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SDO400002 Taxonomical, Anatomical and Chemotaxonomical I - Studhs Of 12 Mosses From Jebel Marra (Darfur State, Sudan), With Special Reference To Their Localities. BY 1kram Madani Ahmed B. Sc. Hon s Botany December 1994), M. Sc. Forestry (Septernber 1997) A tesis submitted in fulfillment of the requirements for the degree of Doctor of Philosophy in Botany Department of Botany Faculty of Science University of Khartoum February 2003

Taxonomical, Anatomical I - and Chemotaxonomical State, … · 2004. 2. 18. · Philonotis longiseta j Barbula ehrenbergii jj Thiddium furfurosum _ Ceratodon purpureus,. Philonotis

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  • SDO400002

    Taxonomical, Anatomical and ChemotaxonomicalI -Studhs Of 12 Mosses From Jebel Marra (DarfurState, Sudan), With Special Reference To Their

    Localities.

    BY

    1kram Madani Ahmed

    B. Sc. Hon s Botany December 1994), M. Sc. Forestry (Septernber 1997)

    A tesis submitted in fulfillment of the requirements for thedegree of Doctor of Philosophy in Botany

    Department of BotanyFaculty of Science

    University of Khartoum

    February2003

  • DISCLAIMER

    Portions of this document may beillegible in electronic image products.

    Images are produced from the bestavailable original document

  • TABLE OF CONTENTS

    ACKNOW LEDGEM ENIS ............................................. i

    ABSTRACT ......................................................... ii

    ARABIC ABSTRACT .......................................... iv

    7CHAPIfER-ONE-INTRODUCTMNAND-LITERATURER-VIEW---]L

    The study area .................................. 4

    Location ..................................... 4

    Topography ...................................... 4

    Geology and geomorphology .................. 7

    Climate ........................................... 9

    Vegetation ....................................... 9

    CHAPTER TWO: MATERIAL AND METHODS .......................... 14

    A: Taxonomical investigation .................. 4

    Taxonomical procedures .................... 15

    The relevant consulted floras and publications... 18

    B: Anatomical investigations .................. 20

    C: Chernotaxonomical. investigations ......... 22

    Qualitative analysis of the cloroplast pigments..22

    Quantitative analysis of the chlorophylls ......... 22

    Nitrogen content ................I..................... 23

    Elemental analysis ...................................... 25

  • CHAPTER THREE: RESULTS ............................................. 26

    A: Taxonomical results ........................ 26

    -Sudan-bryoflor-f�he-k4ist�.--.--.

    Present study taxonomic descriptions ........ 31

    Key to the studied genera ...................... 61

    B: Chemotaxonomical results ............... 62

    Elemental analysis ............................ 62

    Sodium content ............................. 62

    Potassium content ................... 62

    Zinc content ......................... .65

    Cobalt content ....................... 65

    Copper content ............................ 70

    Manganese content ... .................... 70

    Iron content ................................. 75

    Chlorophyll concentrations ........ 84

    Absorption coefficients ..................... 84

    Nitrogen content ............................. 90

    C:Numerical taxonomy ......................... 93

    CHAPTER FOUR: DISCUSSION ............................................... 97

    References ............................................................ 102

  • ACKNOWLEDGEMENTS

    I would like to express my sncere thanks and appreciations to my

    supervisor Dr. S. El Tiganii for her advice and encouragement throughout this

    research.

    I'm keenly 'indebted to Prof E. Ban', Prof W. E. El Saadawi, Prof T.

    Heddrson and Prof B. Osl�a for their valuable help various ways in te

    course of this work.

    I wish to thank my friends and colleagues for their support and special

    thanks are extended to M. Mubarak and M. Khalid for their valuable help in

    photo. sanning and pinting.

    More thanks are also extended to my family.

    i

  • ABSTRACT

    This research represents taxonomical, anatomical, and

    cheniotaxonomical stud-fdof mosses of Jebel Marra, Darfour State, Sudan.

