammals of Botswana. Mus. mem. Natl. Mus. Monum. Rhod. 4, 1-340. — (1983): The mammals of the Southern African Subregion. Pretoria: University of Pretoria Press. Author’s address: Prof. Dr. B. R. Near, Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan, Canada S7N OWO Z. Säugetierkunde 56 (1991) 112-118 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Habitat relations in Ctenomys talarum (Caviomorpha, Octodontidae) in a natural grassland By VIvIana M. COMPARATORE, N. ©. MACEIRA and CRISTINA BuscH Universidad Nacional de Mar del Plata, Facultad de Ciencias Exactas y Naturales, Mar del Plata, and Facultad de Ciencias Agrarias, Balcarce, Pcia. de Buenos Aires, Argentina Receipt of Ms. 19. 2. 1990 Acceptance of Ms. 12. 9. 1990 Abstract Tested the hypothesis that the spatial distribution of Ctenomys talarım (“tuco-tuco”) in a natural grassland on a coastal clıff at Mar de Cobo, Argentina, is determined by topographic, soil and vegetation characteristics of the habıtat. In summer, there was no correlation between the number of “tuco-tucos” and any of the vegetation, soil or topography variables. In contrast, in autumn and winter, the number of animals was positively correlated with the density of grasses with small reserve organs (definite rhizomes). In spring, the number of animals was positively correlated with soil humidity, density and height of the vegetation, and density of grasses without reserve organs and with small reserve organs, and was negatively correlated with the sand percentage in the soil. "The results indicate that the spatial distribution of Ctenomys talarum is associated with the studied variables, although their relative importance varıes seasonally. Introduction Rodents of the genus Ctenomys (“tuco-tucos”) are subterranean and herbivorous. Popula- tions are distributed throughout Argentina, southern Brazil, and parts ot Paraguay, Bolivia, Uruguay, Perü and Chile (Woops 1984). The “tuco-tucos” dig sinuous burrows with several openings that are plugged with vegetation and/or sand during the greatest part of the day and night. These burrows consist of a main gallery connected with shorter ones that end on or below the surface (CoNTRERAS 1973; ALTUNA 1983). Because movement underneath the earth is energetically very expensive, characteristics related to energetic balance and foraging costs are presumably under high selection pressure (VLEcK 1981). Since the habitat productivity determines the foraging benefits and the soil hardness determines the costs, foraging economies should favor a maximum body sıze for each type ot habıtat such that benefits and costs are equal (VLEcK 1981). The larger anımals should occupy zones with high productivity and/or lighter soils. The subadults should occupy less favorable habitats, migrating to favorable zones when these are vacated because of the death of their previous occupants (PEARson et al. 1968). Other habitats should be unsuitable because of excessive hardnesss ot the soil or low food availabiılity. Soil texture ıs also important in determining burrow temperature and ventilation, with sandy soils favoring gaseous exchange (McNag 1966; WırLson and KıLGorE 1978). Consequently, factors such as topography and exposure of the occupied zones relative to dominant winds, and vegetation structure, should also affect Ctenomys habitat selection. Because of the high energetic cost of digging, soil hardness could be the most limiting factor affecting the dispersion of subterranean rodents (Best 1973; ABRAHAM 1980). Species with larger body sıze would be the most affected (Best 1973; VLEcK 1981). Along the Atlantic coast of Buenos Aires Province, Argentina, there is a clear halt of Ctenomys distribution between Santa Clara del Mar and Necochea (CoNTRERAS and REıG 1965). U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0112 $ 02.50/0 Habitat relations in Ctenomys talarum in a natural grassland 113 CONTRERAS (1973) found a positive correlation between the textural type “sandy-clay- loam” of the second horizon and the presence of Ctenomys. Tuco-tucos at Mar de Cobo (Partido de Mar Chiquita), where the habıtat ıs hetero- geneous, have a clumped distribution (Busch# et al. 1989). The present study tests whether Ctenomys spatial distribution is associated with topography, soil and vegetation charac- teristics of the habitat; and evaluates their relative importance. Study site The experimental area covered a surface of 100 by 70 m divided ın 10 by 10 m squares in Mar de Cobo (Partido Mar Chiquita, Buenos Aires Province, Argentina) on a coastal cliff where the vegetation was a natural grassland with predominance of species wıth rhizomes. To characterize the vegetation of the site, the frequency of occurrence of each species was calculated season- nentlly mean precipltation (mm) „]]y as the number of touches of green parts over the number of total temperature(‘C) 220 touches. The dominant species ın summer were Panıcum racemosum (32%), Ambrosia _tenuifolia 180 (17%), and Distichlis scoparia mm (16%). In autumn, Panıcum racemosum (19%), Distichlis 140 scoparia (12 %), Bromus unioloides (8 %) and Stipa neesiana (7 %). In winter, Bromus unioloides (17 %), 100 and Stipa neesiana (8 %); ın this season, 61 % of the vegetation was dry, and warm season species such 30 60 as Panicum racemosum and Distich- lıs scoparia dropped to 2% and r „2 a 4 % of green material, respectively. 20 In spring, there were 19% Panıcum racemosum, 19 % Bromus u | unioloides, 10 % Ambrosia JASONDJ FMAM J tenuifolia, 10 % Distichlis scoparia, and 8 % Stipa neesiana. Fig. 1. Cilmatic data (1986). Source: Estacıiön A&rea Mar del In sixteen measurements, the Plata. Data are plotted according to WALTER (1977) sand percentage in the soıl varıed from 88 % to 99 %, winter soil moisture varıed from 4 % to 19 %, spring soil moisture varied from 4 % to 15 %, and soil hardness varied from 7 to 52 kg/cm?. The topographic map showed that the difference in altitude between the lowest and the highest point of the study area was 2.4 m. Fig. 1 shows climatic data for the year when the studies were conducted (1986). Data are plotted according to WALTER (1977). Material and methods The vegetation of the experimental area was sampled in March, May, August, and December 1986 by means of a non-destructive method which consisted of positioning a vertical needle and recording the canopy contacts (LEvy and MADDEN 1933; GREIG-SMITH 1964; HESLEHURST 1971; Jong et al. 1983). The needle was lowered systematically, each 20 cm, along four 70 m transects perpendicular to the coast. For each touch we recorded: the plant species, its height above the surface, and whether it was living. This method of sampling vegetation underestimates forbs and overestimates grass cover (GREIG-SMITH 1964), and may explain the low percentages of Hydrocotyle bonariensis, which never exceeded 5 % in the plots. Samples of soil were collected at a depth of 20cm in August and December along the same transects used to study vegetation. Four 500 g samples were obtained systematically in each transect. Organic matter from soil samples was removed using hydrogen peroxide. Sand percentages were determined by leaching the silt and clay fractions. These latter percentages were determined only in the winter samples, because they do not vary throughout the year. Soil moisture was determined by 114 Vivıana M. Comparatore, N. O. Maceira and Cristina Busch differential weight readings before and after drying the soil samples at 105 °C. Soil hardness was measured along the transects with a soil penetrometer. The contours of the topographic map were traced with respect to the highest point that was set at the arbitrary value of zero. The greater values are the lower points of the study site. Density and spatial distribution of Ctenomys were obtained in a sımultaneous study in the same grid (BuscH et al. 1989) by capture-recapture methods. Population size on the grid was estimated as the minimum number known alive between trapping intervals (PErrusewicz 1983). The Jolly-Seber method (CAuUGHLEY 1977) was also used to estimate population size, but in the present study we used the former situation (Fig. 2), and data on their spatial distribution to estimate the number of animals in each square of the grıd. Poisson distribution analysıs indicated that the distribution of animals on the study area was clumped (df. = 69; P = 0.01). The following varıables were measured for each square in each season: a. Vegetation: Total cover (total number of touches); Green cover (number of touces of green parts); Mean maximum height (cm); Cover of grasses with large reserve organs (indefinite rhizomes); Cover of grasses with small reserve organs (definite rhizomes); Cover of grasses without reserve organs; Cover of forbs with reserve organs (rhizomes); Cover