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ISSN: 1578-665 X An international journal devoted to the study and conservation of animal biodiversity

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  • Form

    erly M

    iscelln

    ia Zoolgica

    2001

    AnimalBiodiversity Conservation24.1

    and

  • Editor executiu / Editor ejecutivo / Executive EditorJoan Carles SenarJoan Carles SenarJoan Carles SenarJoan Carles SenarJoan Carles Senar

    Secretria de Redacci / Secretaria de Redaccin /Managing EditorMontserrat FerrerMontserrat FerrerMontserrat FerrerMontserrat FerrerMontserrat Ferrer

    Consell Assessor / Consejo asesor / Advisory BoardOleguer EscolOleguer EscolOleguer EscolOleguer EscolOleguer EscolEullia GarciaEullia GarciaEullia GarciaEullia GarciaEullia GarciaAnna OmedesAnna OmedesAnna OmedesAnna OmedesAnna OmedesJosep PiquJosep PiquJosep PiquJosep PiquJosep PiquFrancesc UribeFrancesc UribeFrancesc UribeFrancesc UribeFrancesc Uribe

    Editors / Editores / EditorsAntonio Barbadilla Antonio Barbadilla Antonio Barbadilla Antonio Barbadilla Antonio Barbadilla Univ. Autnoma de Barcelona, Bellaterra, SpainXavier BellsXavier BellsXavier BellsXavier BellsXavier Bells Centre d' Investigaci i Desenvolupament CSIC, Barcelona, SpainJuan Carranza Juan Carranza Juan Carranza Juan Carranza Juan Carranza Univ. de Extremadura, Cceres, SpainLus M CarrascalLus M CarrascalLus M CarrascalLus M CarrascalLus M Carrascal Museo Nacional de Ciencias Naturales CSIC, Madrid, SpainAdolfo Cordero Adolfo Cordero Adolfo Cordero Adolfo Cordero Adolfo Cordero Univ. de Vigo, Vigo, SpainMario Daz Mario Daz Mario Daz Mario Daz Mario Daz Univ. de CastillaLa Mancha, Toledo, SpainXavier Domingo Xavier Domingo Xavier Domingo Xavier Domingo Xavier Domingo Univ. Pompeu Fabra, Barcelona, SpainFrancisco Palomares Francisco Palomares Francisco Palomares Francisco Palomares Francisco Palomares Estacin Biolgica de Doana, Sevilla, SpainFrancesc Piferrer Francesc Piferrer Francesc Piferrer Francesc Piferrer Francesc Piferrer Inst. de Cincies del Mar CSIC, Barcelona, SpainIgnacio Ribera Ignacio Ribera Ignacio Ribera Ignacio Ribera Ignacio Ribera The Natural History Museum, London, United KingdomAlfredo Salvador Alfredo Salvador Alfredo Salvador Alfredo Salvador Alfredo Salvador Museo Nacional de Ciencias Naturales, Madrid, SpainJos Lus TJos Lus TJos Lus TJos Lus TJos Lus Tellera ellera ellera ellera ellera Univ. Complutense de Madrid, Madrid, SpainFrancesc Uribe Francesc Uribe Francesc Uribe Francesc Uribe Francesc Uribe Museu de Zoologia de Barcelona, Barcelona, Spain

    Consell Editor / Consejo editor / Editorial BoardJos A. BarrientosJos A. BarrientosJos A. BarrientosJos A. BarrientosJos A. Barrientos Univ. Autnoma de Barcelona, Bellaterra, SpainJean C. BeaucournuJean C. BeaucournuJean C. BeaucournuJean C. BeaucournuJean C. Beaucournu Univ. de Rennes, Rennes, FranceDavid M. BirdDavid M. BirdDavid M. BirdDavid M. BirdDavid M. Bird McGill Univ., Qubec, CanadaMats BjrklundMats BjrklundMats BjrklundMats BjrklundMats Bjrklund Uppsala Univ., Uppsala, SwedenJean BouillonJean BouillonJean BouillonJean BouillonJean Bouillon Univ. Libre de Bruxelles, Brussels, BelgiumMiguel Delibes Miguel Delibes Miguel Delibes Miguel Delibes Miguel Delibes Estacin Biolgica de Doana CSIC, Sevilla, SpainDario J. Daz CosnDario J. Daz CosnDario J. Daz CosnDario J. Daz CosnDario J. Daz Cosn Univ. Complutense de Madrid, Madrid, SpainAlain DuboisAlain DuboisAlain DuboisAlain DuboisAlain Dubois Museum national dHistoire naturelle CNRS, Paris, FranceJohn FaJohn FaJohn FaJohn FaJohn Fa Durrell Wildlife Conservation Trust, Trinity, United KingdomMarco FestaBianchetMarco FestaBianchetMarco FestaBianchetMarco FestaBianchetMarco FestaBianchet Univ. de Sherbrooke, Qubec, CanadaRosa FlosRosa FlosRosa FlosRosa FlosRosa Flos Univ. Politcnica de Catalunya, Barcelona, SpainJosep M GiliJosep M GiliJosep M GiliJosep M GiliJosep M Gili Inst. de Cincies del Mar CMIMACSIC, Barcelona, SpainEdmund Gittenberger Edmund Gittenberger Edmund Gittenberger Edmund Gittenberger Edmund Gittenberger Rijksmuseum van Natuurlijke Historie, Leiden, The NetherlandsFernando HiraldoFernando HiraldoFernando HiraldoFernando HiraldoFernando Hiraldo Estacin Biolgica de Doana CSIC, Sevilla, SpainPatrick LavellePatrick LavellePatrick LavellePatrick LavellePatrick Lavelle Inst. Franais de recherche scient. pour le develop. en cooperation, Bondy, FranceSantiago MasComaSantiago MasComaSantiago MasComaSantiago MasComaSantiago MasComa Univ. de Valencia, Valencia, SpainJoaqun MateuJoaqun MateuJoaqun MateuJoaqun MateuJoaqun Mateu Estacin Experimental de Zonas ridas CSIC, Almera, SpainNeil MetcalfeNeil MetcalfeNeil MetcalfeNeil MetcalfeNeil Metcalfe Univ. of Glasgow, Glasgow, United KingdomJacint NadalJacint NadalJacint NadalJacint NadalJacint Nadal Univ. de Barcelona, Barcelona, SpainStewart B. PeckStewart B. PeckStewart B. PeckStewart B. PeckStewart B. Peck Carleton Univ., Ottawa, CanadaEduard PetitpierreEduard PetitpierreEduard PetitpierreEduard PetitpierreEduard Petitpierre Univ. de les Illes Balears, Palma de Mallorca, SpainTTTTTaylor H. Ricketts aylor H. Ricketts aylor H. Ricketts aylor H. Ricketts aylor H. Ricketts Stanford Univ., Stanford, USAJoandomnec RosJoandomnec RosJoandomnec RosJoandomnec RosJoandomnec Ros Univ. de Barcelona, Barcelona, SpainVVVVValentn SansComaalentn SansComaalentn SansComaalentn SansComaalentn SansComa Univ. de Mlaga, Mlaga, SpainTTTTTore Slagsvoldore Slagsvoldore Slagsvoldore Slagsvoldore Slagsvold Univ. of Oslo, Oslo, Norway

    Secretaria de Redacci / Secretara de Redaccin /Editorial Office

    Museu de ZoologiaPasseig Picasso s/n08003 Barcelona, SpainTel. +34933196912Fax +34933104999Email [email protected]

    "La tortue greque" Oeuvres du Comte de Lacpde comprenant L'Histoire Naturelle des Quadrupdes Ovipares, desSerpents, des Poissons et des Ctacs; Nouvelle dition avec planches colories dirige par M. A. G. Desmarest;Bruxelles: Th. Lejeun, diteur des oeuvres de Buffon, 1836. Pl. 7

    Animal Biodiversity and Conservation 24.1, 2001Animal Biodiversity and Conservation 24.1, 2001Animal Biodiversity and Conservation 24.1, 2001Animal Biodiversity and Conservation 24.1, 2001Animal Biodiversity and Conservation 24.1, 2001 2001 Museu de Zoologia, Institut de Cultura, Ajuntament de BarcelonaAutoedici: Montserrat FerrerFotomecnica i impressi: Sociedad Cooperativa Librera GeneralISSN: 1578665XDipsit legal: B16.27858

  • 1Animal Biodiversity and Conservation 24.1 (2001)

    2001 Museu de ZoologiaISSN: 1578665X

    Tasmanitachoides Erwin glabellus n. sp.from North Queensland, Australia,with a note on Tasmanitachoides lutus(Darlington) (Insecta, Coleoptera,Carabidae, Bembidiinae)

    M. Baehr

    Baehr, M., 2001. Tasmanitachoides Erwin glabellus n. sp. from North Queensland, Australia, with a note onTasmanitachoides lutus (Darlington) (Insecta, Coleoptera, Carabidae, Bembidiinae). Animal Biodiversity andConservation, 24.1: 17.

    AbstractAbstractAbstractAbstractAbstractTasmanitachoides Erwin glabellus n. sp. from North Queensland, Australia, with a note on Tasmanitachoideslutus (Darlington) (Insecta, Coleoptera, Carabidae, Bembidiinae). A new species of the genus TasmanitachoidesErwin from North Queensland is described: T. glabellus n. sp. The species belongs to the T. murrumbidgensisgroup of species that is characterized by its distinctly impressed clypeus, but it is distinguished from all relatedspecies by its glabrous body surface. It is the first Tasmanitachoides from northern Australia to be found inrainforest on high mountains and has thus probably preserved the original habits of the genus that are stillcharacteristic for those species living in southern temperate regions of Australia. Tasmanitachoides lutus(Darlington) so far known from the type locality in southern New South Wales and from the holotype only, isnow recorded from eastern Victoria.

    Key words: Tasmanitachoides, T. glabellus n. sp., Australia.

    ResumenResumenResumenResumenResumenTasmanitachoides Erwin glabellus sp. n. del norte de Queensland, Australia, con una mencin sobreTasmanitachoides lutus (Darlington) (Insecta, Coleoptera, Carabidae, Bembidiinae). Se describe una nuevaespecie del gnero Tasmanitachoides Erwin del norte de Queensland: T. glabellus sp. n. Esta especie perteneceal grupo de especies T. murrumbidgensis que se caracteriza por la impresin diferenciada del clipeo, pero quese distingue de todas las especies relacionadas por presentar una superficie corporal glabra. Es el primerTasmanitachoides del norte de Australia encontrado en la selva pluvial de alta montaa y probablementemantiene los hbitos originales de su gnero, que siguen siendo caractersticos de las especies que viven en laszonas templadas del sur de Australia. Tasmanitachoides lutus (Darlington) conocido hasta el momento a partirde la localidad tipo, en el sur de Nueva Gales del Sur, y nicamente por el holotipo, ha sido registrado ahoraen el este del estado de Victoria.