    This study and for the first time includes the bryoflora checklist of the

    Sudan which is gathered from many previous studies, in addition to te

    present study.

    The fully identified mosses for the study area are 12 species belonging

    to different families and 10 different genera.. Three genera frorn the family

    Pottiaceae 2 from the family Bartramiaceae 2 frorn Funariaceae, and one

    genus recorded for each of the families: Ditrichaceae, Grinuniaceae,

    Bryaceae, Puidiaceae and Polytrichaceae. Key to the studied genera is

    constructed and full species description and illustrations of the most

    taxonomically inportant characters are included.

    In this study, the genus Ceratodon Brid. is recorded for the first tirne in

    Sudan, and to the specific level, species aretheountered for the first time:

    Barbida ehrenberaii, Philonotis fenuis, Philonotis lonkisela, Ceratodon

    purpureusand Thuidiumfurfurosum.

    H

  • For each species determination of the levels of the heavy metals:

    sodimn, potassimn, zinc, cobalt, copper, manganese and iron are studied,

    addition to nitrogen and chlorophyll content.

    W

  • j� t��S' ;A L)

    L)

    3 I Lp 8 12

    ,�Ar Bartramiaceae z�Wi �i L)6-- j Pottiaceae zL;W� L�j

    j Ditrichaceae J< -L-�j L i Funariaceae z�w� L�� �)L--"

    L�s- Polylrichaceae Thuidiaceae j Bryaceae Grimmiaceae

    Y L) Z,2--dl Li -�l Jj �S' Z��

    Y Jj� Ceratodon -,--j Z�-� .J� olz C,Uj

    Philonotis longiseta j Barbula ehrenbergii jj�

    Thiddium furfurosum _ Ceratodon purpureus,.� Philonotis tenuis

    -Y.4jl ZJ�l -,�w L51

    .5-S L�-�- 4 �k--

    iv

  • CHAPTER ONE

    INTRODUCTION AND LITERATURE REVIEW

    -B-ryophyte&-(mosse"verw-arts and hornworts) are small vxeen plants tat

    reproduce by means of spores (or vegetatively) instead of seeds. Most are only a

    few centimetres hgh, although some mosses attain a half meter or more. Although

    often sinall and inconspicuous, bryophytes are remarkably successful. Tey are

    sensitive indicators of air and water pollution, ad play important roles in te

    cycling of water and nutrients and in relationships with many other plants and

    animals. Bryophytes rank second (after the flowering plants) arnong major groups

    of green land plants with an estimated 15,000-18,000 species worldwide Anderson

    et al., 1990).

    Bryophytes have successfully exploited many environinents, perhaps partly

    because-�ey-me--Tm���ect--coinpetition--vMh-highef -plants-.For--such-a---smafl--

    organisms, the climate near the ground (microclimate) is often very different from

    conditions recorded by standard meteorolo ical methods, and shifts in temperature

    and humidity are often extreme. A remarkable adaptation of bryophytes is teir

    ability to remain alive for long periods without water, even under hgh

    temperatures, then resume photosynthesis within seconds after being moistened by

    rain or dew (Anderson, 1980).

  • Mosses are the higher bryophytes. They reproduce trough aternating

    generations, The first generation, the gametophyte, forins the green leafy structure.

    It produces a sperm and an egg (the gametes) which unite, when conditions are

    -favourable; - -to--graw-iM-o-4he-next-- generation,,- te -- sporophyte-or---sporA�-,-bear'mg

    structure. The sporophyte contains no chlorophyll, it grows parasitically o its

    gainetophyte mother. As the sporophyte dries ot the capsule releases spores which

    will grow into a new generation of gametophytes (Ben Hl et. al

    Mosses aid greatly in stabilization and strengthening of te fra 'le soil.

    Rhizoids of mosses have te ability to bind and adhere particles of soil together.