of forbs without reserve organs (Table). We grouped the vegetation this way because we considered that the presence of rhizomes is an important character in Ctenomys nourishment. b. Soil: Percentage sand; Relative humidity; Soil hardness (kg/cm? ): c. Relief: Distance from the coast (m); Relative elevation (m). d. Ctenomys talarum: Number of individuals. Vegetation, soil and relief data were analyzed complementing ordination and classification techniques. Principal Components Analysıs (PCA) (Harrıs 1975) and Cluster Analysis (CA) by the nearest neighbor with simple linkage (OrLöcı 1978) using 1-Pearson correlation coefficient for sample distances, were performed. Simple correlations were established between sample positions along the first seven axes and sample anımal numbers. SYSTAT statistical program was used. Vegetation groups and their principal species Grasses with large reserve organs Grasses with small reserve organs Grasses without reserve organs Panicum racemosum Festuca arundinacea Bromus unioloides Distichlis scoparia Poa bonariensis Stipa neesiana Paspalum vaginatum Poa lanigera Dactylıs glomerata Stenotaphrum sp. Paspalum dilatatum Phalaris platensis Cynodon dactylon Setaria geniculata Lolium multiflorum Lolium perenne Hordeum leporinum Sporobolus indicus Forbs with reserve organs Forbs without reserve organs Ambrosia tennifolia Medicago lupulina Hoydrocotyle bonariensis Mebhlotus officinalıs Solidago chilensis Phylla cannescens Oxalıs cordobensis Stellaria media Anagallıs arvensis Sonchus oleraceus Geranium dissectum Rapıstrum rugosum Brassica campestris Dichondra repens Gamochaeta spicata Results Fig. 3 shows data ordination, grouped according to CA. The figure ıs a bidimensional representation of the first two spring axes. Although analysıs of all the seasons were performed and plotted, spring results are the most interesting since animal spatial distribu- Habitat relations in Ctenomys talarum in a natural grassland 115 70 60 SIZE 50 40 30 POPULATION 20 10 Farin JOLLY-SEBER MINIMUM NUMBER KNOWN ALIVE J A Fig. 2. Estimates of number of Ctenomys talarım on a 100 by 70 m grid in Mar de Cobo, Argentina S 16) N D J F M A M BJ July 1985-May 1986. Vertical lines indicate 2 SE above and below the Jolly Seber estimate. Numbers indicate proportion of males in the sample. (Taken from BuschH et al. 1989) Fıg. 3. Scatter dıagrams of spring samples along axes 1 and 2 of the Principal Com- ponents Analysıs. Group- ing according to Cluster Analysıs. Values between brackets show the percent- age of the variance ex- plained by each axis. GC = green cover; TC = total cov- er; G = cover of grasses without reserve organs; H = vegetation mean maximum height; D = distance from the coast; Gl = cover of grasses with small reserve organs; HU = humidity percentage ın the soil; $S = sand percentage in the soil; GR = cover of grasses with large reserve organs; FR = cover of forbs with reserve organs; SH = soil hardness DENSE VEGETATION INTERMEDIATE VEGETATION 118,30 %) SH SPARSE VEGETATION PARSE VEGETATION 116 Viviana M. Comparatore, N. ©. Maceira and Cristina Busch tion ıs closely related to vegetation heterogeneity at the community level. In the other seasons, anımal distribution is associated with intracommunity heterogeneity (fine grain), and thus, scatter diagrams are poorly informative. A common aspect of the analyzed seasons is that the principal tendency of data varıability (ax 1 ofthe PCA) shows a contrast between samples with sparse vegetation and sandy soil, and samples with dense and high vegetation and humid soil, more distant from the sea. Such contrast always explains over 38 % of the total varıance. In March (summer) no correlation was found between the number of “tuco-tucos” and the position of the samples along the first seven axes. In this season, no soil samples were collected; as a consequence, there were no moisture data. In May (autumn), the number of anımals was significantly correlated (P < 0.05) with the position of the samples along ax 3 of the PCA, and significantly correlated (P < 0.055) with the position of samples along ax 4. No correlation was found with any of the other first seven axes. Ax 3 opposes forbs without reserve organs and grasses that have small reserve organs, with grasses and forbs that have large reserve organs. The loading of these variables in ax 3 ıs 0.730, 0.474, 0.679 and 0.439 respectively, being anımal distribution positively correlated with the first two varıables. Ax 3 explains 13.45 % of the total varıance. Ax 4 shows a gradient of samples wıth and without grasses that have small reserve organs. The loading of this varıable in ax 4 ıs 0.601, being the number of animals positively correlated with the presence of grasses that have small reserve organs. Ax 4 explains 8.07 % of the total varıance. In this season, no soil samples were collected. In August (winter), the number of “tuco-tucos” was significantly correlated (P < 0.05) with the position of samples along ax 4. No correlation was found with any of the other first seven axes. Ax 4 shows a gradient of samples with and without grasses that have small reserve organs. The loading of this varıable in ax 4 ıs 0.793, being the number of anımals positively correlated with the presence of grasses that have short reserve organs. Ax 4 explains 9.80 % of the total variance. In December (spring), the number of anımals was significantly correlated (P < 0.05) with the position of the samples along axes 1 and 6. No correlation was found with any of the other first seven axes. Ax 1 opposes samples of sandy soils (loading 0.826) and low vegetation cover, with samples of high vegetation cover (total cover, green cover, cover of grasses with small reserve organs and without reserve organs, height of the vegetation, whose loadings lie between 0.768 and 0.949), humid soils (loading 0.675), more distant from the sea (loading 0.794). The number of anımals is positively correlated wıth high vegetation cover and humid soils. Ax 1 explains 45.81 % of the total varıance (Fig. 3). Ax 6, which only explains 3.53 % of the total variance, shows a gradient of samples with and without forbs that have reserve organs (loadıng 0.418), being the number of anımals pösitively correlated with the samples without forbs that have reserve organs. Discussion Soil texture and humidity are important factors in the distribution of subterranean rodents. ABRAHAM (1980) found that Ctenomys talarum does not build its burrows in soils containing 25 % or more of clay; Hansen and BEck (1968) found that when the soil moisture exceeds 50 %, pocket gophers (Thomomys talpoides) move to dryer sites. The percentage of sand in the soil of our experimental area ıs high, never below 87 %, therefore, texture and hardness should not be limiting factors in the dispersion and occupation of new zones of the area. As sand percentage is very high, soil humidity ıs low, even in the colder seasons. Soil humidity and probably soil temperature, conditioned by the vegetation cover, can be very important when climatic factors are extreme. The fact that the number of animals in late spring (Fig. 3) was positively correlated with humidity, Habitat relations in Ctenomys talarum in a natural grassland 117 suggests that low soil humidity and high soil temperature can affect the occupation of certain areas during the hot season. Therefore, in warm months, favorable zones should be those of higher humidity, and consequently less sandy and with greater density and height of the vegetation (Fig. 3). Also, in spring, most of the vegetation is green, so that the positive correlation of the “tuco-tucos” with the density of the vegetation can have a dietary explanation. Results of this kind would be expected ın summer. In this season, on the contrary, no correlation was found between anımal density and any of the varıables examined. During this season, green vegetation (warm season species) is ubiquitous. Also, this period coincides with the end of the breeding season and the dispersion of the subadults who will occupy the less favorable areas (PEARson et al. 1968). In this situation, the animal distribution could be mainly determined by social factors. When the aerial productivity of the vegetation is low in autumn and winter, the anımals’ energetic requirements are higher because of the regulation of body temperature and the lactation of the young. The breeding season lasts nine months, from the end of May to the end of February with a peak ın August (Busc# et al. 1989). The autumn (ax 3 and 4 of the principal components analysıs) and winter (ax 4) association of anımal density with grasses that have small reserve organs, suggests a relationship between energetic requirements and animal spatial distribution. Most of this vegetation is composed of cool