    Palabras clave: Tasmanitachoides, T. glabellus sp. n., Australia.

    (Received: 18 IX 01; Final acceptance: 4 X 01)

    Martin Baehr, Zoologische Staatssammlung, Mnchhausenstr. 21, D81247 Mnchen, Germany.

    e-mail: [email protected]

  • 2 Baehr

    Introduction

    While examining the immense bulk of rainforestcarabid beetles collected during the last decadesby staff at Queensland Museum, Brisbane, theauthor recently detected two specimens of thegenus Tasmanitachoides that he was unable toidentify at once. The specimens were quite unusual,because according to the labels they werecollected near a small creek at the highest top of arainforestcoated mountain in far northernQueensland, presumably even at or near the sourceof this creek. Careful examination and comparisonwith all related species revealed that the specimensbelong to an undescribed species that is of specialinterest due to its habits.

    Methods

    Description and measurements follow the styleused in the authors revision of the genusTasmanitachoides (BAEHR, 1990).

    The types are shared with Queensland Museum,Brisbane (QM) and the authors working collectionin Zoologische Staatssammlung, Munich (CBM).

    Studied material

    Genus Tasmanitachoides Erwin

    Erwin, 1972: 2 (ERWIN, 1972); Moore et al., 1987: 144;(MOORE et al., 1987); Baehr, 1990: 868 (BAEHR, 1990)

    Type speciesBembidion hobarti Blackburn, 1901; by subsequentdesignation.

    This genus of small, elongate, Perileptuslike,sand or gravelinhabiting ground beetles wasfounded by ERWIN (1972) who included those speciesthat were combined by DARLINGTON (1962) to thehobartigroup within the genus Tachys s. l. BAEHR(1990) later included additional species notmentioned by Darlington or Erwin, and describedfurther species. At present, this genus includes 16species which are distributed through the east(including Tasmania) and tropical north of Australiaincluding the Kimberleys in northwestern Australia.A single species (T. arnhemensis Erwin), however,apparently ranges far inland into the west ofWestern Australia and also into central Australia(see BAEHR, 1990: fig. 45).

    The genus combines some archaic bembidiinecharacter states as enumerated in ERWIN (1972)with characters comparable with similar states inthe trechine complex. Erwin regarded thesesimilarities as remnants of an archaic pre-bembidiine stock, but analyses using moleculartechniques seem to indicate that Tasmanitachoidesindeed belongs rather to the trechine than tothe bembidiine stock (Maddison, pers. comm.).

    Within the genus, according to BAEHR (1990),the dark coloured species of the T. hobartisubgroup in its restricted sense (T. hobarti, T.leai, T. wattsense) that occur in southeasternAustralia and Tasmania are most basic phylo-genetically, whereas the lightcoloured, moredelicate species of the T. fitzroyigroup are mostadvanced. If this is true, then the genus originatedsomewhere in temperate (montane) southeasternAustralia and derivative stocks later spread toopen, sometimes even rather dry lowlands ofthe north, west and centre.

    Tasmanitachoides glabellus sp. n. (figs. 1, 2)

    TypesHolotype: }, head of Francis Ck 12km WSWMossman, NQ 30 Dec 1989, 1200m ANZSESExpedition (QMT, 93349).

    Paratype: 1 }, same data (CBM).

    DiagnosisDistinguished by almost glabrous surface of elytrafrom all other species of the T. murrumbidgensisgroup that is characterized by anteriorlyimpressed clypeus. Further distinguished frommost similar T. murrumbidgensis (Sloane) ofsouthern New South Wales by larger size; fromT. fitzroyi (Darlington) of tropical Australia bydark colour of surface and dark 2nd4th

    antennomeres; and from T. maior Baehr ofsoutheastern Victoria by smaller size and slightlymore divergent frontal furrows.

    DescriptionMeasurements. Length: 2.452.50 mm; width:0.95 mm; ratio width/length of pronotum:1.321.33.

    Colour. Dark piceous, anteriorly almost black,only disk of elytra with faint brownish lustre.Antenna and palpi piceous, only 1st antennomerereddish. Legs piceous, tibiae in middle slightlylighter.

    Head. Slightly narrower than pronotum. Surfacenitid, with scattered fine punctures and highlysuperficial isodiametric microreticulation. Labrumanteriorly deeply impressed. Frontal furrows deep,slightly divergent and posteriorly curved outwards.Eyes large, protruding, orbits short. Mandiblesshort. Antenna mediumsized, median anten-nomeres slightly longer than wide.

    Pronotum. Wide, though considerably narrowerthan elytra. Heartshaped, fairly convex, distinctlynarrowed to base. Widest at anterior third, sidesevenly convex, shortly sinuate in front of therectangular basal angles. Base in middle produced,anterior angles slightly projecting. Median lineinconspicuous, no lateral channel developed.Transverse basal sulcus deeply impressed, laterallycoarsely punctate, in middle with a longitudinalfurrow. Disk nitid, sparsely punctate, with highlysuperficial, isodiametric microreticulation.

    Elytra. Rather elongate, widest at about middle,

  • Animal Biodiversity and Conservation 24.1 (2001) 3

    surface depressed. Inner five striae at least in basalhalf deeply impressed, 5th stria near base evensulcate. Sixth and 7th striae barely impressed,becoming very weak towards apex. Third stria atposition of anterior discal pore characteristicallyoutturned and shortly interrupted to meet 4th

    stria. Discal pores almost foveiform. Recurrent striashort. Striae anteriorly rather coarsely punctate.Inner five intervals very gently convex, sparselyand very faintly punctate, with superficial traces ofmicroreticulation only, surface remarkably nitid.Marginal pores reduced to four behind humerus,two in middle, and two near apex within thedeeply impressed submarginal sulcus that formsthe apical part of 7th stria.

    Legs. Anterior tibia barely excised at outer edge.Aedeagus. Unknown.Female stylomeres (fig. 1). Both stylomeres very

    slender and elongate. Stylomere 1 dorsoventrallycurved, without any setae at apical margin. Stylomere2 almost straight, with a very elongate and a shorternematiform seta right on apex, and one, respectivelyone or two shorter nematiform setae at internal andexternal margins close to apex.

    Variation. Little variation noted.

    DistributionFar northeastern Queensland. Known only fromtype locality.

    BiologyVery little known. According to the label, bothspecimens were collected at the top of a mountainat the height of 1,200 m, most probably in montanerainforest, at the edge of a creek and perhapseven at or very close to the source of this creek.Probably the habits of this species are similar tothose living in montane regions of temperatesouthern Australia (southern New South Wales,eastern Victoria, Tasmania) where species ofTasmanitachoides likewise occur in sand or gravelof small banks at mountain creeks and small rivers.

    EtymologyThe name refers to the rather glabrous elytralsurface as compared with that of similar species.

    RecognitionThe determination key in the authors revisionof Tasmanitachoides (BAEHR, 1990, p. 869870)has been fully revised.

    Fig. 1. Tasmanitachoides glabellus n. sp.: A. Habitus, length 2.5 mm; B. } left stylomeres 1 & 2,ventral view.

    Fig. 1. Tasmanitachoides glabellus sp. n.: Habitus, longitud 2,5 mm; B. } estilmeros 1 y 2 izquierdos,visin ventral.

    AAAAA B B B B B

    0.1

    mm

    0.1

    mm

    0.1

    mm

    0.1

    mm

    0.1

    mm

  • 4 Baehr

    Revised key to the genus Tasmanitachoides.

    Clave revisada del gnero Tasmanitachoides

    1 Elytral striae, except sutural, reduced. SouthernNew South Wales, eastern Victoria T. lutus (Darlington)Elytra with at least 5th stria marked, otherssometimes superficial 2

    2 Clypeus distinctly impressed at middle (doubtfulspecies under both couplets) 3Clypeus not impressed at middle 11

    3 Larger and wider species, c. 2.5 mm long or more(doubtful species under both couplets) 4Smaller and narrower species, < 2.3 mm long 6

    4 Either reasonably smaller species (< 2.5 mm long)with elytra almost lacking microreticulation; or anteriorbody rufoustestaceous, elytra at apex testaceous;antennae and palpi yellow; anterior angles of pronotumproduced. Northern Queensland, northwestern Australia 5Larger species, 2.9 mm long; colour uniformly piceous;antennae and palpi dark; anterior angles of pronotumnot produced. Eastern Victoria T. maior Baehr

    5 Dark species with dark antennae and palpi; elytraalmost lacking microreticulation; frontal furrows littledivergent. Northeastern Queensland, near rain forestbordered creek on high mountain T. glabellus n. sp.Light species with rufoustestaceous fore body, elytraat apex testaceous, antennae and palpi yellow; frontalfurrows distinctly divergent. Northern Queensland,northwestern Australia, near rivers and creeks in opento sparsely forested lowland T. fitzroyi (Darlington)

    6 Fore body reddish to reddishtestaceous, elytra testaceous.Northern Territory and northwestern Australia 7Either completely piceous or dark reddish, or forebody dark piceous and elytra dark reddish withpiceous borders, suture, base, and apex. Eastern Australiafrom north Queensland to southern New South Wales 8

    7 Larger and wider species, 1.92.15 mm long; frontalfurrows posteriorly slightly divergent; border ofpronotum convex throughout to the small, projectingbasal angle; 2nd4th elytral striae less impressed. Centraland far Northern Territory, northwestern Australia T. arnhemensis ErwinSmaller and narrower species, 1.651.95 mm long;frontal furrows parallel; border of pronotum distinctlysinuate to the right, but nonprojecting basal angle;2nd4th elytral striae more impressed. NorthwesternAustralia north of Great Sandy Desert T. minor Baehr

    8 Clypeus anteriorly deeply impressed; only 1st antennomerereddish, others piceous; colour dark piceous withelytra at most feebly lighter on disk 9Clypeus anteriorly lightly impressed, impression sometimesdifficult to see; 1st4th antennomeres reddish, otherspiceous; colour either completely dark reddish, or forebody dark piceous with contrastingly lighter elytra 10

  • Animal Biodiversity and Conservation 24.1 (2001) 5

    9 Smaller species, < 2.2 mm long; elytra markedlymicroreticulate. Southern New South Wales T. murrumbidgensis (Sloane)Larger species, > 2.4 mm long; elytra barely microreticulate.Far northeastern Queensland T. glabellus sp. n.