    Mosses also aid in water infiltration into the soil, and allow the other organisms in

    the community to collect and take up water. The mosses help creation of a

    roughened, microtopographic Srface that decreases water run-off by creating rain

    buddies wch trap and disperse precipitation through the entire area were growth

    is occtining�--Mile--perffimiing-these-func�fion-s---mosses- -aid-i"e-prevention--of-

    erosion caused by water, wind, animals or even people (BeInap, 1994).

    Mosses are reliable indicator of soil conditions because they tend to

    accumulate chemical elements directly into leaves and stems (Longton, 1980).

    Few researches have been published on the bryoflora of the Sdan Republic.

    Pettet 1967) studied mosses collected from Khartown State, between latitude 15'

    ION and 16' 30'N. He recorded six fully identified mosses from this area.

    Imain and Ghabbour 1972) regarded four Sudanese mosses collected by

    Kassas in I _e__�Wtidhf�ora� �

    2

  • Wickens 1976) recorded another four fully identified osses or

    another aea: Jebel Marra wich is also studied later by Townsend 1984).

    He added 14 ore fully dentified mosses for th-i's area.

    III 1994 Buck, the Curator of Bryophytes i New York Botanic

    Garden, dentified five new moss species for Sudan Tese species were

    collected by Bari from Ekowit area (Red Sea Hills), and kept in te Botany

    Department Herbarluin, University of Khartown.

    ---- Stiabb-a-a--e--a1-(M9) "um. -- His-- -

    study dealt with Khartoum State, in localities different from those eported

    by Pettet ( 967).

    Brain 2002), In 111's bryophyte Sub-Saharan Affican cecklist,

    worked out 41 taxa, sorne of which were already mentioned by other

    authors. He listed all fully dentified mosses for Sudan, wth their literature

    references. The floral elements entioned il all te previous published

    works are ordered and listed as a checklist for Sudan (See Chapter Tree).

    Objectives:

    The objectives of tis work were:

    1. To collect, identify the mosses of Jebel Marr a for documentation.

    3

  • 2. To study te morphology, anatomy and chernotaxonorny of' each

    moss plant

    3.1-o work out te updating bryoflora cecklist for Sudan.

    4. To construct a taxonomic relationships betwee te identified taxa

    using te Chernotaxononic data

    5. With special reference to the mosses localities, this Study also

    aimed to study mosses, each with its corresponding cheimical

    constituents in an attempt to construct a database for further

    pollution moffltorifig studies

  • The Study Area:

    1. Location:

    The study area was "Jebel Marra" which is an isolated, 3042 meters high

    volcanic massif, located about I 00 kilornetres south- west Khartoum. It rests on a

    base of Archaean rocks at the summit of an upwarding between the Chad and the

    Nile Basins. lt lies almost in the center of the African continent, being equidistant

    from the Atlantic and the Indian oceans, the Mediterranean and the Red Sea. It is

    situated in te climatically transitional Sahelian zone between the arid Sahara desert

    and the seasonally-wet tropicaf Savanna (Fig. 1) (Captin, 1959).

    Jebe dfibis about 100 kilometres north of Nyala town in Darfur State, which extends

    for more than 200 kilometres to the north. It is 80 kilometres wide at its broadest

    part. It os the highest massif in the Sudan. It lies at the intersection of two

    transcontinental tectanovolcanic zones "East North East" from Cameroon to Red

    Sea and South South East" from Libya to Kenya as shown in (Fig. 2 (Vail, 1972).

    2. Topopraphy:

    The Marra Mountains rise steeply from the flat plains of Western Sudan at

    about 1000-3042 meters elevation above sea level. The highest point occurs around

    the serrated rm of a spectacular Caldera ,some kilometres n dameter. Wthin the

    floor of this basin, at an elevation of about 2000 meters, are the two Deriba lakes,

    the smaller of which occupies a small cone as a deep Crater Lake approximately one

    kilometre across. The other is a shallow, saline, 25 kilornetres in length. A number

    of streams have cut steep gorges down the flanks of the mountains and spectacular

    waterfalls and canyons have developed (Vail, 1972).