season species, and is therefore green when most of the other vegetation ıs dry. Further, Reıc (1970) and LOHFELDT et al. (1989), suggest that Ctenomys eats much more aerıal than underground vegetation. Also, in autumn, the positive correlation of anımals wıth the presence of forbs without reserve organs (ax 3 of the PCA), can be explained by anımal continuous earth movements that favor the implantation of annual forbs. Although the tendency of the varıable along the axes ıs subtle, the negative correlation of anımal spatial distribution with the presence of forbs that have reserve organs in autumn (ax 3 of the PCA) and spring (ax 6), suggests a rejection of these species, which are principally Ambrosia tennifolia and Hydrocotyle bonariensis. This rejection can have a dietary explanation, since LOHFELDT et al. (1989) suggest that Ctenomys talarum prefers grasses in autumn and spring. In conclusion, the clumped distribution of Ctenomys talarum ın the area, can be correlated with environmental factors, the presence of grasses with small reserve organs being the most important determinant ın autumn and in winter, and soil humidity, density and height of the vegetation in spring. Also, data suggest that other factors, like animal behavior, may be important determinants of anımal spatial distribution during certain seasons. Acknowledgements We wish to thank the “Centro de Geologia de Costas, Universidad Nacional de Mar del Plata”, for their assistance in the processing of the soil samples and the realization of the topographic map of the study area. Also to “Grupo Pasturas, Unidad Integrada Facultad de Ciencias Agrarıas, Universidad Nacıonal de Mar del Plata, e EEA Inta Balcarce”, for the use of the computer programs. For many helpful discussions we thank Dr. OsvaLdo A. Reıc and M. Sc. MönıcA AGNUSDEI. CRISTINA BUSCH is as member of CONICET, and VIvIana COMPARATORE holds a fellowship from CONICET. Zusammenfassung Habitat-Beziehungen von Ctenomys talarum (Caviomorpha, Octodontidae) auf natürlichem Grasland Auf einer bestimmten Fläche von natürlichem Grasland an der Küste von Argentinien nahe Mar de Cobo, Provinz Buenos Aires, wurde untersucht, in welcher Weise die räumliche Verteilung von Individuen der Art Ctenomys talarum (Tukotuko) durch Topographie, Bodenbeschaftenheit und Vegetation bestimmt ist. Für den Sommer ergab sich keine Korrelation von Individuenanzahl zu irgendeinem der untersuchten Faktoren. Im Herbst und Winter war demgegenüber die Anzahl der 118 Viviana M. Comparatore, N. O. Maceira and Cristina Busch Tiere positiv korreliert mit der Dichte von Gräsern, die mit kleinen Speicherorganen ausgestattet sind. Auch für den Frühling ergab sich eine positive Korrelation, diesmal jedoch mit der Bodenfeuchtigkeit, mit Dichte und Höhe der Vegetation, sowie mit der Dichte von Gräsern sowohl ohne besondere, als auch mit kleinen Speicherorganen. Andererseits wurde eine negative Korrelation zum prozentualen Sandanteil des Bodens festgestellt. Die Ergebnisse deuten an, daß die räumliche Verteilung von Tukotukos zu den untersuchten Faktoren in Beziehung steht, im Laufe des Jahres jedoch in unterschiedlicher Weise. Literature ABRAHAM, M. A. (1980): Factores edafıcos limitantes de la distribuciön del “tuco-tuco” de Magdalena (Ctenomys talarum). Ecologia Argentina 4, 14. Arruna, C. A. (1983): Sobre la estructura de las construcciones de Ctenomys pearsoni Lessa y Langguth, 1983 (Rodentia, Octodontidae). Resümenes y comunicaciones de las Jornadas de Ciencias Naturales, Uruguay. Best, T. L. (1973): Ecological separation of three genera of pocket gophers (Geomyidae). Ecology 54, 1311-1319. BuscH, C.; MarızIa, A. 1.; ScAaGL1a, ©. A.; Reıc, O. A. (1989): Spatial distribution and attributes of a population of Ctenomys talarum (Rodentia, Octodontidae). J. Mammalogy 70, 204-208. CAUGHLEY, G. (1977): Analysıs of vertebrate populations. New York: John Wiley. CONTRERAS, J. R. (1973): El tuco-tuco y sus relacıones con los problemas del suelo en la Argentina. Idia 29, 14-36. GREIG-SMITH, P. (1964): Quantitative plant ecology. 2nd ed. London: Butterworths. Hansen, R. M.; Beck, R. F. (1968): Habitat of pocket gophers in Cochetopa Creek Drainage, Colorado. Am. Midl. Natural. 79, 103-117. Hareıs, R. ]J. (1975): A primer of multivariate statistics. London: Academic Press. HESLEHURST, M. R. (1971): The point quadrat method of vegetation analysıs: a review. Reading: Univ. Jong, P. DE; AARsEn, L. W.; TURKINGTON, R. (1983): The use of contact sampling in studies of association in vegetation. J. Ecol. 71, 545-559. Levy, E. B.; Mappen, E. A. (1933): The point method of pasture analysıs. New Zealand ]. Agriculture 46, 267-279. LOHFELDT, M.; COMPARATORE, V.; BuscH, C. (1989): Preferencias alımentarıas de Ctenomys talarum. Resümenes de la 14° Reuniön Argentina de Ecologia, abrıl de 1989, San Salvador de Jujuy, Argentina. McNas, B. K. (1966): The metabolism of fossorial rodents: a study of convergence. Ecology 47, 712-733. Orröcı, L. (1978): Multivariate analysis in vegetation research. The Hague: Dr. W. Junk Publ. PEARson, ©. P.; BinszTEin, H.; Boiry, L.; BuscH, C.; DiPAce, M.; GALLOPIN, G.; PENCHASZADEH, P.; PrANTANIDA, M. (1968): Estructura social, distribuciön especial y composiciön por edades de una poblaciön de tuco-tucos (Ctenomys talarum). Invest. Zool. Chilenas 13, 47-80. PETRUSEWICZ, K. (1983): Numbers: the concept and definitions. Acta Theriol. 28, 89-101. Reıc, ©. A. (1970): Ecological notes on the fossorıal octodont rodent Spalacopus cyanus (Molina). ]. Mammalogy 51, 592-601. VLEck, D. (1981): Burrow structure and foraging costs in the fossorial rodent, Thomomys bottae Oecologia 49, 391-396. WALTER, H. (1977): Zonas de vegetaciön y clima. Barcelona: Ed. Omega, $. A. Wırson, K. J.; KıLGore Jr, D. L. (1978): The effects of location and design on the diffusion of respiratory gases in mammal burrows. J. Theor. Biol. 71, 73-101. Woops, Ch. A. (1984): 14. Histricognath rodents. In: Orders and families of recent mammals of the world. Ed. by $. Anperson and ]J. K. Jones. New York: John Wiley. pp. 389-446. Authors’ addresses: Vıvıana M. COMPARATORE and CRISTINA BuscH, Departamento de Biologia, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata, C. C. 1245 (7600) Mar del Plata, Pcia. de Buenos Aires, Argentina; NEsToR ©. MaceIRA, Facultad de Ciencias Agrarias, Universidad Nacional de Mar del Plata, C.C. 276, (7620) Balcarce, Pcia. de Buenos Aires, Argentina Z. Säugetierkunde 56 (1991) 119-120 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 NZISSIINSIEEPASENNSECTTESKURZNI EI EIBUÜNIGEN: Use of an active badger sett by Egyptian mongooses, Herpestes ichneumon, in Southwest Spain By F. PALOMARES Estacion Biolögica de Donana, Sevilla, Spain Receipt of Ms. 23. 4. 1990 Acceptance of Ms. 30. 10. 1990 Badgers (Meles meles) use dens dug by themselves for rest and protection (Kruuk 1978; Near 1986). Badgers’ setts are often used as a refuge by other mammals, including several carnıvores (NEAL 1986). Some of the carnıvores use deserted setts (see NEAL 1986, for a review). Small carnıvores also use active setts, although only casually, whilst foxes (Vulpes vulpes) use them more regularly and even for breeding in. Here I report for the first time deserted badger setts being used by Egyptian mongooses (Herpestes ichneumon) and describe in detail the regular use of an active sett by several radio-tracked mongooses ın Donana National Park, SW Spain, from September 1987 to March 1989. Mongooses in this area are typically dıurnal and usually use burrows at various locations to rest in at night and for resting periods of several hours during the day (DELIBES and BELTRAn 1985; PALOMARES 1986). During the two years of study at least nine mongooses have used badger setts as a resting area and overnight refuge. In some situations the density of the vegetation made it difficult to establish the type of burrow used. Four deserted badger setts which displayed two to five entrance holes were used by seven mongooses (3 males and 4 females). One active sett was used by 4 males and 4 females as follows (between the parentheses the tracking day number is shown): males: HMI, at least 3 times (117); HM2, 2 times (205); HM3, 1 time (24); HM7, 26 times (53); females: HH4, 2 times (38); HH6, 1 time (163); HH10, 30 times (154); HH12, 19 times (103). Three individuals, HH10, HH12 and HM7 were also located in the sett during the daytime, both resting and active. Of the mongooses captured whose home range included the sett, only one was never recorded inside it. The above mentioned badger sett, in the bank of a small stream, ıs dug out of earth and covers an area of approximately 400 m?. Eleven entrances are visible, although there appear to be more between the vegetation below which the sett is found. Of these 11 only 6 showed signs of use