    10 Smaller species, 1.752.1 mm long; eyes large,protruding, orbits almost wanting; fore body piceous,elytra lighter on disk, colour of body and antennaerather contrasting; pronotum less narrowed to base,dorsally more convex. Northeastern Queensland T. bicolor BaehrLarger species, 2.02.3 mm long; eyes smaller, lessprotruding, orbits perceptible, oblique; completelydark reddish to light piceous, elytra, at most, slightlylighter, colour of body and antennae not muchcontrasting; pronotum rather narrowed to base,dorsally more depressed. Eastern New South Wales,Australian Capital Territory . T. rufescens Baehr

    11 Elytra parallel, depressed; eyes small, depressed,with well developed orbits; posterior supraorbitalseta situated far behind eye; mandibles veryelongate, decussate; pronotum trapezoid, widestslightly behind anterior angles; colour testaceous.Northeastern New South Wales, eastern Queensland T. obliquiceps (Sloane)Elytra less parallel and depressed; eyes larger,more protruding, orbits small; posterior supraorbitalseta situated immediately at posterior border of eye;mandibles shorter, not decussate; pronotum laterallymore convex, widest far behind anterior angles;colour reddishtestaceous to black 12

    12 Only 1st and 5th elytral striae well impressed, othersbarely recognizable; elytra nitid 13All elytral striae present, though 2nd4th sometimessuperficial; elytra distinctly microreticulate 15

    13 Short, convex species; elytra considerably less than 1.5xas long as wide; frontal furrows short; pronotum wide,base (at basal angles) as wide as apex, basal anglesover 90, not projecting. Southern Queensland T. wilsoni (Sloane)More elongate, less convex species; elytra more than1.5x as long as wide; frontal furrows elongate,conspicuous; pronotum narrower, base (at basal angles)considerably narrower that apex, basal angles acute,laterally projecting 14

    14 Smaller species, less than 1.7 mm long; eyes lessprotruding, orbits perceptible; basal angles ofpronotum c. 90, less acute and projecting. Tasmania T. kingi (Darlington)Larger species, 1.82.0 mm long; eyes more protruding,orbits almost reduced, basal angles of pronotumacute, less than 90, laterally distinctly projecting.Eastern Victoria and New South Wales T. angulicollis Baehr

    15 Elongate, depressed, very small species, 1.51.7 mm long;colour testaceous to light reddish. Far Northern Territory,northwestern Australia, northeastern Queensland,northeastern New South Wales T. katherinei ErwinMore convex, larger species, 1.72.6 mm long; colourdark reddish to black 16

  • 6 Baehr

    16 On the average smaller species, 1.72.3 mm long,rather depressed; clypeus faintly impressed; coloureither rather uniformly dark reddish, or piceouswith disk of each elytron contrastingly lighter 17On the average larger species, 2.12.6 mm long,more convex; clypeus not at all impressed; colouruniformly dark piceous to black, or piceous withelytra slightly (not contrastingly) lighter 18

    17 Smaller species, 1.752.1 mm long; eyes large,protruding, orbits almost wanting; fore body piceous;elytra lighter on disk, colour of body and antennaerather contrasting; pronotum less narrowed to base,dorsally more convex. Northeastern Queensland T. bicolor BaehrLarger species, 2.02.3 mm long; eyes smaller, lessprotruding, orbits perceptible, oblique; completelydark reddish to light piceous, elytra, at most, slightlylighter, colour of body and antennae not muchcontrasting; pronotum rather narrowed to base,dorsally more depressed. Eastern New South Wales,Australian Capital Territory T. rufescens Baehr

    18 Elytral striae, including 5th, strongly impressed 19Elytral striae, especially 5th, rather superficial.Eastern Victoria, southern New South Wales T. wattsense (Blackburn)

    19 Colour uniformly dark piceous to almost black;antennae completely dark. Tasmania T. hobarti (Blackburn)Colour piceous, disk of elytra slightly lighter;basal antennomeres reddish. Northeastern NewSouth Wales T. leai (Sloane)

    RemarksWith respect to the distinctly impressed clypeus, T.glabellus clearly belongs to the T. murrumbidgensisgroup within the genus Tasmanitachoides. Thecombination of its dark colour and almost glabroussurface, however, at once distinguishes this speciesfrom all known species. Moreover, the dark colouris also unique within all Tasmanitachoides knownso far to occur in northern tropical Australia. Allthose species are either completely reddish ortestaceous (arnhemensis, fitzroyi, katherinei, minor,obliquiceps), or are at least bicolourous with darkforebody though lighter elytra (bicolor). In contrast,almost all of the southern species are completelydark.

    According to ERWINs (1972), DARLINGTONs (1962:gravel by brooks), and the authors observations,in temperate Australia Tasmanitachoides arecommonly found at small rivers and mountainbrooks, even in shaded places, and commonly alsoat high altitudes. The species living in tropicalAustralia, however, are generally found in gravelsand sands of lakes, rivers and creeks of openlowlands, commonly even in comparatively arid

    regions. Here, while exposed to the bright sun,their reddish or testaceous colour corresponds wellwith the colour of the substratum they live on andin namely light coloured, at most light reddishgravels and sands.

    It has been postulated that the habits near streamsin temperate montane regions is regarded theoriginal mode of life for the genus Tasmanitachoides,whereas their occurrence in the tropical regions ofnorthern Australia is secondary (BAEHR, 1990). If thisassumption is true, then the occurrence of a darkcoloured species living at shaded rainforest creeksin montane northern Queensland would be quitesurprising, because this would mean a relictoccurrence with an ancient mode of life far northof the roots of this ancient genus that most probablyoriginated somewhere in temperate southeasternAustralia.

    This assumption seems rather unlikely at firstglance, though within recent years a number ofexamples of definitely southern groups weredetected that have members far north in thetropics and subtropics well outside of theirrecognized range. Carabid examples for this

  • Animal Biodiversity and Conservation 24.1 (2001) 7

    distribution pattern are two merizodine species ofthe genus Sloaneana Csiki which occur onLamington Plateau of southeastern Queensland(BAEHR, in press), or the occurrence and remakabletaxonomic radiation of the psydrine generaRaphetis Moore, Sitaphe Moore, and of amblytelinePsydrinae in the wet tropics of North Queensland(unpublished records), or even the discovery of apeculiar (yet undescribed) new genus of thedefinitely antarctic subfamily Migadopinae,likewise in tropical North Queensland.

    It follows from these examples, which couldbe complemented by certain noncarabidexamples, that remnants of the southerntemperate Antarctic faunal element ofAustralia are still present even in tropicalnorthern Queensland, and furthermore that thisdistribution pattern is probably more commonthan was believed to date. If related to thegeographic history of Australia, these examplesdemonstrate that various elements of thesouthern fauna were somehow trapped onmountains and tablelands of eastern and northeastern Queensland during Australias drift tothe north during the Tertiary period. As a result,they can now be found high up in environmentswhich although allowing them to survivethere prevent their contact with their southerncounterparts and also prevent any furtherspreading.

    When seen in the light of the biogeographicalhistory of northeastern Australia, the unexpecteddiscovery of the new Tasmanitachoides addsvaluable information towards understanding thecomplexity of the montane fauna of the wettropics of northern Queensland.

    Tasmanitachoides lutus (Darlington)

    Tachys lutus Darlington, 1962: 120 (DARLINGTON, 1962)Tasmanitachoides lutus, Erwin 1972: 5 (ERWIN, 1972);

    Moore et al. 1987: 145 (MOORE et al., 1987); Baehr1990: 877 (BAEHR, 1990)

    This remarkably and easily recognized speciesthat lacks all but the sutural, elytral striae was

    only known to date from the holotype collectedat Termeil, near the coast of southeastern NewSouth Wales. During ecological studies on ripariangravel bank arthropods carried out by V.Framenau on rivers in eastern Victoria (FRAMENAU,et al., in press) this species has been now recordedfrom Cann River and Castleburn Creek, both insoutheastern Victoria. At both localities, a singlespecimen each was found on gravel banks withinclosed forest.

    New recordsVIC: Cann River at Chandlers Ck Bridge, 37.20 S,149.12 E, 8 XII 1998; Castleburn Ck, Junctionwith Mitchell River, 37.31 S, 147.12 E, 26 XI 1998.

    References

    BAEHR, M., 1990. Revision of the Australiangroundbeetle Genus Tasmanitachoides Erwin(Insecta: Coleoptera: Carabidae: Bembidiinae),with special regard to the tropical species.Invertebr. Taxon., 4: 867894.

    (in press). Two new species of Sloaneana Csikifrom southern Queensland (Coleoptera,Carabidae, Merizodinae). Mem. QueenslandMus.

    DARLINGTON, P. J. JR, 1962, Australian Carabidbeetles XI. Some Tachys. Psyche, Cambridge69: 117128.

    ERWIN, T. L., 1972. Two new genera of BembidiineCarabid beetles from Australia and SouthAmerica with notes on their phylogenetic andzoogeographical significance (Coleoptera).Breviora, 383: 119.

    FRAMENAU, V., MANDEBACH, R., & BAEHR, M. (inpress). Riparian gravel banks of upland andlowland rivers in Victoria (South East Australia):Arthropod community structure and life historypatterns in a longitudinal gradient. Aust. J.Zoology.

    MOORE, B. P., WEIR, T. A. & PYKE, J. E., 1987.Rhysodidae and Carabidae. In: ZoologicalCatalogue of Australia, 4: 17320. AustralianGovernment Publishing Service, Canberra.

  • 9Animal Biodiversity and Conservation 24.1 (2001)

    2001 Museu de ZoologiaISSN: 1578665X

    Bells, X., 2001. Description of Sphaericus selvagensis n. sp. from the Salvage Islands, and new data onSphaericus bicolor Bells (Coleoptera, Ptinidae). Animal Biodiversity and Conservation, 24.1: 913.

    AbstractAbstractAbstractAbstractAbstractDescription of Sphaericus selvagensis n. sp. from the Salvage Islands, and new data on Sphaericus bicolor Bells(Coleoptera, Ptinidae). Sphaericus (Sphaericus) selvagensis n. sp. is described from the Salvage islands. WithSphaericus (Sphaericus) bicolor Bells, this new species is only the second ptinid beetle reported from theseislands. S. selvagensis belongs to the Sphaericus pilula group, which also includes S. bicolor. However, thetransverse shape of the pronotum (with its maximal breadth near the base) and the peculiar morphology of theaedeagus, distinguish S. selvagensis from all other members of the S. pilula group. S. selvagensis lives in all themajor islands of the Selvagens archipelago: Selvagem Grande, Selvagem Pequena and Ilhu de Fora.

    Key words: Coleoptera, Ptinidae, Sphaericus, Salvage Islands.

    ResumenResumenResumenResumenResumenDescripcin de Sphaericus selvagensis sp. n. del archipilago de las Salvajes, y nuevos datos sobre Sphaericusbicolor Bells (Coleoptera, Ptinidae). Se describe Sphaericus (Sphaericus) selvagensis sp. n. del archipilago delas Salvajes. Junto a Sphaericus (Sphaericus) bicolor Bells, esta nueva especie es el segundo coleptero ptnidoregistrado en esas islas. S. selvagensis pertenece al grupo de Sphaericus pilula, que tambin incluye S. bicolor,aunque la forma transversa del pronoto (con anchura mxima cerca de la base) y la peculiar morfologa deledeago distinguen a S. selvagensis de los restantes miembros de grupo de S. pilula. S. selvagensis vive en todaslas islas principales del archipilago de las Salvajes: Salvaje Grande, Salvaje Pequea (o Pitn Grande) y LaSalvajita (Ilhu de Fora).