    4

  • 20 00 200 400 6.040* 40'�-

    -2 + 2,cL-

    Jebel Marra mP SUDAN

    -00

    -20*

    20' 00 2e 40' 6

    Fla. Location of tile sudy area.(after Caphil, 1959).

  • REGIONAL SETTING OF

    J. MARRA

    0 kin

    Volcanic rocks

    a Fracture zones

    .............. International

    boundaries

    N '.MARRA

    Regional setting of Jebel Marra in north - east Affica.(afterV�11, 1972).

    6

  • 3. Geolou and Geomorpholoyy:

    The following components represent the main outlines of the geology of Jebel

    Marra:

    3. L Crystalline Basement Rocks:

    These rocks are exposed in the west around Zalingel and Kebkabeya bt are

    obscured in the east and south by a younger cover of unconsolidated alluvial silts

    and clays and sands.

    3.2 Sedimentary cover:

    Overlying the basement are a group of sediments belongngto teNi.Nan

    Sandstone Forination. Generally, poorly sorted, cross-bedded and flat-lying

    sandstone, silts and shales crop out in the north eastern part of the Marra ranges. In

    protected basins and troughs, they reach several hundreds of meters tick bt ae

    usually much thinner.

    3.3 Volcanic Complex:

    The volcanic rocks are in the form of a very large stratiform volcanic

    complex. The outpourings from several centers have coalesced to forin ranges of

    igneous rocks, and obviously great many flows from different eruptions have bilt

    up more than 2000

    meters extrUSve and pyroclassic rocks which remain today. The main

    constituents of the volcanic rocks are: Basalts, Tuffs, and trachytes (Fig. 3 (Vail,

    1972).

    7

  • 0 b

    15

    ,0

    NOEL FASHER

    GENEINA

    lk

    A L\0

    1, Q

    I NYA�A 12:x

    5PkT

    2TE 26'1 2?1

    MVIokanic rocks plugs Nubian Sandstone �Metamorphosed basement with trends

    Fig. Regional geology ofthe Jebel Marra area, Darfur State, Sudan,

    (Abbreviations- Zal,= Zalingel-, Keb = Kebkabeya).

    (After Vail, 1972).

  • 4. Climate:

    The Jebel Marra iiiassif strongly odifies te regional cmate and Increases

    tile aniount of precipitation- Rainfall is almost entirely dUring te period May to

    September, with 60 during te 111011tilS Of Rtly and August. Te total

    precipitation is below 94 or above I 10 of te normal to be expected

    every three years. Variability of te rainfall fom year to year is shown ]II (Fig. 4)

    as te total rainfall in nil.) dUring te period 1960-1999.

    The average temperatUre at Jebel Marra varies between 20'C in winter and

    27'C a( (lie beginning of te rains n May-hine. Te average diurnal variation is

    greatest dUrilIg te winter dry season (28'C) and least during tile rains August,

    10T.). Te average niaminuin temperatures are lowest in te rains (ALIgUSt, 29'C),

    rising to WC in December and 38'C in April. Te average mininium temperatures

    -are--lawest-HY-December m&January-(6C)undAlighest-in--J-Lme-ancf-J-uly,-tt-9-C)�.-14iglit--

    fi-osts alsoOCCLII- III tle lower valleys especially In te depressions.

    Mean SUrface winds are less strong within Jebel Marra than n te

    SUrrOUndings. Jebel Marra acts as a shield to te northern winds. Te relative

    humidity is gnerally low Wickens, 1976).

    5. Vegetation:

    The vegetation of Jebel Marra inassif is divided by Wickens( 976)

    -it"u-Ce type--,�--

    9

  • r" "-%rn�. 1000

    1000

    800800

    600 600

    400 400

    200 200

    0 - - - 0w (D C4 CI) t- co

    at m �m t- t- I'. -6 t" 0) OD w m 0) co Om 0) CD m qm

    1000 1000

    Boo I

    600" :0000

    400" roll- 400200 200

    0 I I 0 L I " el) W r.-

    00 co co co 00 co co co co 00 0) Im C" a) 0) 0) a) a)C) Ca -9� Cn -4 co w m 0) a,

    -FV�,.-4--TotaLraw-ULdiwing-the-period 1.-960--- fQ994SOUrce: Nyrtete Metrological Station.