in March 1989. From earlıer visits, no important changes in the entrances in active use were observed. Throughout the study the sett was ın daily use by badgers as confirmed by their tracks, and it can be considered the principal sett of a badger clan (Kruuk 1978). Usually the mongooses used 3 entrances for arrıval and departure, whilst the badgers used only 2; one of them being utilised by both species. The resting site of two of the marked mongooses, HH10 and HM7, was located very close to the entrances most frequently used by the badgers. Different behaviour is usually observed with foxes, since they tend to use the parts of the sett the badgers do not (NEAL 1986). WIJNGAARDEN and PEppEL (1964) have speculated that the use of badger setts by foxes is due to the absence of optimal refuge sites. For mongooses, which frequently use the abundant rabbit warrens of Donana, this is not the case. In the study area, badgers and mongooses display different activity patterns. The U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0119 $ 02.50/0 120 F. Palomares mongoose is dıurnal (DELIBEs and BELTRAN 1985; PALOMARES 1986), the badger noctur- nal. Despite this difference, both species overlap underground for four to ten hours a day. The appreciable size difference between the two species in Donana (approximately 7-9 kg for badgers compared to 3 kg for mongooses) makes such great mutual tolerance unexpected, since confrontations are known to be frequent between other carnivores (e.g. Rogers and MEcH 1981). However, badgers, which are very aggressive among themselves (Kruuk 1978), generally show little interspecific aggression (NEAL 1986). Moreover, there appears to be no disadvantage for the mongoose, in spite of the fact that the two species have a certain sımilarıty of diet and therefore possibly compete for resources in the area (MARTIN-FRANQUELO and DELIBES 1985; PALOMARES 1986). I am grateful to Dr. M. DeLiBEs for review and valuable comments on the manuscript, and to N. BUSTAMANTE, for reviewing the English. The research was supported by DGICYT (project PB87-0405). References Derıses, M.; BELTRAN, ]J. F. (1985): Activity, daily movements and home range of a Ichneumon or Egyptian mongoose (Herpestes ichneumon) in southern Spain. J. Zool., London 207, 610-613. Kruuk, H. (1978): Spatial organization and territorial behaviour of the European Badger Meles meles. J. Zool., London 184, 1-19. MARTIN-FRANQUELO, R.; DELIBES, M. (1985): Ecology of the badger in Donana, Mediterranean Spain. IV Intern. Ther. Congress. Edmonton. August 1985. Near, E. (1986): The natural history of badger. London, Sidney: Croom Helm. 1-238. PALOMARES, F. (1986): Ecologia de la gineta y el meloncillo en el Parque Nacıonal de Donana. Tesina de Licenciatura. Fac. de Ciencias. Univ. de Granada. Rotes, L. L.; MecH, L. D. (1981): Interaction of wolves and black bears in Northeastern Minnesota. ]- Mammalogy 62, 434-436. WIJNGAARDEN, A. VAN; PEPPELM J. VAN DE (1964): The badger (Meles meles) ın the Netherlands. Lutra 6, 1-60. Author’s address: Francısco PALOMARES, Estaciön Biolögica de Donana (CSIC), Apdo. 1056, E-41080 Sevilla, Spain Z. Säugetierkunde 56 (1991) 121-123 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Fully formed first lower premolars in a White-tailed deer (Odocoileus virginianus Zimmermann, 1780) By CyYRILLE BARRETTE Departement de Biologie, Universite Laval, Quebec, Canada Receipt of Ms. 7.6. 1990 Acceptance of Ms. 13.9. 1990 The first artiodactyls (e.g. Diacodexis) trom the lower Eocene of North America, Europe and Asia, had a full dentition (I 3/3, C VI, P 4/4, M 3/3), without diastema (CARROLL 1988). The first ruminants (e.g. Archaeomeryx) from the upper Eocene of Asıa had already lost P!, were on the way to losing their upper incisors, and P, was canıniform and located in a short diastema separating P, from the lower canine (VIRET 1961). Then in the lower Miocene, from Blastomeryx onwards, P, had disappeared almost completely being seen in only a few specimens of Blastomeryx itself, then not at all afterwards (MATTHEw 1908; Frick 1937). Ever since that time, ı.e. for the last 20 million years, most artiodactyls (i.e. the bovids and cervids) have had a tooth formula reduced to 1 0/3, C 0/1, P 3/3, M 3/3, with a long diastema between P, and the incısitorm lower canıne. Even though this has been the condition for about 20 million generations, one occasionnally sees one of the long lost tooth being expressed in the phenotype of a present- day artiodactyl. This is the case for P, for instance that has been reported for some nine species of bovids and cervids (Gazella thomasii: ROBINETTE and ARCHER 1971; Sylvicapra grimmia: CHILD and Rıney 1964; Bison bison: FULLER 1954; Capra pyrenaica: VIGAL and MACHORDOM 1987; Odocoileus virginianus: MECH et al. 1970; O. hemionus: ROBINETTE et al. 1977; Rangıfer tarandus: BANFIELD 1961; MiLLeEr and Tessıer 1971; Cervus elaphus canadensis: MORAN and FAIRBANKS 1966; and Capreolus capreolus: BUBENIK and WURT- ZINGER 1967; MEYER 1985). From 15 studies that have reported on supernumerary teeth ın bovids and cervids (including the ones lısted above), I could compute that the overall frequency of occurence ot P} in the two families is less (and sometimes considerably less) than one ın 3000 individuals. The case described here ıs a young adult male ©. virginianus, hıt by a car north of Montreal (45° 45’ N, 74° 45’ W) during the winter 1988. The anımal was healthy, weighing 78.5 kg, and having a total length of 181 cm. Its skull has a condylobasal length of 289 mm and a bizygomatic width of 116 mm. Its upper tooth row measures 74 mm while the lower (excluding P,) ıs 81.1 mm long. This case is particularly interesting not only because of ıts rarıty, but because both its permanent P, are very similar to what those teeth looked like in pre-cervid artiodactyls. In most of the cases reported above the first premolars are small, peg-like, single rooted, and sometimes limited to one side, 1.e. it is not only a vestigial tooth but a rather stunted form of it. On the contrary, the P, of the specimen presented here, are symmetrical in size, shape, and placement, have two fully formed roots (both measure 7.4 mm long on the left P, that came loose after the drying of the jaw), and have the size and shape that one would predict by extrapolating the trend seen from P, to P; in any normal white-tailed deer (Fig.). The length and height of the crown of the P, are 6.7 and 3.7 mm on the right side, and 7.3 and 3.5 mm on the left. Finally, each P, has three cusps in a straight line (Fig.). On both sides P, is separated from P, by 6.5 mm, a placement very similar to the one seen in some modern suids like Sus scrofa, one of the few present artiodactyls that have U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0121 $ 02.50/0 22 Cyrille Barrette Vestigial Pı in an adult male white-tailed deer seen from the left side. The black bar is 10 mm long retained their P,. The diastema separating P, from the lower canine is 78.5 mm long, 96.8 % of the length of the lower tooth row (excluding P,). That is about the same value as in white-tailed deer without P, (mean + S.D. for 5 males and 5 females = 94.3 + 5,3 %). Thus P, has not caused a lengthening of the diastema, it merely occupies a short segment of it. The form of the P, of this white-tailed deer is premolar-like. They are not the canıniform P| seen in living camelids or suids, in Archeomeryx, or in the Oligocene Hoypertragnlus (CARROLL 1988; VIRET 1961). A specimen such as the one described here lends concrete support, from a contempor- ary anımal, to Rın£y’s (1951) proposal to consider the first premolar present in a deer jaw as being P;. It ıs also a stricking case of genetic atavism. It is however unclear whether the genes responsible for the production of P, are present in only a few contemporary deer, or else are present ın all, but are being repressed in some way. Finally, the present case gives us a chance to examine, in a fresh specimen, what the dentition was presumably like millions of years ago. I am grateful t0 ROLAND LEMIEUX of the Ministere du Loisir, de la Chasse et de la Peche of Quebec who provided the deer and to MıcHEL Bourassa who photographed the mandible. Literature BANFIELD, A. W. F. (1961): A revision of the reindeer and carıbou, genus Rangıfer. National Museum of Canada, Bulletin 177, 1-137. BUBENIK, A.; WURTZINGER, H. (1967): Beidseitiger erster Prämolar im Unterkiefer des Rehes, Capreolus capreolus Linne, 1758. Säugetierkdl. Mitt. 15, 35-39. CARROLL, R. L. (1988): Vertebrate paleontology and evolution. New York: W. H. Freeman. CHıLD, G.; Rıney, T. (1964): Abnormal dentition in the common duiker (Sylvicapra grimmia), impala (Aepyceros melampus) and Sharp’s grysbuck (Raphicerus sharpei). Occasıonal Papers National Museum, Southern Rhodesia, no. 27B. Frick, C. (1937): Horned ruminants of North-America. Bull. Amer. Mus. Nat. Hist., 69, 1-669. Fully formed first lower premolars in a White-tailed deer 123 FuULLER, W. A. (1954): The first premolar and the canıne tooth in bison. J. Mammalogy 35, 454-456. MATTHEw, W. D. (1908): Osteology of Blastomeryx and phylogeny of the American Cervidae. Bull. Amer. Mus. Nat. Hist. 24, 535-562. MecnH, L. D., FRENZEL, JR., L. D.; Karns, P. D.; KueHn, D. W. (1970): Mandibular dental anomalies in white-tailed deer from Minnesota. J. Mammalogy 51, 804-806. MEyeEr, P. (1985): Beidseitiger P, im Unterkiefer eines Rehs (Capreolus capreolus L., 1758). Z. Jagdwiss. 