    Palabras clave: Coleoptera, Ptinidae, Sphaericus, Islas Salvajes.

    (Received: 1 X 01; Conditional acceptance: 10 X 01; Final acceptance: 20 X 01)

    Xavier Bells, Dept. of Physiology and Molecular Biodiversity, Inst. de Biologia Molecular de Barcelona (CID, CSIC),c/ Jordi Girona 18, 08034 Barcelona, Espanya (Spain).

    e-mail: [email protected]

    Description of Sphaericus selvagensis n. sp.from the Salvage Islands, and new data onSphaericus bicolor Bells(Coleoptera, Ptinidae)

    X. Bells

  • 10 Bells

    Introduction

    The Salvage Islands lie in the Atlantic Oceanbetween the wellknown archipelagos ofMadeira and Canaries (BRAVO & COELLO, 1978).Up to now, the only ptinid beetle reportedfrom the Selvagens is Sphaericus (Sphaericus)bicolor Bells, described from Selvagem Pequena(= Pito Island) (BELLS, 1982) and later recordedby ERBER & WHEATER (1987) from SelvagemGrande and Ilhu de Fora. However, the studyof the ptinid beetles collected during a campaigncarried out in the Salvages in May 1999, in thecontext of the Project Macaronesia 2000 ofthe Museo de Ciencias Naturales de Tenerife,has lead to the discovery of a new species ofSphaericus, which is described in the presentpaper. The data on the arthropods collectedduring this expedition of 1999 have beenreported by ARECHAVALETA et al. (2001).

    The genus Sphaericus was proposed byWollaston as early as 1854, but has been thesubject of a relatively recent synopsis by BELLS(1994), who divided it into three subgenera:Sphaericus, the members of which are charac-terized by having 11segmented antennae,5segmented male metatarsi, the base ofthe pronotum simple and the parameres ofthe aedeagus slender and pubescent only atthe apex; Nitpus Jacquelin du Val, whosetwo species have 9segmented antennae and4segmented male metatarsi; and DoramasusBells, described in the same synopsis (BELLS,1994) as similar to Sphaericus s. str. but showingthe base of the pronotum protuberant and theparameres of the aedeagus robust and evenlypubescent.

    With the exception of Sphaericus (Sphaericus)gibboides (Boieldieu), which is anthropophilousand nearly cosmopolitan (HINTON, 1941), andSphaericus (Sphaericus) niveus (Boieldieu),Sphaericus (Sphaericus) exiguus (Boieldieu) andSphaericus (Nitpus) ptinoides (Boieldieu), whichare known from sparse localities in theMediterranean area (BOIELDIEU, 1856; PIC, 1912;BELLS, 1994), all the other species of thesethree subgenera are endemic to islands ofAtlantic archipelagos.

    The island groups include the Canaries(10 species), Madeira (nine species), Cape Verde(two species), Salvages (two species, includingthat described herein), and Aores (one species)(BELLS, 1994). More recently, the new subgenusLeasphaericus Bells (1998) (BELLS, 1998) hasbeen proposed for two Australian species. Thesetaxa, in contrast with the Palaearctic Sphaericus,have a triangular scutellum easily visible fromabove.

    Due to the morphology of the aedeagus andthe pronotum, the number of the segments inthe antennae and tarsi, and the hidden scutellum,the new species described below falls into thesubgenus Sphaericus Wollaston.

    Description

    Sphaericus (Sphaericus) selvagensis n. sp.

    TypesHolotype: 1{ labelled Islas Salvajes, SelvagemGrande, 21/26V1999, M. Arechavaleta leg.(Museo de Ciencias Naturales, Santa Cruz deTenerife).

    Paratypes: 84 specimens of both sexes with thesame label as the holotype; 6 specimens of bothsexes with the label Islas Salvajes, SelvagemPequena, 25V1999, M. Arechavaleta leg.; 1}labelled Selvagem Pequena, Pico Veado, 21870,Maul leg.; 18 specimens of both sexes with thelabel I. Selvagens, Pito, 5VI1970, Maul leg.; 1}labelled Islas Salvajes, Ilhu de Fora, 25V1999,M. Arechavaleta leg. (Museo de Ciencias Naturales,Santa Cruz de Tenerife; Departamento de BiologaAnimal, Universidad de La Laguna; Museo Nacionalde Ciencias Naturales, Madrid; Museu de Zoologa,Barcelona; colls. Orom, Bells, Arechavaleta andGarca Becerra).

    Description of the male (fig. 1)Length: 1.21.8 mm (n = 12)

    Broadly oval; pronotum black, elytra darkbrownishred, appendages and sternal part ofbody testaceous. Head clothed with short,recumbent, golden hairs; eyes moderately convex,round, about twothirds as broad as first antennalsegment; space between antennal fossae narrowand acute; antennae 11segmented, short (abouthalf as long as body) and robust, segments 3 to10 oval, slightly longer than width. Pronotumevenly convex, transverse, sides feebly arcuate,maximal breadth near base; surface with verydense to contiguous granules each about twiceas large as eye facets; covered with flat, obovate,recumbent yellowish scales (easily abraded), eachabout same size as granules, also sparse, shortand suberect dark rufopiceous hairs; legsrelatively short, robust; tarsi 5segmented. Elytrabroadly oval, sides regularly rounded, maximalbreadth at middle; surface shiny, with moderatelydense, irregularly distributed punctures slightlynarrower than pronotal granules; covered withscales like those of pronotum (also easilyabraded), but ellipsoidal in shape, also withsparse, short and recumbent yellowish irregularlydistributed hairs. Scutellum much reduced,hidden from above. Aedeagus (figs. 23)symmetrical, with the median lobe slender, indorsal view slightly shorter than parameres;parameres elongate, slightly broader than medianlobe, with sparse, short and erect setae at apex.

    Description of the femaleThe female is externally similar to the male.

    Distribution and habitatS. selvagensis is known from the three majorislands of the Salvage Islands: Selvagem Grande,

  • Animal Biodiversity and Conservation 24.1 (2001) 11

    Selvagem Pequena (= Pito Island ) and Ilhude Fora. Specimens from the campaign of May1999 were collected with pitfall traps andsifting leafmould from different plant species.Moreover, the label of the female collected byMaul at Pico Veado, in Selvagem Pequena, inAugust 1970, indicates that it was collectedfrom sifted dry foliage and leafmould underBassia tomentosa.

    In the Salvages, B. tomentosa (Lowe)(Chenopodiaceae) is a relatively rare speciesobserved in the Selvagem Pequena, and localizedonly in two spots, one on the West slope of thePico Veado and the other one in the Eastern partof the island (PREZ DE PAZ & ACEBES, 1978).

    Comparative notesWithin the subgenus Sphaericus, the generalshape and the typical scaliform pubescence ofS. selvagensis reminds one of the speciesbelonging to the Sphaericus gibboides andSphaericus dawsoni groups (sensu BELLS, 1994).However, the narrow interantennal spacedistinguishes S. selvagensis from the species ofthese groups. Moreover, the pronotum evenlyconvex and the morphology of the aedeagus,especially that of the basis of the median lobe,

    easily separates S. selvagensis from the speciesof the Sphaericus gibbicollis group (sensu BELLS,1994). The new species appears to belong tothe Sphaericus pilula group (sensu BELLS, 1994),which includes S. bicolor, previously known fromthe Salvage Islands. The members of this grouphave the interantennal space narrow, thepronotum evenly convex and the elytra irregularlypunctated. In the case of S. selvagensis, thetransverse shape of the pronotum (with itsmaximal breadth near the base) and the peculiarmorphology of the aedeagus, distinguish it fromall other members of the S. pilula group. Thedifferences are well apparent between S.selvagensis and S. bicolor, as shown by the keyshown below.

    New data on Sphaericus bicolor Bells, 1982

    Up to now, S. bicolor was the only known ptinidspecies from the Salvage Islands. It was describedfrom Selvagem Pequena (= Pito Island), on thebasis of abundant material collected in February1976 by P. Orom (BELLS, 1982). The specimenswere found in leaf mould under Suaeda veraGmelin (Chenopodiaceae) (OROM et al., 1978),

    Figs. 13. Sphaericus (Sphaericus) selvagensis n. sp., a typical specimen from Selvagem Grande(21/26 V 1999, M. Arechavaleta leg.): 1. Habitus; 2. Aedeaguss, dorsal view; 3. Aedeagus, lateralview.

    Figs. 13. Sphaericus (Sphaericus) selvagensis sp. n., un ejemplar tpico de isla Salvaje Grande(21/26 V 1999, M. Arechavaleta leg.): 1. Habitus; 2. Edeago, vista dorsal; 3. Edeago, vista lateral.

    11111 22222 33333

  • 12 Bells

    which is one of the most abundant and typicalplants of the Salvages, either in the SelvagemGrande or in the Selvagem Pequena (PREZ DE PAZ& ACEBES, 1978). Interestingly, no specimens of S.selvagensis were collected during this 1976campaign. Almost simultaneously, SERRANO (1983)recorded an undetermined species of Sphaericusfrom the Selvagem Grande (1 specimen) andSelvagem Pequena (549 specimens). Seventytwospecimens were examined by the author fromthis large series and all were S. bicolor. Thespecimen from Selvagem Grande was collectedon S. vera and those from Selvagem Pequena onElytrigia junceiforme A. et D. Lve (Poaceae)(SERRANO, 1983). A. junceiforme is relatively rarein Selvagem Pequena, being found in a singlelocality on the Eastern part of the island. Morerecently, ERBER & WHEATER (1987) have reportedthe identification of 89 specimens of S. bicolorfrom Selvagem Pequena, 4 from Ilhu de Foraand 1 from Selvagem Grande, which had beencollected by Backhuys in 1968 and deposited inthe Museum of Funchal. Materials from theexpedition in 1999 studied in the present workincluded specimens of S. bicolor mixed with thenew S. selvagensis, and was collected using pitfalltraps and sifting leaf mould from different plants.The number of specimens of both species presentin these and in other samples studied by theauthor is indicated in table 1. These data suggestthat any of the two species may be very abundant

    Key to the Sphaericus of the Salvage Islands.

    Clave para los Sphaericus de las Islas Salvajes.

    1 Antennae long and slender, clearly longer thanhalf the body, with the segments 210 subcylindrical,nearly longer than width. Pronotum longer thanwidth, with the maximal breadth near the middle.Legs long and slender. Elytra ellipsoidal (fig. 1from BELLS, 1982). Aedeagus in dorsal view withthe median lobe much shorter than the parameres;parameres clearly broader than the median lobe(figs. 34 from BELLS, 1982) S. bicolor Bells, 1982Antennae short and robust, about half as long asthe body, with the segments 210 oval, slightly longerthan width. Pronotum transverse, with the maximalbreadth near the base. Legs short and robust.Elytra broadly oval (fig. 1, present paper).Aedeagus in dorsal view with the median lobealmost as long as the parameres; parameresslightly broader than the median lobe(figs. 23, present paper) S. selvagensis n. sp.