    I

  • 5. I. Hill Savanna:

    The hill savanna is described as different cominunities with marked

    transitions: Acacia mellifiera hill thorriland occurs on the drier eastern slopes of the

    if and merges with the Anogeissus leiocarpus hl savanna. Elsewhere o te

    massif there-lis a-transition-wifl-i-inereasing---altitude-ftoin-Anogeissus-leH�curpu,,�-a-s-a--

    dominant tree species, through Terminalia brownie to Combretum n7olle. However,

    zonation is so interfered by man's activities.

    5.2. Gallery Forest:

    There are several true gallery forest on the south western slopes of Jebel

    Marra massif. The larger gallery forest is at Saur, above Golol and extends for

    several kilometres. The main body of the forest lies in a deep precipitous-sided

    gorge into wich access is impossible, both the upper and the lower extremities are

    protected by waterfalls. There is an abrupt change in the vegetation frorn open

    An th�ealk� forest. Anogeissus

    leiocarpus, Combretum molle and Entada abyssinica frequently occur along the

    forest margin

    The main constituent of the forest is Terma orientalis. Ferris are also plentiffil

    along the stream banks in shaddy moist places. To the east of Saur, there is a

    minor gallery forest where Anogissus leiocarpus forms a closed canopy.

    The trd gallery forest is at Mortagello, by the Forest Department Rest

    House. The forest is in a deep bassalt gorge; protected upstream by a waterfall bt

    merging downstream into woodland where the gorge widens into. V -shaped valley.

    Combretum malle, Lannea kerstingii and Anogeissus leiocarpus border the gorge

    rim.

  • .3. Montane Grassland:

    The transition from hill tree savanna to montane grassland takes place

    between 1800-2000 ineter. An interesting feature of this upland area is te apparent

    absence of any altitudinal zoning of the vegetation. Two distinct corni'minities are

    recognized in ts grassland zone: upland grassland which occurs' on the steeper

    slopes and upland meadows whichstf, the flatter ash plains, sei-ni-waterlogged dring

    rainy season. Olea laperrinei and Acacia albida are thinly scattered troughout te

    two grassland communities. Fig. shows the vegetation of different geornorphic

    and soil combination in Jebel Marra.

    2

  • 22' 30'

    13' 30' w e

    'la

    Al

    uld X/ 4",

    Ze,

    Z cr a 1:12,Ned ratcr. .. .. ....... ...

    k

    KaJokitting w

    ...........

    Kas I 2 30'

    '24' 45'

    8

    2 9

    i Acacia meIIifrr&-C&WMiOkPr,, africana on indurated soils

    3 .10 2 Acacia Zero on hill soils of the basement complex

    3 Acacia inelfifere-Anageissui kiocarpas mm-tic on basemcsit

    4 complex soils

    F7777771 4 Acacia styal- Balanites aegyptiace on clay soils

    1 .... 12 5 Acacia sed- Anageiismi kiwarpui mosaic on basement comph-

    soils

    6 ---- 1 3 6 Anogeissai leiararpwi on basement complex soils

    D�. 7 hill soils8 Acacia albida - Balanitej aegipliaca on alluvial soils

    9 Acacia albida on ash piedmont soils

    to Coatbretun; Slutinasum- Terminalia laxiflora on ash picclinont -.0,

    i jAcacia welfifera on volcanic soils of the Jebel Marra musif

    12 Anageirm kiocarpus on volcanic soils of the Jebel Marta mad

    13 Upland grassland and UP12nd meadow on volcani( soils oflhr

    Jebel Marra massif

    Sketcb map of the vegetation associations.(After Wickens, 1976).