31, 120-123. MILLER, F. L.; TEssıer, G. D. (1971): Dental anomalıes in carıbou, Rangifer tarandus. J. Mammalogy 52, 164-174. MOoRAN,R. J.; FAIRBANKS, T. (1966): Vestigial first premolar in Michigan elk. J. Mammalogy 47, 514. Rıney, T. (1951): Standard terminology for deer teeth. J. Wild. Manage. 15, 99-101. ROBINETTE, W. L.; ARCHER, A. L. (1971). Notes on ageing criteria and reproduction of Thomsons’s gazelle. E. Afr. Wild. J. 9, 83-98. ROBINETTE, W. L.; Hancock, N. V.; Jongs, D. A. (1977): The Oak creek mule deer herd in Utah. U. S. Fish and Wild. Ser. Pub. No. 77-15, 148 pp. Vıcar, C. R.; MACHORDOM, A. (1987): Dental and skull anomalies in the Spanish wild goat, Capra pyrenaica Schinz, 1838. Z. Säugetierkunde 52, 38-50. VIRET, J. (1961): Artiodactyles. In: Traite de Paleontologie. Ed. by. J. PivErEAu. Paris: Masson et Cie. 6(2), 887-1084. Author’s address: CYRILLE BARRETTE, Departement de Biologie, Universite Laval, Ste-Foy, Quebec, GI1K 7P4, Canada Z. Säugetierkunde 56 (1991) 124-125 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Chromosomes of the Argentine Andean mouse, Akodon andinus (Cricetidae: Sigmodontinae) By Marıa BEATRIZ Espıinosa, A. D. VrtULLo and MarıA $. MERANI Centro de Investigaciones en Reproduccion, Facultad de Medicina, Universidad de Buenos Aires, Buenos Aires, Argentina Receipt of Ms. 30. 7. 1990 Acceptance of Ms. 30. 10. 1990 Cytogenetic analysis was performed on six specimens of A. andinus (Philippi, 1858) (two males and four females) collected at Plaza de Mulas, in the eastern slope of the Aconcagua, province of Mendoza, Argentina, at 4,200 m above sea level. Specimens were identified as A. andinus by external and cranial characteristics by Prof. JuLIO CONTRERAS, and deposited at the Systematic Collection of the CECOAL, province of Corrientes, Argen- tina. Chromosomes were obtained from bone marrow by routine methods, and Giemsa- Stained G-banded according to SEABRIGHT (1971). All specimens studied showed a 2n = 52 (NF = 60) karyotype with three distinctive pairs of medium- and small-sızed metacentrice autosomes, and 22 pairs of telocentrics decreasing gradually ın size (see Fig. ). Besides minimal variations in positioning of metacentric autosomes, this karyotype ıs similar both in diploıd number, chromosomal morphology and lengths to those previously reported for A. xanthorhinus, A. olivacens, A. longipilis (RODRIGUEZ et al. 1983), and A. ıllnteus (Lıascovich et al. 1989). G-banded chromosomes were arm-to-arm compared with a banded karyotype of A. olivaceus (kındly provided by Dr. L. VıDAaL-RıojJa from the Instituto Multidisciplinario de Biologıa Celular, La Plata, Argentina), showing entire sımilarıty between banding patterns Giemsa-stained karyotype of Akodon andınus,; 2n = 52, NF = 60 U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5602-0124 $ 02.50/0 Chromosomes of the Argentine Andean mouse, Akodon andinus 125 of both species. Similarly, whole G-band identity has also been reported for A. xanth- orhinus, A. olivaceus, and A. longipilis from Chile (RODRIGUEZ et al. 1983), and A. illuteus and A. longipilis from Argentina (LıascovichH et al. 1989). These results strongly support the contention that speciation in the southern lineage of Akodon proceeded without chromosomal change from a generalized 2n = 52 karyotype, which lıkely ıs the primitive condition for Akodon (Reıc 1987). Although bandıng patterns had not been reported, the same 2n = 52 karyotype has been described for A. sanborni and A. jelski, and the related genera Chelemys and Geoxus (PEARSON 1984). We thank Dr. A. CAastro-VAzQuEz for providing the anımals employed in this study, Prof. ]. CONTRERAS for the identification of specımens, and Dr. L. Vıpar-RıoJA for providing us with original pictures of G-banded karyotypes of A. olivaceus. Thanks also to Dr. E. RoLpän, for his comments and improving of the Englısh. References LıascovIcH, R.; BARQUEZ, R. M.; Reıc, ©. A. (1989): A karyological and morphological reassess- ment of Akodon (Abrothrix) ıllutens Thomas. J. Mammalogy 70, 386-391. Pearson, ©. P. (1984): Taxonomy and natural history of some fossorial rodents of Patagonia, southern Argentina. J. Zool., London 202, 225-237. Reıc, ©. A. (1987): An assessment of systematics and evolution of the Akodontini, with the description of new fossil species of Akodon, (Cricetidae: Sigmodontinae). Fieldiana: Zoology, New Series 39, 347-399. RODRIGUEZ, M.; MonNToOYA, R.; VENEGAS, W. (1983): Cytogentic analysıs of some Chilean species of the genus Akodon Meyen (Rodentia, Cricetidae). Caryologia 36, 129-138. SEABRIGHT, M. (1971): A rapıd banding technique for human chromosomes. Lancet 2, 971-972. Authors’ addresses: MARIA B. EspınosA, and MARIA $. MERANI, Centro de Investigaciones en Repro- duccıön, Facultad de Medicina, Universidad de Buenos Aires, Paraguay 2155, pıso 10, RA-1121 Buenos Aires, Argentina; ALFREDO D. VITULLO, Station de Phy- sıologie Anımale, Unite Biologie de la Fecondation, INRA, F-78 350 Jouy-en-Josas, France BEKANNTMACHUNG CNEVA Laboratoire d’Etudes sur la rage et la pathologie des anımaux sauvages Symposium on Health and Management of free ranging Mammals Nancy/France, 14-18 October 1991 Co-sponsored by: WHO-OIE-IUCN Topics Invasion, spread and persistence of contagıious diseases Toxic, Food selection Behaviour and ecology of hosts, vectors, reservoir and victims Population biology, population dynamics of diseases Host/parasites coevolution Diseases of threatened mammals, conservation biology Control of vectors, pest and feral or stray anımals Immunizatıon against rabies, baits, biomarkers, aversive conditioning, repellent Send your tentative title of communication to: CNEVA/LERPAS 20e annıversaire BP 9 54220 Malzeville (FRANCE) el @3) 83 292803 Tacsımıle (Bo) 83 2933 13 BU@EIBESERECTUNGEN PoLLAK, G. D.; CAsseDay, J. H.: The Neural Basis of Echolocation in Bats. Zoophysiol- ogy, Vol. 25. Ed. by W. BuRGGREn, $. Isis, H. LANGER, G. NEUWEILER, D. J. RANDALL. Berlin-Heidelberg-New York-London-Paris-Tokyo-Hong Kong: Springer Verlag 1989. 143 pp., 82 figs. DM 128,-. ISBN 3-540-50520-2 This monograph reviews the physiology of the auditory system in bats, from periphery to midbrain. The authors focus on recent neuroanatomical and electrophysiological investigations ın the horseshoe bats (Rhinolophus) and the mustache bats (Pteronotus); comparisons between these constant-fre- quency bats and frequency-modulated species are somewhat rare. The first chapter gives an introduction to echolocation by characterizing the different types of ultrasonic calls and their use in various orientation situations. The main parts deal with different aspects of the major auditory nuclei of the bats’ brainstem, the cochlear nucleus, olıvary complex, lateral lemniscus and inferior colliculus. It is shown that the principal characteristic of the system is a tonotopic organization, remapping the frequency arrangement of the cochlea in the nuclei of the auditory system. Intensively described are the monaural and binaural pathways, emanating from the anteroventral cochlear nucleus. These parallel pathways extract ditferent information from the acoustical signal and converge at the inferior colliculus. A great number of investigations deal with the physiological properties of auditory neurons. The characteristic neurons of the different nuclei are considered. For instance the E-I units (excited by the contralateral, inhibited by the ipsilateral ear) in the lateral superior olive and the colliculus inferior, that are sensitive to interaural intensity disparities. It is shown that these neurons play a crucial role for the localızation of sounds at different azımuthal positions. Equally important are “constant latency neurons” that encode range information. This volume is written with great scientific competence, and gives an excellent insight in the neural processes associated with hearing in bats. Due to the instructive introductions, the complex subjects of the different chapters are easy to understand, even for the interested reader who is not professionally involved in neurophysiology. U. ScHMIDT, Bonn HEIDECKE, D.; StuBBE, M.: Populationsökologie von Fledermausarten. Teil I u. II. Kongreß- und Tagungsberichte der Martin-Luther-Universität Halle-Wittenberg, WB 1989/20 (P36); 368 S. August-Bebel-Str. 13, O-4010 Halle-Saale. ISBN 3-86010-194-3 Dieses ist der Referateband der 4. DDR-Tagung zur Fledermausforschung vom 11.