    Table 1. Number of specimens of Sphaericusbicolor and Sphaericus selvagensis collectedin the Salvage Islands and studied by theauthor: M. Maul, 5 VI 1970; O. Orom, 26/29II 1976; S. Serrano, 20 IV15 V 1980; A.Arechavaleta 21/26 V 1999; SP. SelvagemPequena; SG. Selvagem Grande. (* From atotal sample of 549 specimens identified bySERRANO, 1983 as Sphaericus sp., 72 werestudied by the author and identified as a S.bicolor.)

    Tabla 1. Nmero de ejemplares de Sphaericusbicolor y Sphaericus selvagensis recogidosen las Islas Salvajes, estudiados por el autor:M. Maul, 5 VI 1970; O. Orom, 26/29 II 1976;S. Serrano, 20 IV15 V 1980; A. Arechavaleta,21/26 V 1999; SP. Salvaje Pequea; SG. SalvajeGrande. (* De un total de 549 ejemplaresidentificados por SERRANO, 1983 comoSphaericus sp., 72 fueron estudiados por elautor e identificados como S. bicolor.)

    SP SG

    M O S A A

    S. bicolor 13 75 72(549)* 6 6

    S. selvagensis 18 0 0 6 91

  • Animal Biodiversity and Conservation 24.1 (2001) 13

    depending on the time and eventually on theprecise site of collection. All data (BELLS, 1982;ERBER & WHEATER, 1987; present results) indicatethat both S. bicolor and S. selvagensis arewidespread in the three main islands of thearchipelago: Selvagem Grande, SelvagemPequena (= Pito Island) and Ilhu de Fora.

    Acknowledgements

    Thanks are due to Pedro Orom for critical readingof the manuscript and for sending abundantmaterial of Sphaericus from the Salvages,especially those collected by M. Arechavaletaduring the expedition of May 1999, in the contextof the Project Macaronesia 2000 of the Museode Ciencias Naturales de Tenerife. Keith Philipsalso reviewed the manuscript. Artur R. M. Serranosent a large sample of S. bicolor from SelvagemPequena collected during the ExpedioZoolgica aos Arquiplagos da Madeira e dasSelvagens (30 de Abril15 de Maio, 1980).

    References

    ARECHAVALETA, M., ZURITA, N. & OROM, P., 2001.Nuevos datos sobre la fauna de artrpodos delas Islas Salvajes. Rev. Acad. Canar. Cienc., 12(34): 8399 (2000).

    BELLS, X., 1982. El primer representante de lafamilia Ptinidae (Col.) de las Islas Salvajes:Sphaericus bicolor n. sp. Vieraea, 11: 103108.

    1994. El gnero Sphaericus Wollaston, 1854(Coleoptera: Ptinidae). Boln. Asoc. esp. Ent.,18: 6179.

    1998. A new subgenus and two new species ofSphaericus Wollaston (Coloptera, Ptinidae)from western Australia. Eur. J. Entomol., 95:263268.

    BOIELDIEU, A., 1856. Monographie des Ptiniores. Annls.Soc. ent. Fr., (3)4: 285315, 487504, 629686.

    BRAVO, T. & COELLO, J., 1978. Descripcingeogrfica del Archipilago de las Salvajes. In:Contribucin al estudio de la historia naturalde las Islas Salvajes: 914. Aula de Cultura deTenerife, Santa Cruz de Tenerife.

    ERBER, D. & WHEATER, C. F., 1987. The Coleopteraof the Selvagem Islands, including a catalogueof the pecimens in the Museu Municipal doFunchal. Bol. Mus. Mun. Funchal, 39(193):156187.

    HINTON, H. E., 1941. The Ptinidae of economicimportance. Bull. ent. Res., 31: 331381.

    OROM, P., BAEZ, M. & MACHADO, A., 1978.Contribucin al estudio de los artrpodos de lasIslas Salvajes. In: Contribucin al estudio de lahistoria natural de las Islas Salvajes: 178194.Aula de Cultura de Tenerife, Santa Cruz deTenerife.

    PREZ DE PAZ, P. L. & ACEBES, J. R., 1978. Las IslasSalvajes: Contribucin al conocimiento de suflora y vegetacin. In: Contribucin al estudiode la historia natural de las Islas Salvajes: 79104. Aula de Cultura de Tenerife, Santa Cruz deTenerife.

    PIC, M., 1912. Ptinidae. In: ColeopterorumCatalogus, 41: 146 (W. Junk & S. Schenkling,Eds.). W. Junk, Berlin.

    SERRANO, A. R. M., 1983. Os coleopteros doArquiplago das Selvagens. In: Act. I Congr. IbricoEnt., 2: 759776. Servicio de Publicaciones de laUniversidad de Len, Len.

  • 15Animal Biodiversity and Conservation 24.1 (2001)

    2001 Museu de ZoologiaISSN: 1578665X

    Survival of a small translocatedProcolobus kirkii population onPemba Island

    A. Camperio Ciani, L. Palentini & E. Finotto

    Camperio Ciani, A., Palentini, L. & Finotto, E., 2001. Survival of a small translocated Procolobus kirkii populationon Pemba Island. Animal Biodiversity and Conservation, 24.1: 1518 .

    AbstractAbstractAbstractAbstractAbstractSurvival of a small translocated Procolobus kirkii population on Pemba Island. A survey to evaluate thedistribution of Procolobus kirkii on Pemba island (Tanzania) was conducted, 20 years after they had beentranslocated from Zanzibar in the Ngezi forest park. A team of both expert and trained observers, guided bythe authors, censused 68.3 linear km of forest, corresponding to an estimated area of 3.5 km2 (63.6%) of theprotected Ngezi forested area of 5.5 km2. Nineteen groups of Cercopithecus aethiops were observed, with atotal of 166 animals and an estimated density of 47.43 individuals per km2, and only one troop of Procolobuskirkii. Supplemented by interviewing the local people we obtained an estimate of 1530 P. kirkii, including asmall troop outside the protected area. This small population survived but did not increase, possibly due toadverse relations with humans.

    Key word: Procolobus kirkii, Translocated population, Density, Conservation, Pemba Island.

    ResumenResumenResumenResumenResumenSupervivencia de una pequea poblacin trasladada de Procolobus kirkii en la isla de Pemba. Se realiz unestudio para evaluar la distribucin de Procolobus kirkii en la isla de Pemba (Tanzania), veinte aos despus deque fuera trasladada desde Zanzbar al Parque Ngezi. Un equipo de observadores expertos y entrenados,guiados por los autores, efectu un censo a lo largo de 68,3 km lineales de bosque, correspondiente a un reaestimada de 3,5 km2 (63,6%) del rea protegida del bosque de Ngezi de 5,5 km2. Se observaron 19 grupos deCercopithecus aethiops, con un total de 166 animales y una densidad estimada de 47,43 individuos/km2, y sloun grupo de Procolobus kirkii. Complementando los datos con entrevistas a la poblacin local se obtuvo unaestimacin de 1530 ejemplares de P. kirkii, incluyendo un pequeo grupo localizado fuera del rea protegida.Este pequeo grupo sobreviva pero no se incrementaba en nmero, posiblemente debido a las relacionesadversas con los humanos.

    Palabras clave: Procolobus kirkii, Poblacin trasladada, Densidad, Conservacin, Isla de Pemba.

    (Received: 23 VII 01; Final acceptance: 2 X 01)

    Andrea Camperio Ciani(1), Loris Palentini & Enrica Finotto, Dip. di Psicologia Generale, Universitadegli Studi diPadova, 8 via Venezia, 35139 Padova, Italy.

    (1)e-mail: [email protected]

  • 16 Camperio Ciani et al.

    Introduction

    Procolobus kirkii, member of the Colobinae family,represents one of Africas most endangeredprimate species.

    It is mainly an arboreal and folivorous species,sympatric but not in competition with Cercopitecusaethiops which is mainly frugivorous (SIEX &STRUHSAKER, 1999a). It has been reported that tocontrast the toxins contained in certain fruit P.kirkii eats a small quantity of charcoal which allowa slower, but otherwise impossible, digestion(STRUHSAKER et al., 1997). Its ideal habitats inZanzibar are areas with ground water, swampforest, scrub forest or mangrove swamp.

    Troops are numerous and can include morethan 80 individuals. They have a multimalestructure which is unusual for the Colobinaefamily, with a 1:2 sex ratio with adult females.Fecundity is about 1.5 newborn every 2 yearsand infant care is intense and shared by severalrelated females. Infanticide is common as inmost Colobinae when a new male joins thegroup.

    P. kirkii is endemic and confined to the islandof Zanzibar. It is present in 3 different forestswith a total population of about 1,500 individuals(Zanzibar Unpublished Government Census,1981). Two decades ago specimens were movedto new areas, mostly small islands, in order totry to inhibit their decline leading to a rapidextinction. These animals are threatened bymassive deforestation and furthermore arehunted for their meat and for pet markets(STRUHSAKER & SIEX, 1996).

    An assessment of the present survival rate andthe diffusion of the small Procolobus population(14 individuals) translocated from Jozani Park andintroduced in the region of Ngezi Forest, in 1974(STRUHSAKER & SIEX, 1998) in the north of PembaIsland is reported in this study.

    Methods

    Data were collected from 15th20th October2000. To census the region as thoroughly aspossible the forest was divided into 14 transects(fig. 1) varying in length from about 2 to 8 km(totally 68.3 km). Each transect segment wasidentified by a 1:50,000 topographical map, andlocated in the field with a GPS and compass.

    Teams included volunteers who underwentprior training in the Jozani Forest of Zanzibar toidentify the different species of monkeys untilconsensus with the trainers reached completeagreement.

    Transects were walked with a fixed departure,arrival and direction. Each transect was walkedby a rotating team of 3 to 4 people, scaled inexperience in the field, and randomly changedeach day in order to avoid individual bias in datacollection (CAMPERIO CIANI et al., 2001).

    Forest quality was classified into five mainhabitat types: gallery forest, mangrove, savannah,swamp and cultivations. To estimate the densityof monkeys in each different habitat wecalculated the width of our transects in eachhabitat. To assess the width, as for the case oftransects of indefinite width (CAUGHLEY, 1977),we used the average distance at first sighting ofthe Cercopithecus aethiops in that habitat.

    Field survey was supplemented with interviewsamong the local people living in villages aroundand within the Ngezi Forest in search of witnessesand information about the presence of P. kirkii.

    Results and discussion

    A distance of approximately 68.3 km was walkedin the five various habitats inside the park.Considering the length and the relative width ofour transects, during the study about 3.5 km2,63.6% of the total forested area of the park wasmonitored (5.5 km2) (table 1).