    13

  • CHAPTER TWO

    MATERIALS AND METH

    In this study, taxonomical, anatomical and chemotaxonomical investigations

    -were carried out.,

    A) Taxonomical Investii!ation:

    Study area:

    A general srvey was carried out in Jebel Marra area (Darftir State, Western

    Sudan) during the season July-August 1998 in which sites of different altitudes

    were h-of'-fl forAhis-�dy-.-

    1) Togi: on the south west slope at an average altitude of approximately 2600

    meter above sea level (m.a.s.l.)

    ii) Beldong: on the south west slope at an average altitude of approximately

    2500 m.a.s.l.

    iii) Golol: on the southwest slope at an average altitude of approximately

    1500 m.a.s.l.

    _iv-) \41,rtagdlG--. -����wer-slope-at-an-altitude efWfom

    I 000 in. a. s.

    1 4

  • V) Nyrtete: on the western lower slope at an altitude of approximately I 00

    ni.a.s.l.

    Materials:

    In each site the encountered mosses were collected as mats frorn dfferent

    locations alongside "Wadis" and depressions. They were kept in paper bags labelled

    for site, date of collection and collector's name. Another, collection following te

    same procedure was carried out in the same sites in the second season July-August

    I 999.

    Methods:

    Taxonomical rocedures:

    The two field collections were then sorted out as different groups and all the

    niacrotaxonoinic caracters of each group were examined sng tile binocular

    microscope.

    For te investigations of the microtaxonomic characters, the collected inosses

    were prepared as whole mount specimens on slides using the following techniqUe:-

    Plant materials were firstly soaked in water contained in an open Petri-dish

    for 23 hours so as to become turgid, then cleaned 23 times from undesired iterns,

    such as sands and other plants depris.

    15

  • Fixation was carried out using formalin: acetic acid: alcohol solution whic is

    prepared as 5% fon-naldehyde, 5% glacial acetic acid and 90% ethyl alcohol. The

    materials were kept in the fixative until the chlorophyllW completely eached out.

    Dehydration was carried out after stretching each plant individual between

    two slides before putting tem in staining jars. Pa--n--ts-wer-etliensuT3et&(]-t-6-�e�i&s

    of concentrations of ethyl alcohol (50%, 70% and 90%). Then the slides were

    removed and the plants were transferred to wide-mouth bottles and Sbjected to 95-

    98% absolute alcohol. The last step was repeated to ensure complete dehydration.

    Each dehydration step was left for overnight.

    Cleaning of the material was done by transferring the plants to a mixture of I

    1 - xylene and cedar oil for overnight and pure xylene for half an our only. The

    last step was also repeated to ensure clearing.

    Mounting was carried out for all plants by taking them by a dry bnish ad

    placing each one on a drop of Canada balsam mountant. Lastly, using a oreceps a

    suitable-sized glass covers were used, and all slides - placed in an oven adjusted

    at 60'C for 3 days as recommended by (Johansen, 1940). The whole inount slides

    were examined microscopically for the taxonomically valued caracters. Tile

    characters were then tabulated and used later for both describing the ndividual. -tks�

    mosses and constructing� identification key. Also, fully-labelled drawings of

    different parts for each individual plant were done using a drawing tube fixed to a

    microscope.

    16

  • Photographs were taken from the whole mount slides using Leitz DalLIx 20

    -Aens mici!aseope-w��MFIS-H-eamem-

    For presentation of the whole mount samples, enlargement modified

    technique was used as follows:

    Using Focomat 11 enlarger, the prepared whole mount slides were situated on

    the negative holder and well focused. Then sensitive papers were exposed for 20

    secs. and processed as usual (developed by Metol for I min., washed by water for

    30 secs., fixed by 30% sodium thiosulphate for 10 min., washed by running tap

    water for 30 rnin. and dried) (Langford, 1975).