-13. 11. 1988 in Meisdorf, Kreis Aschersleben. Veranstalter waren die Sektion Biowissenschaften der Martin-Luther- Universität in Halle-Wittenberg, die Biologische Gesellschaft der DDR (hier die AG-Fledermaus- forschung) sowie das Institut für Landschaftsforschung und Naturschutz in Halle. Die beiden Bände enthalten insgesamt 33 Referate und sind dem großen Fledermausforscher JOHANN CHRISTIAN DANIEL SCHREBER (250. Geburtstag) gewidmet. Einleitend wird die Bedeutung von Halle für die klassische Fledermausforschung vom 18. bis zum 20. Jahrhundert dargestellt (Biographischer Anhang). Eine Gruppe von Vorträgen befaßt sich mit dem Status der Fledermausarten in den verschiedenen Bezirken der DDR. Berichtet wird ferner über die Situation einzelner Arten, über Arealgrenzen, Erstnachweise und über besondere Wochenstuben. In 8 Referaten wird der Einsatz von Fledermauskästen behandelt (Arbeitsmethoden). Den Abschluß bilden Vorträge über die Echoortung (Einsatz von Bat-Detektoren). Beide Bände geben einen guten Überblick über den gegenwärtigen Stand der Fledermausforschung in der DDR. E. Kurzer, Tübingen Kıörzuı, F.: Ökosysteme: Aufbau, Funktionen, Störungen. 2., völlig überarb. Aufl. Stuttgart, New York: Gustav Fischer Verlag 1989. UTB Nr. 1479. 464 S., 166 Abb., 87 Tab. DM 44,80. ISBN 3-437-20403-3 Die 1. Auflage erschien 1980 unter dem Titel „Unsere Umwelt und wir“. Wie schon aus dem Literaturverzeichnis zu ersehen, das etwa zur Hälfte nach 1979 erschienene Zitate enthält, ist die jetzt vorliegende Fassung nicht nur im Titel verändert, sondern auch erheblich umgestaltet. Das Buch ist in vier Abschnitten gegliedert: Struktur, Kreisläufe, organismische Beziehungen und menschliche Ein- flüsse. Im Vordergrund stehen weniger Okosysteme (= Biozönosen + Biotope) als vom Menschen ausgehende Umweltgefahren, die mit vielen Zahlen in Tabellen und Diagrammen untermauert werden. So erfährt man eine Menge zu den Themen saurer Regen, Ozonschild, Treibhauseffekt, Müllentsorgung und Recycling, Schwermetalle, Herbizide und Radionuclide in der Natur, Ausbrei- tung von Wüsten oder Nutzungsänderung in der Landwirtschaft. Zu diesen Fragen dürfte das Buch auch wertvoll und wichtig sein. Leider gilt das weniger für zoologische Aspekte. So werden bei den 128 Buchbesprechungen guten, alten Klimaregeln inter- und intraspezifische geographische Variabilität vermischt, Rehe sollen in den Polargebieten wesentlich größer sein als in südlicheren Breiten. (Wo kommen sie in den Polargebieten vor? Skandinavische Rehe sind nicht größer als mitteleuropäische.) Unter den Beispie- len für konvergente Tierformen findet man „eurasisches Steppenmurmeltier und Präriehund“ oder „Oppossum/Ratte“ (S.7). Nach S.22 mußten beim Übergang aufs Land Lungen und Nieren entwickelt werden. Tabelle 115 zeigt die Beziehungen zwischen Lemming-Häufigkeit und Wollgras- Dichte nach Tastr und Kareıa (nicht Karena). Die Unterschrift behauptet, die Arbeit sei 1971 durchgeführt worden, sie bezieht sich aber, wie die zugehörige Graphik ergibt, auf den Zeitraum von 1962-1970. Die Ordinate gibt nicht die in 100 Nächten in Fallen gefangenen Lemminge, sondern die Fangzahlen bei 100 Fallennächten wieder. Mit der Definition von Fitneß als „die Fähigkeit auch unter ungünstigen Bedingungen Nachkom- men zu erzeugen“ kann man ebenso wenig einverstanden sein wie mit RNS „als wichtiger Transfer- oder Botensubstanz für organische Stoffe (z.B. Eiweiße)“. Unter der „Erklärung der Fachausdrücke“ findet man eine ganze Reihe weiterer zumindest sehr mißverständlicher Formulierungen bei ATP, Aminosäuren, Chromosom, Assimilation, organische Substanz, Parasit, Polypeptid, Sukzession, Trittum, Wirkungsgrad. Auf Seite 93 liest man, daß Wasser „mit Säuren und Basen gepufferte Lösun- gen“ bildet. Auf Seite 239 regt Streß die Nebennierenrinde zu vermehrter Adrenalinausschüttung an. Bei den in Tab. 45 aufgeführten Nahrungsketten handelt es sich um Beispiele aus Nordamerika, bei denen es oft schwer fällt, die betreffende Art zu identifizieren, weil nur deutsche Trivialnamen vorkommen. Eine Nahrungskette „Tanne-Reh-Wolf“ (S. 225) dürfte es kaum geben. In der Nah- rungspyramide Abb. 108 findet man als Zusatz zur Stufe der „Ruderfußkrebschen“: z.B. B. Krill, Euphausia superba“. Auf Seite 217 wird Stellaria media als Sternmiere bezeichnet. Selbst bei dem Schachspiel stimmt etwas nicht, wenn Patt als Situation erklärt wird, „wo beide Könige auf ihrem Feld festsitzen und sich gegenseitig blockieren“. Die Quellen der Abbildungen sind zwar genannt, aber nicht ım Literaturverzeichnis zitiert. Die zahlreichen Lapsus machen mißtrauisch hinsichtlich der Richtigkeit sonstiger Angaben. Deshalb kann das in Druck und bildlicher Gestaltung (abgesehen von den oft zu kleinen Fotos) sehr ansprechende Buch leider nur mit erheblichen Bedenken empfohlen werden als ein Werk, dessen Aussagen man sich nicht ungeprüft zu eigen machen darf. J. NIETHAMMER, Bonn EISENBERG, J. F.: Mammals of the Neotropics. Vol. I: The Northern Neotropics. Panama, Columbia, Venezuela, Guyana, Suriname, French, Guiana. Chicago, London: University Chicago Press 1989. 449 pp., num. ıillustr., 21 colour plates, num. tables. Cloth US $ 97.75 (ISBN: 0-226-19539-2); paper US $ 40.45 (ISBN: 0-226-19540-6) Based on his own extensive fieldwork in South and Central America, the well-known North American mammalogist JOHN F. EISENBERG has published a remarkable book, which will certainly become a MUST on the bookshelves of people interested in neotropical mammals. After a short introduction to historical biogeography and contemporary habitats of the northern part of South America, the author deals with the order Marsupialia and 12 eutherian orders that live ın the above-mentioned part of the continent. It is very helpful that the author organızes the text according to a strict format. The orders are presented by giving a diagnosis, a short summary on their geographical distribution, followed by a brief account on palaeontological history and on classifica- tion. Each family that belongs to one of the mammalian orders living in the northern neotropices is introduced by an account on taxonomy, a diagnosis, the discription of its geographical distribution and a summary on its natural history. The different species belonging to the famılıes are described; essential measurements are given in these descriptions and colours as well as other physical charac- teristics are summarized. A clear description of the range and habitat of the respective species is given and a standardized map shows its distribution. An account of the natural history concludes the section on each species. At the end of each chapter on a mammalian order, a modern, useful and truely international list of references ıs supplied. The text of this most noteworthy book is concluded by two interesting chapters. One deals with speciation and faunal affinities of mammals in the northern neotropics, the other wıth mammalıan community ecology. As islands are useful indicators of speciation rates, EISENBERG discusses the mammals of Trinidad, Tobago, Isla Margarıta, Grenada, and three of the Netherland Anuilles. According to the author little specific distinctiveness has evolved on the above-mentioned islands, but sub-specific differences can be noted. To conclude, an index of scientific and one of common names helps the reader to make full and easy use of the materials compiled in this volume. The referee is certain that this book, and, most probably, also the forthcoming companion, will soon become most useful sources of reference, not only for scientists working on that continent, but also for those interested in general zoogeography, ecology or taxonomy. Researchers might wish to apply the remarkable wealth of data that was made available by JoHn F. EISENBERG for comparative studies or for specialized investigations in neotropical mammals. P. LANGER, Gießen Deutsche Gesellschaft für Säugetierkunde: Referate, Vorträge und Posterdemonstrationen der 64. Hauptversammlung 1990 E; Hauptziel der Deutschen Gesellschaft für Säugetierkunde ist, auf ihren Jahrestagungen über Säugetiere arbeitende Wissenschaftler verschiedenster Fachrichtungen zusammenzuführen, den Gedanken- und Erfahrungsaustausch anzuregen, um so Erkenntnisse aus den einzelnen Forschungsgebieten zu integrieren. Die Kurzfassungen der Vorträge und Posterdemonstrationen von der 64. Haupt- versammlung, die in diesem Jahr in Osnabrück stattfand, liegen wiederum als Sonderheft der »Zeitschrift für Säugetierkunde« vor. Die Tagung war durch drei Themenschwerpunkte gekennzeichnet, die ökologische, ontogenetische und ernährungs- physiologische Fragestellungen behandeln. Zusätzlich wurde eine große Zahl freier Beiträge angeboten. Diese auch in der Vergangenheit geübte Art der Themen- darbietung soll die Breite der aktuellen Säugetierforschung widerspiegeln. Die säugetierbiologische Arbeit war in der Ethologie von Anfang an ein Forschungsschwer- punkt. Im Vordergrund stehen Projekte mit öko-ethologischer Fragestellung. Da sich dafür stenöke Säugetiere besonders eignen, werden bevorzugt silvicole und ripicole Arten in Gehegen und im Freiland beobachtet. Eine intensive Freilandarbeit, auch auf Exkursionen, förderte die Untersuchung von Kleinsäugetiergemeinschaften. Die Kurzfassungen der Vorträge und Posterdemonstrationen der Deutschen Gesellschaft für Säugetierkunde sind ab der 58. Hauptversammlung 1984 in Göttingen noch lieferbar. Zu beziehen durch jede Buchhandlung. % Deutsche Gesellschaft für Säugetierkunde, 64. Hauptversammlung in Osnabrück, 23. bis 27. September 1990. Kurzfassungen der Vorträge und Posterdemonstrationen. Heraus- gegeben von Dr. Christel Schmidt, Bonn, und Prof. Dr. Rüdiger BAJL Schröpfer, Osnabrück. 