    Sightings almost exclusively regard C. aethiops.A total of 19 troops were located from amongall habitats except swamps. A total of 166 animalswere observed inside the forested region(table 1), mainly sighted in the gallery forest.The estimated total density of C. aethiops in thepark area is 47.43 individuals per km2.

    Only an elusive sighting of P. kirkii was noted,this occurring in the gallery forest in the south

    Table 1. Distribution and habitat preferenceof C. aethiops: Tl. Transect lenght (in km);V. Visibility (in m); Nt. Number of troops;Ni. Number of individuals; D. Estimateddensity; * Distance not calculated becauseit was a sighting from the boat.

    Distribucin y preferencia de hbitat deC. aethiops: Tl. longitud del transecto (enkm); V. Visibilidad (en m); Nt. Nmero degrupos; Ni. Nmero de individuos; D.Densidad estimada; * Distancia no calculadapor tratarse de una observacin realizadadesde el barco.

    Tl V Nt Ni D

    Gallery forest 41.5 20 12 95 57.23

    Mangrove 5.1 1* 27

    Savannah 8.7 43 4 25 33.42

    Swamp 3.3 0 0

    Cultivation 9.7 43 2 19 22.78

    Total 68.3 19 166 47.43

  • Animal Biodiversity and Conservation 24.1 (2001) 17

    Fig. 1. Pemba island with insert indicating the study area and the 14 transects walked. Circlesshow the location of recent Procolobus kirkii sightings.

    Fig. 1. Isla de Pemba con el rea de estudio indicada y las 14 transecciones realizadas. Los crculosindican la localizacin de avistamientos recientes de Procolobus kirkii.

    of the Ngezi Park near Bandarikuu village with acount of three individuals. Our field observation,however, was supplemented by frequentinterviews with local people regarding recentsightings of the red monkeys (as the Procolobusmonkeys are known). These interviews confirmedthe presence of a small troop of 5 to 7 individualsin the Bandarikuu area corresponding to oursighting. Furthermore, most people interviewedreported recent sightings in two other locationsin the park: the first in the Makangale schoolarea, in a mosaic habitat of forest and rubberplantation, with counts of 5 to 8 individuals; anda second sighting, confirmed by most interviews,indicated an area near the east section of theWumawimbi beach in a mosaic of mangrove andgallery forest, with counts of 5 to 7 individuals.Finally, various people interviewed reported thepresence of another small troop of red monkeys,4 to 6 individuals, 6 km south of the Ngezi Forestpark, in a region with abandoned clove plantation,between the town of Conde and city of Wete.

    The home range of Procolobus is particularlysmall and all these sightings are too far fromeach other to be the same troop shifting around(fig. 1). A small population of P. kirkii can thus be

    confirmed that still survives in the Ngezi Forest ofPemba, and some individuals have even movedoutside the park area. However, the estimatedabundance of the whole population in the NgeziForest region does not exceed 1530 units (lessthan 6 individuals/km2), confirming difficulties inthe diffusion of these translocated Procolobuskirkii populations (STRUHSAKER & SIEX, 1998).

    A sympatric cohabitation with a relatively highdensity of C. aethiops should not be a majorproblem for P. kirkii which has very differentdietary preferences, and favors mangrove andswamp areas little used by C. aethiops (STRUHSAKERet al., 1997). Most problems and risks for theirsurvival and growth in number were suggestedthat comes from the local people, as in the caseof the Jozani park population in Zanzibar (SIEX &STRUHSAKER, 1999b). In the interviews with locals,it emerged that as the result of local superstition,farmers in Pemba fear and occasionally harassthis species of monkey as they are considered tobring bad luck.

    To promote the conservation of this beautiful,unique and elusive Colobinae population, wesuggest the interest to develop awarenessamongst the local people that these animals are

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  • 18 Camperio Ciani et al.

    not only harmless but that their protection andan increase in numbers will eventually bebeneficial in attracting tourists to the NgeziPark, as occurred in Zanzibar.

    Acknowledgements

    We wish to thank K. Siex for suggesting thisstudy and introducing us to the Jozani Park. Wethank the direction of the Ngezi Forest Park fortheir enthusiastic collaboration in the field.Special thanks to all the members of the GEAPemba expedition who funded and volunteeredin this project.

    References

    CAMPERIO CIANI, A., MARTINOLI, L., CAPILUPPI, C.,ARAHOU, M. & MOUNA, M., 2001. Effect ofWater Availability and Habitat Quality on Bark-Stripping in Barbary Macaques. ConservationBiology, 15(1): 259265.

    CAUGLEY, G., 1977. Analysis of vertebratepopulations. John Wiley and Sons Ltd., NY.

    COONEY, D. O. & STRUHSAKER, T. T., 1997. Adsorptivecapacity of charcoal eaten by Zanzibar redcolobus monkeys: implications for reducingdietary toxins. International Journal ofPrymatology, 18(2): 235246.

    SIEX, K. S. & STRUHSAKER, T. T., 1999a. Colobusmonkeys and coconuts: A study of perceivedhumanwildlife conflicts. Journal of AppliedEcology, 36(6): 1,0091,020.

    1999b. Ecology of the Zanzibar red colobusmonkey: demographic variability and habitatstability. International Journal of Prymatology,20(2): 163192.

    STRUHSAKER, T. T., COONEY, D. O. & SIEX, K. S., 1997.Charcoal consumption by Zanzibar red colobusmonkey: its function and its ecological anddemographic consequences. InternationalJournal of Prymatology, 18(1): 6172.

    STRUHSAKER, T. T. & SIEX, K. S., 1996. The Zanzibarred colobus monkey Procolobus kirkii:conservation status of an endangered islandendemic. African Primates, 2(2): 5461.

    1998. Translocation and introduction of theZanzibar red colobus monkey: Success andfailure with an endangered island endemic.Oryx, 32(4): 277284.

  • 19Animal Biodiversity and Conservation 24.1 (2001)

    2001 Museu de ZoologiaISSN: 1578665X

    DomingoRoura, X., Marmi, J., LpezGirldez, J. F. & GarciaFranquesa, E., 2001. New molecular challengesin animal conservation. Animal Biodiversity and Conservation, 24.1: 1929.

    AbstractAbstractAbstractAbstractAbstractNew molecular challenges in animal conservation. The contribution of genetics to wildlife conservation hasbeen stressed often forgetting the existing theoretical and empirical limitations in the use of genetic informationto solve ecological and demographic problems. The possibilities of molecular analyses are extensive and theautomation of procedures is increasing the efficiency and reducing the cost of molecular technology. With largeamounts of molecular data already available, the interest is switching towards the analysis of these data andthe interpretation of genetic variability within and across species from a functional perspective. The understandingof the link between genetic variation and fitness or survival is essential in conservation biology and thisunderstanding needs the combination of molecular data with nonmolecular (e.g. physiological, behaviouraland ecological) data. Progress in this promising field will depend on the trust and collaboration betweenmolecular and field biologists.

    Key words: Review, Molecular techniques, Animal conservation, Fitness, Genetic variation.

    ResumenResumenResumenResumenResumenNuevos retos moleculares en la conservacin animal. La contribucin de la gentica a la conservacin de lavida salvaje ha sido enfatizada, olvidndose a menudo que existen limitaciones tericas y empricas sobre el usode la informacin gentica para solucionar problemas ecolgicos y demogrficos. Los anlisis molecularesofrecen numerosas posibilidades y la automatizacin de los procesos est incrementando la eficiencia yreduciendo los costes de la tecnologa molecular. Con grandes cantidades de datos moleculares ya disponibles,el inters se est desplazando hacia el anlisis de dichos datos y la interpretacin de la variabilidad genticaintraespecfica e interespecfica desde una perspectiva funcional. La comprensin del vnculo entre variabilidadgentica y eficacia biolgica o supervivencia es esencial en la biologa de la conservacin, requiriendo estacomprensin la combinacin de datos moleculares con datos no moleculares (por ejemplo fisiolgicos, decomportamiento y ecolgicos). El progreso en este campo tan prometedor debe basarse en la confianza y lacolaboracin entre bilogos moleculares y de campo.

    Palabras clave: Revisin, Tcnicas moleculares, Conservacin animal, Eficacia biolgica, Variacin gentica.

    (Received: 17 IX 01; Final acceptance: 10 X 01)

    Xavier DomingoRoura(1), J. Marmi & J. F. LpezGirldez, Unitat de Biologia Evolutiva, Dept. de CinciesExperimentals i de la Salut, Univ. Pompeu Fabra, Dr. Aiguader 80, 08003 Barcelona, Espanya (Spain). XavierDomingoRoura, Wildlife Conservation Research Unit, Dept. of Zoology, Univ. of Oxford, South Parks Road,Oxford OX1 3PS, UK. Eullia GarciaFranquesa, Museu de Zoologia de Barcelona, Passeig Picasso s/n, 08003Barcelona, Espanya (Spain).

    (1) email: [email protected]

    New molecular challenges in animalconservation

    X. DomingoRoura, J. Marmi, J. F. LpezGirldez& E. GarciaFranquesa

  • 20 DomingoRoura et al.

    Rationalising the use of molecular biology

    The current diversity of molecular techniquesoffers a wide range of possibilities to supportdecision makers, and genetic studies are becominga primary argument in wildlife conservation. Theimportance of genetic variation in biodiversityevaluation has been recognised (EHRLICH & WILSON,1991). Molecular biology tools have already beenused to guide expensive conservation programs,including risky reintroduction projects (e.g. brownbear Ursus arctos [TABERLET & BOUVET, 1994];bearded vulture Gypaetus barbatus [NEGRO &TORRES, 1999]). The protection of genetic diversityhas been incorporated into national andinternational legislation.

    To optimise the use of molecular biology inconservation, a wise rationalisation of thetechniques and a realistic interpretation of thedata produced are needed. Technologicalseduction and the availability of numerousinformative techniques should not interfere withthe recognition of the actual limitations of thesetechniques, both in the theoretical ground andin supporting the real problems that nature isfacing (HEDRICK, 1996). For instance, it is importantto recognise that molecular information mightnot be as critical for the immediate survival of aspecies as improving its habitat (CAUGHLEY, 1994)and reducing the exploitation of natural resourcesin this habitat (BEGON et al., 1999). Currentlimitations are also evident from the recognition,for instance, that no agreement has yet beenreached on how to incorporate genetic diversityinto landuse planning (MORITZ & FAITH, 1998).

    It is also important to note that special careneeds to be taken before reaching managementconclusions in endangered species, where in spiteof the urgency implied, erroneous recommendationscould be detrimental to a species and ecosystem.Recommending the separate management ofalreadyreduced populations could promoteinbreeding. Proposing population intermixing couldpromote the hybridisation of specific adaptationsto a particular environment (WAYNE et al., 1994).

    In this work, the wide variety of moleculartechniques available to support wildlifemanagement are reviewed and relevant examplesare provided in order to better understand whenthese techniques are used (table 1). The gap thatexists between technological possibilities andtheir use can thus be recognized to interpret thecomplexity of life is noted. Finally, molecularand nonmolecular biologists are appealed tocollaborate in tracing the link between genesand adaptation so as to progress in many fieldsof life sciences including conservation biology.