1990. 55 Seiten. Kartoniert 24,- DM Verlag Paul Parey - Spitalerstraße 12 - 2000 Hamburg 1 P/REY Subscription information for USA, Canada and Mexiko obtainable from Paul Parey Scientific Publishers/IR Unltd., 35 West 38th Street, Suite 3 W, New York, NY 10018, USA. Erscheinungsweise und Bezugspreis 1991: 6 Hefte bilden einen Band. Jahresabonnement Inland: 338,- DM zuzüglich 13,80 DM Versandkosten, Gesamtpreis 351,80 DM einschließlich 7% Mehrwert- steuer. Jahresabonnement Ausland: 338,—- DM zuzüglich 18,- DM Versandkosten. Das Abonne- ment wird zum Jahresanfang berechnet und zur Zahlung fällig. Es verlängert sich stillschweigend, wenn nicht spätestens am 15. November eine Abbestellung im Verlag vorliegt. Die Zeitschrift kann bei jeder Buchhandlung oder bei der Verlagsbuchhandlung Paul Parey, Spitalerstraße 12, W-2000 Hamburg 1, Bundesrepublik Deutschland, bestellt werden. Die Mitglieder der „Deut- schen Gesellschaft für Säugetierkunde“ erhalten die Zeitschrift unberechnet im Rahmen des Mitgliedsbeitrages. Z. Säugetierkunde 56 (1991) 2, 65-128 .. Eikmeier/Fellmer/Moegle Hans Eikmeier ee Lehrbuch Lehrbuch der jer gerichtlichen use Gerichtlichen recht - Öffentliches Veterinärwesen Herausgegeben von Prof. Dr. Hans Eikmeier, Gießen, Rechtsanwalt Eberhard Fellmer, Hanseatisches Oberlandesgericht, Hamburg, und Prof. Dr. Horst Moegle, Regierungspräsidium Tübingen. Mit Beiträgen von Prof. Dr. Hartwig Bostedt, Gießen, Rechtsanwalt Alexander Busse, Hamburg, Prof. Dr. Hans Eikmeier, Gießen, Dr. Ursula Eikmeier, Gießen, Rechtsanwalt Eberhard Fellmer, Hamburg, Rechtsanwalt Peter Kiel, Hamburg, Prof. Dr. Horst Moegle, Tübingen, Prof. Dr. Heinrich Müller, Gießen, Dr. Priska Müller, Wiesbaden, Dr. Marie-Luise Nagel, Gießen, und Dr. Gottfried Schreiber, Fritzlar. 1990. 256 Seiten. 24 x 16,5 cm. Gebunden DM 78,- ISBN 3-489-56616-5 Ein neues Buch unter dem bewährten Titel „Lehrbuch der Gerichtlichen Tierheilkunde“: Es setzt die Tra- dition des in 11 Auflagen erfolgreichen und seit Jahren vergriffenen Werkes von Eugen Fröhner fort. Die Konzeption dieses langerwarteten, modernen Lehrbuches entspricht den heutigen Anforderungen an die Gerichtliche Tierheilkunde. Das gilt für den Abschnitt Tierkaufrecht und die ausführliche Darstel- lung des Haftpflichtrechts und der tierärztlichen Sorgfaltspflicht ebenso wie für die Kapitel Tierarzt und Gericht, Tierschutzrecht, Berufs- und Standesrecht, öffentliches an u Tierhalter- und Tierhü- terhaftung, Tierversicherungen und Doping. Aus dem Inhalt: Tierarzt und Gericht - Gerichtliche Verfahren - Tierarzt als Angeklagter, Beschuldigter, Betroffener - Tierarzt als Sachverständiger und (sachverständiger) Zeuge - Tierärztliche Atteste, Proto- kolle, Gutachten - Tierkaufrecht - Tierkaufrecht in angrenzenden deutschsprachigen Ländern - Gewähr- leistung beim Tierkauf - Kaufvertrag — Gesetzliche Regelung / Vertragsschluß / Vertragspflichten - Ab- weichung von der gesetzlichen Regelung durch Parteivereinbarung - Vertragliche Anderung der gesetz- ı lichen Gewährleistung und die Zusicherung von Eigenschaften - Tierarzt und Tierkauf - Gesetz zur Re- gelung des Rechts der Allgemeinen Geschäftsbedingungen (AGBG) und seine Bedeutung für Tierkauf- jerheilkunde: Tierarzt und Gericht - Tierkauf - Haftpflicht - Sorgfaltspflichten - Tierversicherung - ‚Tier- schutz - Doping - Berufs- und Standes-' m 7B (TEE ER en verträge und Tierarztverträge - Berufshaftpflicht des Tierarztes - Haftung für das Verschulden des tier- ärztlichen Hilfspersonals (Erfüllungsgehilfen) - Allgemeine Sorgfaltspflichten des Tierarztes - Haftung des Tierarztes bei hoheitlichen Aufgaben - Berufshaftpflichtversicherung des Tierarztes - Haftpflicht des Tierhalters und Tierhüters - Tierarzt und Tierversicherung - Tierschutzrecht - Verordnung über die Mel- dung von in Tierversuchen verwendeten Wirbeltieren vom 1. August 1988 - Verordnung zum Schutz von Tieren bei der Beförderung in Behältnissen vom 20. Dezember 1988 - Doping - Berufs- und Standes- recht - Bundestierärzteordnung - Niederlassung in eigener Praxis - Öffentliches Veterinärwesen - Aufbau der Veterinärverwaltung - Zugang zum amtstierärztlichen Dienst - Tierärzte in internationalen Organisa- tionen - Übersicht über die Gewähr-, Anzeige-, und Verjährungsfristen, Gerichte und Zuständigkeiten - Übersicht über die Inkubationszeiten und Nachweismethoden der forensisch wichtigsten Infektions- krankheiten. PAJL Berlin und Hamburg P/REV lo 56 (3), 129-192, Juni 1991 ISSN 0044-3468 C21274F EITSCHRIFT FÜR AUGE TIERKUNDE INTERNATIONAL JOURNAL FMAMMALIAN BIOLOGY 2 rgan der Deutschen Gesellschaft für Säugetierkunde | doper, T. J.; Tait, A. l.; Christian, S. F.; Fee, D.: Excavation of three badger (Meles meles L.) seits. — Die Struktur dreier Baue des Dachses (Meles meles 9) | 129 ‘aylor, P. J.; Campbell, G. K.; Meester, J. A. J.; van Dyk, Deborah: A study of allozyme evolution in African mongooses (Viverridae: Herpestinae). — Untersuchungen zur Evolution afrikanischer Mangusten (Viverridae: Herpestinae) aufgrund von Allozymen 135 (vam, T.: Reproduction in the European Iynx, Lynx Iynx. — Fortpflanzung beim europäischen Luchs, Lynx Iynx 146 Zeif, Ursula; Klingel, H.: Hiding behaviour in wild Gerenuk (Litocranius walleri) fawns. — Abliegeverhalten bei Gerenuk-Kitzen (Litocranius walleri) 159 aranjon, L.; Vassart, M.; Greth, A.; Cribiu, E.-P.: Genetic study of Sand gazelles (Gazella subgutturosa marica) from Saudi Arabia. Chromosomal and isozymic data. — Genetische Untersuchungen an Sandgazellen (Gazella subgutturosa marica) aus Saudi-Arabien. Chromosomale und elektrophoretische Daten 169 erzig-Straschil, Barbara; Herzig, A.; Winkler, H.: A morphometric analysis of the skulls of Xerus inauris and Xerus princeps (Rodentia; Sciuridae). - Morphometrische Analyse der Schädel von Xerus inauris und Xerus princeps (Rodentia, Sciuridae) na ‚Nissenschaftliche Kurzmitteilungen | nsorge, H.: Diphyodont dentition of first premolar in the Red fox Vulpes vulpes (Linne, 1758). — Diphyodonte Dentition des ersten Prämolaren beim Rotfuchs Vulpes vulpes (Linne, 1758) 188 Rookmaaker, L. C.: The scientific name of the bontebok. — Der wissenschaftliche Name für den Buntbock 190 Buchbesprechung 192 | LE Paul Parey Hamburg und Berlin FIERAUSGEBERVEDIENORS P. J. H. van BreE, Amsterdam - W. FIEDLER, Wien — H. Frick, München — W. HERRE, Kiel - H.-G. Krös, Berlin - H.-]. Kuhn, Göttingen - E. KULZER, Tübingen — B. Lanza, Florenz -— W. Maıer, Tübingen - J. NIETHAMMER, Bonn — Anne E. Rasa, Bonn - H. ReıcHstein, Kiel - M. Rönrs, Hannover — H. SCHLIEMANN, Hamburg — D. STARCcK, Frankfurt a. M. - E. THEnIUs, Wien - P. VoGEL, Lausanne SSENUILIDTTINEAEIDITTONRIANL OBIEILGIE D. Kruska, Kiel - P. LANGER, Gießen This journal is covered by Biosciences Information Service of Biological Abstracts, and by Current Con- tents (Series Agriculture, Biology, and Environmental Sciences) of Institute for Scientific Information Die Zeitschrift für Säugetierkunde veröffentlicht Originalarbeiten und wissenschaftliche Kurzmittei- lungen aus dem Gesamtgebiet der Säugetierkunde, Besprechungen der wichtigsten internationalen Literatur sowie die Bekanntmachungen der Deutschen Gesellschaft für Säugetierkunde. Verantwort- licher Schriftleiter im Sinne des Hamburgischen Pressegesetzes ist Prof. Dr. Dieter Kruska. Zusätzlich erscheint einmal im Jahr ein Heft mit den Abstracts der Vorträge, die auf der jeweiligen Hauptversammlung der Deutschen Gesellschaft für Säugetierkunde gehalten werden. Sie werden als Supplement dem betreffenden Jahrgang der Zeitschrift zugeordnet. Verantwortlich für ihren Inhalt sind ausschließlich die Autoren der Abstracts. Manuskripte: Manuskriptsendungen sind zu richten an die Schriftleitung, z. Hd. Prof. Dr. Dieter Kruska, Institut für Haustierkunde, Biologie-Zentrum, Neue Universität, Olshausenstr. 