    Information contained in the DNA

    Variation at a given DNA region is a consequenceof evolutionary forces such as mutation, selection,

    genetic drift or recombination that have actedover the DNA and the species (GRAUR & LI, 2000;BERTRANPETIT, 2000). Within and across populationsand species the coalescense of genomic regionscan be traced back and the time when genes orgenome separated can be infered. Similarityrelationships between DNA segments can alsobe evaluated to infer relationships betweengenes, individuals and groups of individuals. Ifwe compare derivative characters with theirgeographic distribution, we can infer gene flowand colonisation events. In addition, thedistribution of alleles and the structure of thegenetic variation might be used to inferdemographic parameters such as population sizeand subdivisions (LUIKART & ENGLAND, 1999).

    A wide variety of polymorphic DNA regionswith different mutation patterns and rates havebeen recognised. The choice of one or anotherregion will depend on the objectives of ourresearch. Most nuclear genome regions arediploid and inherited in an autosomal andcodominant fashion affected by recombination.They can code for RNA or be noncoding regions.In wildlife studies, microsatellites or STRs havebeen widely used (QUELLER et al., 1993; LUIKART &ENGLAND, 1999). They consist of a short string ofone to ten base pairs repeated in tandem andare dispersed throughout the genome. They arehighly polymorphic due to the variation in thenumber of repeat units and most behave asneutral markers. Minisatellites are also tandemlyrepeated strings of longer repeat units (JEFFREYSet al., 1985). The number of repeats is inheritedand variable among individuals. This variabilitycan be detected with a probe that will attach toa single or several complementary DNA fragmentsamong all DNA fragments distributed throughan electrophoresis gel, providing a pattern ofbands for comparison.

    Some microsatellites and minisatellites areassociated with mobile genetic elements, anotherDNA class that is currently gaining support forphylogenetic inference (BUCHANAN et al., 1999).These mobile or interspersed elements ofdifferent families and subfamilies occurthroughout the genome. Short InterspersedElements (SINEs) are excellent markers formolecular phylogeny since their integration at aparticular position in the genome can beconsidered an unambiguous derived homologouscharacter (TAKAHASHI et al., 1998). MitochondrialDNA (mtDNA) sequences include the other majorgroup of markers widely used in wildlife analyses(AVISE, 1994). Mitochondrial DNA is haploid,recombination free and maternally inherited. Ithas a low frequency of insertion, deletion andduplication events and an evolutionary rate 510times higher than single copy nuclear genes(BROWN et al., 1979).

    Conclusions in animal conservation should besupported by the analyses of several independentdata sets (WAYNE et al., 1994). If we use different

  • Animal Biodiversity and Conservation 24.1 (2001) 21

    Table 1. References with examples on the applications of molecular biology technologies towildlife management and conservation.

    Tabla 1. Referencias con ejemplos de las aplicaciones de tecnologas de biologa molecular a lagestin y conservacin de la vida salvaje.

    Technique

    Allozymes

    Reference: MERENLENDER et al. (1989)

    Purpose: quantification of genetic variation and differentiation in African rhinoceroses(Ceratotherium simum and Diceros bicornis) and Asian rhinoceroses (Rhinoceros unicornis)

    Results: low levels of intraspecific variation found below the levels expected incomparisons among subspecies

    RFLPs

    Reference: WATKINS et al. (1988)

    Purpose: quantification of Major Histocompatibility Complex (MHC) polymorphism incottontop tamarin (Saguinus oedipus )

    Results: very low levels of polymorphism found in its MHC class I

    DNA Fingerprinting

    Reference: PACKER et al. (1991)

    Purpose: study of the kinship structure in lion (Panthera leo) social groups

    Results: female within the same group are closely related, whereas males can be eitherrelated or unrelated. Reproductively active males are usually unrelated to group females.Males only act as nonreproductive helpers in coalitions composed of close relatives

    Sequencing

    Reference: BAKER et al. (2000)

    Purpose: determine the origins of whale products purchased from markets in Japan andthe Republic of South Korea

    Results: some protected species, such as baleen whales and sperm whales, wereidentified among the commercial products analysed

    SSCP & Sequencing

    Reference: SHAFFER et al. (2000)

    Purpose: screening population structure and identification of management units inYosemite toad (Bufo canorus)

    Results: different genetic substructure and no shared haplotypes among animals fromYosemite and Kings Canyon National Parks. Animals from the two parks should bemanaged as different units

    RAPDs, DGGE & Sequencing

    Reference: NORMAN et al. (1994)

    Purpose: analysis of population structure and identification of management units ingreen turtles (Chelonia mydas)

    Results: IndoPacific rookeries include a number of genetically differentiated populations,with minimal femalemediated gene flow among them

    RAPDs

    Reference: NEVEU et al. (1998)

    Purpose: comparison of the genetic diversity of wild and captive populations of mouselemur (Microcebus murinus)

    Results: captive groups have lost genetic variation in comparison with wild groups

  • 22 DomingoRoura et al.

    types of molecular data with different mutationrates we might be able to separate ancient fromrecent events. Another alternative is the comparisonof maleinherited DNA regions (i.e. nonrecombining regions of the Ychromosome) versusfemaleinherited DNA regions (such as mito-chondrial DNA) to understand the contribution ofeach sex in determining genetic diversity (MELNICK& HOELZER, 1992; PREZLEZAUN et al., 1999). Thisanalysis can contribute to understanding how abalance is achieved between the proportion ofindividuals leaving the natal area and theproportion remaining philopatric to minimiseinbreeding and resource competition (GOMPER etal., 1998). To identify individuals, populations orspecies it is often recomended to work with geneticmarkers that are neutral and therefore goodindicators of ancestry or relationship (HEDRICK, 1996).

    However, there is some concern regarding howneutral characters obtained from noncodingregions reflect the diversity of functional attributes(WILLIAMS et al., 1994; LYNCH, 1996).

    Technology available

    The main goal of molecular techniques is todetect the variation in DNA sequences, directlythrough sequencing or indirectly through othermethods sensitive to sequence variations. Thisvariation can be detected using a wide range oftechniques. A first group of techniques includingisozymes and restriction fragment lengthpolymorphisms (RFLP) is based on the differentialmobility of proteins and DNA fragmentsrespectively (due to their different charge or size)

    Technique

    AFLPs

    Reference: GIANNASI et al. (2001)

    Purpose: exploring the possibilities of AFLPs for phylogenetic reconstruction in thesnake Trimeresurus albolabris

    Results: T. albolabris is not monophyletic

    Microsatellite analysis

    Reference: CIOFI & BRUFORD (1999)

    Purpose: assess the level of genetic variability and gene flow among populations ofKomodo dragon (Varanus komodoensis)

    Results: high levels of genetic diversity and gene flow between Rinca and Flores Islands,highest levels of genetic divergence in Komodo Island and low levels of geneticvariability and gene flow in Gili Motang Island

    Microarrays

    Reference: TROESCH et al. (1999)

    Purpose: genotyping and identification of Mycobacterium species

    Results: the array can identify species within the genus Mycobacterium and detect drugresistance

    Minisequencing

    Reference: MORLEY et al. (1999)

    Purpose: assay the effectivity of fluorescent minisequencing of mtDNA for forensic usein animal, bacterial and fungal species extracts

    Results: the technique is reliable, reproducible and suitable for forensic uses in a widerange of organisms

    Quantitative PCR

    Reference: FELDMAN et al. (1995)

    Purpose: detection of malaria infection in Hawaiian birds

    Results: avian malaria was more widespread in Hawaii than previously thought

    Table 1. (Cont.)

  • Animal Biodiversity and Conservation 24.1 (2001) 23

    in an electrophoretic field (MLLERSTARK, 1998;BRETTSCHNEIDER, 1998). Hybridisation between alabeled DNA fragment or probe and a target DNAis the principle involved in many other techniques(SAMBROOK et al., 1989).

    With the discovery of the polymerase chainreaction (PCR) (SAIKI et al., 1988), a new wave ofmolecular techniques appeared. One importantadvantage of the PCR is that a given DNA fragmentcan be isolated and copied millions of timesreliably and quickly using temperature cycles anda thermally stable polymerase. This allows the useof minute amounts of DNA in molecular studies,such as those obtained from biological remnantsobtained non-invasively (WOODRUFF, 1993).

    Sequencing

    The complete sequencing of the whole genomeis the most detailed method to detect geneticvariability. However, sequencing completegenomes is tedious and expensive and moststudies rely on the sequencing of a minute portionof the genome and the assumption that variationwithin the fragment sequenced represents thevariation along the whole genome. Sequencingof PCR products of up to several hundred basepairs is a widely used methodology in life sciences.During the sequencing reaction of a PCR product,a large number of fragments differing by anucleotide in length and with the last baselabelled with a specific fluorochrome dependingon its identity are obtained (WEAVER & HEDRICK,1992). When these sequencing products ofdifferent length are electrophoresed in a DNAsequencer, the ladder of fluorochrome signalsobtained will indicate the nucleotide sequenceof the PCR product under analysis. It is commonpractice to deposit the sequences obtained inpublic databases, facilitating both the comparisonand complementation of ones own data withthe data from the same or other species obtainedby other researchers.

    Sequencing can be combined with othermethods to reduce its cost. A first group of PCRbased methods (Heteroduplex analysis, SingleStrand Conformation Polymorphisms, DenaturingGradient Gel Electrophoresis and TemperatureGradient Gel Electrophoresis) consists of screeningtechniques for detecting sequence variation in PCRproducts of identical sizes, without the need to gothrough sequencing. These protocols are based onthe physical behaviour of DNA during electrophoresis in acrylamide gels. The use of thesemethods is adequate when dealing with a largenumber of samples and when alleles are shared bymany individuals (LESSA & APPLEBAUM, 1993).

    Heteroduplex analysis

    Heteroduplex analysis starts with the denaturingof the PCR product at 95C and its subsequentrenaturation before electrophoresis (LESSA &

    APPLEBAUM, 1993). Using this technique it ispossible to distinguish between homozygous andheterozygous DNA fragments. If a samplecontains two different alleles, heteroduplexmolecules (hybrids of the two strands belongingto different alleles) are obtained. Since theseheteroduplexes have one or more mismatches intheir double strands, they migrate onto the gelmore slowly than the homoduplex moleculesobtained from the hybridization of strandscontaining the same allele.

    SingleStrand Conformation Polymorphism (SSCP)

    SSCP is a simple and fast method for screeningDNA fragments for nucleotide sequence poly-morphisms. PCR products that have beendenatured by temperature and/or chemicals areloaded and run onto a nondenaturing polya-crylamide gel. The electrophoretic mobility ofeach singlestranded DNA fragment depends onits secondary structure, which in turn depends onits nucleotide sequence (JORDAN et al., 1998). SSCPcan distinguish DNA fragments that differ only byone base-pair substitution in a fragment of up toseveral hundred nucleotides (ORITA et al., 1989).

    Denaturing Gradient Gel Electrophoresis (DGGE) andTemperature Gradient Gel Electrophoresis (TGGE)

    DGGE and TGGE work over double stranded DNA.In these methods, PCR products are loaded onto apolyacrylamide gel and run in a linear gradient ofconcentration of denaturing solvents (urea,formamide) or temperature respectively (LESSA &APPLEBAUM, 1993). The point along the gradientwhere the DNA fragment is partially denatured iscalled the melting point. This point depends onthe overall base composition and the interactionsacross the molecule and can be modified by pointmutations that will be reflected in the gel.

    Randomly Amplified Polymorphic DNAs (RAPDs) andAmplified Fragment Length Polymorphisms (AFLPs)

    The principle of the RAPD technique is thesimultaneous amplification of DNA regions byusing a single randomly chosen primer whichacts as both forward and reverse (GROSBERG et.al., 1996). This primer is able to hybridise withmany sites of target DNA, but amplification onlyoccurs when the primer anneals at two sites onopposite strands separated by a reasonabledistance for the PCR to work (20 to 2000 bp).These fragments are then separated in anelectrophoresis gel and stained with chemicalssuch as ethidium bromide or silver nitrate. Thegels can be scored as the presence or absence ofa band of a specific molecular weight. Bands ofdifferent sizes usually represent independent loci.RAPDs are treated as neutral and anonymousmarkers, can be generated quickly and a largenumber of individuals can be processed in a

  • 24 DomingoRoura et al.

    short time. However, results are difficult torepeat, a band can contain more than oneamplification product that can not be distinguish-ed and it is difficult to estimate allelic frequenciesbecause homozygotes can not be distinguishedfrom heterozygotes. In addition, it is sometimesdifficult to know whether the variation is neutralor whether it follows Mendelian inheritance.

    In AFLPs, genomic DNA is digested withrestriction enzymes and the goal is to reduce thecomplexity of the initial mixture of fragments.To achieve this reduction a subset of fragmentsis biotinylated and selected by union tostreptavidin-coated paramagnetic beads (sincebiotine binds covalently to streptavidine)(MATTHES et al., 1998). The unbound fragmentsare washed and discarded. A subset of thebiotinylated fragments is then amplified by PCRto further reduce complexity. Finaly, PCR productsare analysed by denaturing polyacrilamide gelelectrophoresis and revealed by autoradiography.AFLPs are more informative and easier toreproduce than RAPDs.

    Automation required

    Automation is a key issue in molecular biologyand the machinery used in the automated analysesof humans and model animals is later adapted towildlife research. Automated procedures arecurrently used for standard procedures such asDNA isolation or library construction and spottingbut also for the fast scoring of genetic variabilityamong individuals with technologies such asmicroarrays or quantitative PCR.

    Microsatellite multiplexing

    Several microsatellite loci can be amplified in asingle PCR reaction containing different primers(GILL et al., 1995). The primers are labeled withdifferent fluorochromes and amplify fragmentsof different lengths. When the multiplex PCRreaction is run in an automated sequencer it ispossible to sequentially detect the length of thedifferent PCR products corresponding to thealleles of the different microsatellite loci.

    DNA array technology

    A DNA array consists of up to thousands of DNAstrings attached in order over a solid support(SOUTHERN et al., 1999). An unknown sample ispassed over the array and it will hybridize uponthe immobilised probes when finding acomplementary sequence. The reverse is alsopossible when a known probe hybridises uponunknown immobilised fragments. The fullmicroarray equipment consists of a machine toproduce the array and a machine with afluorescence laser scanner to read the signal andtranslate this signal to a computer. The great

    advantage of microarray technology is that itallows the fast detection of sequence informationfrom a large number of loci or individuals at thesame time. Paradoxically, one of the mainproblems encountered with microarray technol-ogy is that it generates such a large amount ofinformation that results are often difficult tointerpret. Microarrays are used, for instance, tomonitor RNA expression and gene function (DESAIZIEU et al., 1997; WODICK et al., 1997; CHO etal., 1998) or to detect single nucleotidepolymorphisms (SNPs) (CHAKRAVARTI, 1999). Allstudies published used model species and, as faras we know, no study using microarraytechnology has yet been performed in any specieswith a conservation perspective.

    Minisequencing

    The technique consists of a PCRbased mini-sequencing reaction where the polymerase addsa single nucleotide. Primers finalise just beforethe polymorphic position that needs to beinterrogated. The polymerase extends the firstbase position after the primer with labelled newnucleotides and the identity of the incorporatednucleotide can be determined with an automatedsequencer. Several reactions can be performedsimultaneously with primers of different sizes. Itis also possible to conduct a minisequencingreaction in a DNA array (HACIA, 1999; RAITIO etal., 2001).

    Quantitative PCR

    Quantitative PCR consists of a reaction thatdetects and quantifies nucleic acid sequenceseither as a final product or while the reaction isbeing produced. The protocol is based on thedetection of fluorescence emitted by thedegradation of an internal labelled oligocomplementary to our sample when the PCRproceeding is being produced. The outcome isthe quantification of a PCR product that can beused in gene expression studies (DE KOK et al.,2000), to evaluate viral load (LIMAYE et al., 2001),and to detect transgenes (FAIRMAN et al., 1999),duplications and deletions (AARSKOG & VEDELER,2000; WILKE et al., 2000) and SNPs (BREEN et al.,2000). Quantitative PCR and minisequencing canbe cheaper alternatives to microarrays for thestudy of SNPs if a moderate number of SNPs andindividuals are to be analysed.

    Looking for the link between molecular dataand conservation

    Technological resources are available, but theconnection between molecular variability andthe needs of endangered species is notstraightforward. Gene dynamics is complex, mostphenotypic characters are multigenic, and the

  • Animal Biodiversity and Conservation 24.1 (2001) 25

    genetic machinery is loaded with complicatedgene interactions and epistases (HEDRICK, 1996).More than one protein can be translated from asingle gene due to alternative splicing (GRAUR &LI, 2000). Genetic linkage can also mask the roleof important genes. In addition, the relationshipbetween gene and environment is often difficultto discern (FALCONER, 1989). All levels of lifeexpression and population processes are complexand manifold and quick fixes to animalmanagement questions based on simplemolecular biology analyses should be avoided.

    Genetic diversity has been linked to speciesrichness and to better chances to cope efficientlywith enviromental change (HEDRICK & MILLER, 1992;OBRIEN et al., 1985). Consanguineous matingspromote the existence of deleterious genes inhomozygosis, which can be detrimental forsurvival and reproduction. In theory, fitness insmall populations will decline due to theaccumulation of detrimental mutations (LYNCHet al., 1995a, 1995b). However, the importanceof genetic variability for species survival is notclearly defined. In practice, at least somepopulations can survive in spite of having lowgenetic variability (e.g cheetahs [Acinonyxjubatus] [OBRIEN et al., 1987], molerats[Heterocephalus glaber] [FAULKES et al., 1990;REEVE et al., 1990] and Eurasian badgers [Melesmeles] [DOMINGOROURA, 2000]). The empiricalrelationship between genetic distance and fitnessis likely to be speciesspecific and is unlikely tobe linear (LYNCH, 1991).

    In the past, considerable effort has beendevoted to describe key demographic numbersrequired to maintain the necessary geneticvariability needed for species survival (SOUL, 1987).However, key numbers are unlikely to beapplicable across populations or habitats. In thelast decade, polemics concerning the existence ofkey numbers for survival have often given way toother discussions, not often based on molecularinformation. Within species, conservationstrategies have been proposed on the basis of theexistence of Evolutionary Significant Units (ESUs)which have been defined as population units thatmerit separate management and have high priorityfor conservation (RYDER, 1986). The use of ESUs inconservation has signified an upgrade fromprevious strategies that only gave importance toindividual numbers without considering differences among individuals of the same species.However, a compromise has not yet been reachedregarding the relative importance of ecologicaladaptation and genetic variability to determinethese units (MACE et al., 1996). Furthermore, theinadequacies of the dichotomy implied in the ESUconcept in a world ruled by a continuum ofpopulation differentiation have been noted(CRANDALL et al., 2000).

    Across species, molecular techniques are alsoat the base of new strategies to support anintegrated approach to conservation, focusing on

    the preservation of evolutionary diversity insteadof focusing on species number (MAY, 1990; MACE etal., 1996) or singlespecies management. In thiscase, molecular data should play a predominantrole in the selection of areas that containevolutionarily distant lineages and areas ofpotential evolutionary novelty, such as multispeciescontact zones (MORITZ & FAITH, 1998). The protectionof these areas is likely to preserve large amountsof evolutionary heritage and will maximise theevolutionaryresponse potential to perturbations(PETIT et al., 1998). In fact, discussing key numbersand even singlespecies conservation strategiesmight be naif in the face of the immense complexityof nature.

    Looking for the link between gene andfunction

    The rapid development of molecular geneticsfor biomedical and industrial purposes facilitatesthe access to molecular technology. Resolution isalso increased with new techniques and a highernumber of markers. The increase in the numberof markers known in any species means greaterprobabilities to detect major loci that influencequantitative traits.

    As we learn more about DNA, molecularinformation will be better understood when usedin combination with physiological, demographic,ecological and behavioural data collected in thefield (HAIG, 1998). Data can originate from anyparameter that can group individuals in relation totheir evolutionary origin and/or ecological needs. Inanimal conservation it is not enough to understandand describe molecular variation or even ecologicaland demographic characteristics using moleculartools. We need to find loci that have variants thatare responsible for low fitness and survival.Ecologically relevant heritable traits might need tobe emphasised (CRANDALL et al., 2000).

    Nevertheless, fitness measurement might bedifficult in endangered species. Since a selectivedifference smaller than the reciprocal of twicethe effective population size (1/2Ne) is effectivelyneutral (KIMURA, 1979), small selective differencesare unlikely to be of adaptive significance in mostendangered species. A further complication arisesfrom the possible differences between formerand current selection and adaptation processes.The habitat currently used by a rare species canbe marginal and might no longer reflect theenvironmental condition in which the traitsevolved (JOHNSON et al., 2000). This is especiallytrue for carnivores since human expansion hasconsiderably altered their distribution and ecology(GRIFFITHS & THOMAS, 1993).

    To unravel the link between gene and functionor adaptation is not a goal exclusive toconservation genetics. For instance, to clarifythe function of genes that are likely to beresponsible for diseases is a major enterprise in

  • 26 DomingoRoura et al.

    current biomedicine. Since the link betweengenotype and phenotype is still widely unknown,the potential of molecular biology in wildlifemanagement and conservation is still at a veryearly stage. At this point, still far from applyingthe functional interpretation of genetic variationto wild species, to advance the understanding ofenhanced fitness and the evolutionary paths ofphysiological systems, several approaches can beconsidered. T