40-60, W-2300 Kiel, Bundesrepublik Deutschland. Für die Publikation vorgesehene Manuskripte sollen gemäß den „Redaktionellen Richtlinien“ abgefaßt werden. In ihnen finden sich weitere Hinweise zur Annahme von Manuskripten, Bedingungen für die Veröffentlichung und die Drucklegung, ferner Richtlinien für die Abfassung eınes Abstracts und eine Korrekturzeichentabelle. Die Richtlinien sind auf Anfrage bei der Schriftleitung und dem Verlag erhältlich. Sonderdrucke: Anstelle einer Unkostenvergütung erhalten die Verfasser von Originalbeiträgen und Wissenschaftlichen Kurzmitteilungen 50 unberechnete Sonderdrucke. 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For copying from back volumes of this journal see “Permissions to Photo-Copy: Publisher’s Fee List’ of the Eee: Fortsetzung 3. Umschlagseite © 1991 Paul Parey. Verlag: Paul Parey, Hamburg und Berlin. Anschriften: Spitalerstr. 12, D-2000 Hamburg 1; Seelbuschring 9-17, D-1000 Berlin 42, Bundesrepublik Deutschland. — Printed in Germany by Westholsteinische Verlagsdruckerei Boyens & Co., Heide/Holst. Z. Säugetierkunde 56 (1991) 129-134 © 1991 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-3468 Excavation of three badger (Meles meles L.) setts By 1. ]. Roper, A, I. TAıs, S. F./Cartstian and D. FEE School of Biology, University of Sussex and East Sussex County Council, Lewes Receipt of Ms. 9. 8. 1990 Acceptance of Ms. 30. 10. 1990 Abstract Excavated and described three badger setts in the south of England. Two of the setts were classified as subsidiary setts and the third as an annex. Each consisted of a main network of interconnected tunnels associated, in two of the setts, with 2-3 small separate burrows. Total tunnel lengths were 28, 30 and 53 m respectively and estimated total volumes of the setts were 1.1, 1.1 and 2.1 m’. No sett exceeded a depth of more than 1 m and mean depths of the tunnel systems were 61, 56 and 52 cm respectively. All three setts contained bedding material (dry grass) but only one (the annex) contained enlarged chambers, faeces and badger bones. Introduction One of the most strıking features of the European badger ıs ıts habıt of constructing complex underground burrows or “setts”. Within the confines of their setts badgers sleep, breed, overwinter and take refuge when alarmed; and ın addition the sett forms a focus for social interactions between the different members of a badger group (NEAL 1977; KRUUK 1989). Although a few setts have been accurately described and mapped (LIKHACHEV 1956; JEnsEn 1959; CowLin 1967; FREwIN 1976; LEEson and Mırıs 1977; ROPER et al. 1991) we still know comparatively little about the structure or internal environment of setts; and little serious thought has been given to the question of why badgers need elaborate setts in the first place (for a recent review see NEAL and RoPpEr 1991). In the present study we describe three setts that were about to be destroyed by a road development. Prior to excavating the setts in question an attempt was made to determine whether or not they belonged to separate social groups and whether they should be classıfied as main setts, subsidiary setts, annexes or outliers (Kruuk 1978; Bock 1986; THORNTON 1988). Material and methods Study area The excavated setts were located in open farmland immediately to the north-west of Brighton, about 3.5 km inland from the south coast of England. The landscape consisted of a range of rounded chalk hills (the South Downs) containing patches of permanent pasture and scrub (mainly hawthorn Crataegus monogyna, elder Sambucus nigra, blackthorn Prunus spinosa and brambles Rubus spp.) in between larger areas of arable planting (mostly of wheat Triticum aestivum). Location of setts A survey carried out in April 1989 revealed sıx setts falling into two separate spatial groups, one of four sets and the other of two (see Fig. 1). Of the group of four setts, sett 1 was by far the largest, having 9 entrances of which 7 showed clear signs of use. Setts 2 and 3, by comparison, had only 3 and 4 entrances respectively at the time of the survey, while sett 4 had a single entrance; and none of these setts appeared heavily used. Given their close proximity to one another it seemed lıkely that all four setts were within the territory of a single social group, and this was subsequently confirmed when the territory boundaries were mapped using the technique of bait-marking (Kruuk 1978). U.S. Copyright Clearance Center Code Statement: 0044-3468/91/5603-0129 $ 02.50/0 130 MS ERoper ASIA SEER@hnstansandeD Mhee Fig.1. Map of the study area showing position of setts (numbered 1-6). Dotted areas: grassland. Hatched areas: housing. Open areas: arable fields. Solid line: estimated territory boundary surround- ing setts 1-4. Double broken line: route of new road Of the pair of setts in the second group, sett 5 had 11 entrances of which 7 showed clear signs of use, while sett 6 had 5 entrances of which 4 showed clear signs of use. The two setts were connected by a well-worn path and although the territory boundaries surrounding the setts were not determined, it was clear from the proximity of the setts and their interconnectedness that both belonged to a single social group. Of the sıx setts ıdentified in Fig. 1 only setts 2, 3 and 6 were excavated since the remaining setts were not threatened by the new road. Setts 2 and 3 were located on a north-east facıng slope, sett 2 at the top of the slope (altitude 95 m) and sett 3 near the bottom (altitude 80 m). Sett 2 was surrounded by dense scrub containing occasional mature sycamore trees (Acer pseudoplatanus), while sett 3 was ın open grassland dotted with occasional hawthorn trees (Crataegus monogyna). Sett 6 was located on a west-facing slope at an altitude of 102 m in open grassland dotted with hawthorn trees. In all three cases the soil consisted of an open sandy loam to a depth of about 1.5 m, over chalk subsoil. Method of excavation The excavations were carried out in April 1990. Several months prior to this (in October 1989) all three setts were fenced off with one-way gates to exclude badgers from them (for a description of the method see Harrıs et al. 1988). When it was clear that the setts were no longer occupied the fences were removed and excavation was immediately begun. The excavation itself was carried out with a small mechanical digger (Kıbota KH20) or by hand when features such as tunnel intersections and nest chambers were encountered. Sett entrances were first identified with numbered surveyor’s poles and plotted on a scale map. A 1.5 m-deep trench was then dug across one end of the sett, 2 m beyond the furthest entrance. The wall of the trench nearest the sett was gradually dug away, thus moving the trench progressively through the sett and exposing the underground tunnel system. As excavation proceeded tunnels and chambers were measured and plotted on a scale map, and the depth of the floor of each tunnel relative to the soil surface was measured to the nearest 10 cm, at intervals of approximately 2 m. The length of each sett was measured as the furthest distance between any two extremities of the tunnel system. Sett width was measured orthogonal to the line joining these two points. Sett volume was calculated from the dimensions of tunnels and chambers, using the formulae of LEEsoN and Mırrs (1977). Table. Dimensions and contents of the three excavated setts Badger remains Chambers Bedding Latrines Entrances "® S s S) - =: Ss KB} = Total tunnel Estimated Excavation of three badger (Meles meles L.) setts 131 Results The Table summarises the dimensions and contents of the three setts and Fig.2 shows a plan view of each sett. Sett 2 This consisted of four separate burrow systems, with tunnels totallıng 28 m ın length. Three of the burrows consisted of sımple shallow tunnels only a few metres long, with single entrances, all of which were disused. The fourth system was larger and more complex, consisting of an interconnected net- work of tunnels radıating off from a central entrance, with four additional entrances around its outer limits. Two of the latter entrances were disused. No obvious chambers were found, though tunnels tended to widen out where they intersected. Nor was there any evidence of badger faeces or badger remaıns (bones, teeth etc). A small amount of bedding material was found at one point in the main tunnel system, consisting of dry grass (about 1 litre) together with several chocolate wrappers and potato-crisp packets. Sev- eral of the tunnels had been extended by rabbits and at one point the excavation disclosed a rabbit nest containing four juveniles. Sett 3