    -Finally, -individual-samples--wer -leane"tretched-and car fatly-presstd-as-

    herbarium specimens kept at the Herbarium of te Botany Department, University

    of Khartoum. Table (1) represents the relevant consulted floras and publications.

    17

  • Table (1): The relevant consulted floras and publications

    Floras and publications References

    The bryophy-ta of South Africa Sim 1926)

    Illustrated moss flora of Fennoscandia Nyhlom 1969)

    Bryophyte flora of the Huon Peninsula, papua New Norris avi& 1984)

    Guinea K, �'3"P'r\

    Illustrated flora of Nordic mosses Nyhlom 1987)

    Mosses and other Bryophytes- An illustrated glossary Malcolm O\a 2000)N�w"S!:NI.J

    How to know mosses and liverworts Conard. 1956)

    A preliminary account of mosses of Zambia Phirl-a-,J 1988)

    Sorne mosses from Kuwait El Saadawl 1976)

    Index nuscorum Van Der Wijk et al (I 969)

    Contributions to the bryophyta of the United Arab Shabbara and El Saadawi

    Emirates (UAE) (I 999)

    Identity of North African endemic bryophytes I Cano et al 2000)

    1 8

  • Contribution to the bryophyte flora of Morocco: The Ros et al.(2000)

    Jbel Toubkal.

    Three new records of Funariaceae from Egypt Shabbara 1999)

    New Synonyms in Grimmia (Grimmiaceae) Munoz 2000)

    A cladistic overview of the Byaceae (usci) based Pederson 2000)

    on orphological and anatomical data and with

    emphasis on the genus Bryurn.

    Continue table (1): The relevant consulted floras and publications

    19

  • B) Anatomical Investh!ations:

    Transverse sections through the stems of the mosses under investigation were

    done using te following technique

  • maximum expansion of the tissues. After that, dewaxing was done by iminersing

    -tlieslid-esin�fn-ire xylene-foirone -i-inu-te-.

    Staining was done by i-nmersing the slides in series of solutions as follows-

    Xylene I for five ininutes, xylene 2 for three ininutes, 95% alcohol for two minutes,

    90% alcohol for two minutes, 70% alcohol for one and a half minute, Haematoxilin

    mier's stain which is prepared according to Mahoney(1973) for seven ininutes,

    ninning water for ten minutes and arnmonified water for one minute. Then a back

    dehydration was carried out and the slides were washed by xylene for two minutes.

    This last step was repeated twice.

    --Mo-untmg-ww-&ne-u&MA:�-P-X--mountant--ne-slide�-were--phwed-4ito-a-

    warm plate for five minutes and were left at roorn temperature overnight. After tat

    the slides were transferred to an oven adjusted at 60' C for one hour (Johansen,

    1940).

    The pen-nanent slides were examined microscopically, illustrations were done

    using the drawing tube, and photographs were also taken for the best sections as

    shown in the whole mount preparation technique.

    21

  • C) Chemotaxonomical Investip-ations:

    For the chernotaxonomy of the studied mosses, the following analyses were

    done:-

    1. Oualitative analysis of the chloroplast i-ments:

    I. I. Extraction:

    The plant tissue was ground in a mortar in the presence of excess acetone

    until all the pigment was released from the tissue. Few amounts of calcium

    carbonate were added to prevent pheophytin fori-nation. Te extract was

    centriffiged and made up to a known volume (Harborne, 1973).

    1 2 Separation of pigments:

    Paper chromatography:

    Chromatography was carried ot on Whatmari filter paper No. I in the

    dark sing petrolew-n ether (B.P.60 - 80' Q - acetone -- n-propanol 90: 0:

    0.45 v/v). The chromatograms were developed for 30 minutes as

    recommended by Amon, 1949). The chromatograms were examined both

    in daylight and U.V. light. Also, for each sample the chromatograrn was

    documented.

    Z. Ouantitative analysis of the chlorophylls:

    2.1. Estimation of the concentrations of the chlorophylls:

    Measurements of concentrations of both chlorophyll a and b were made

    by direct determination of the-absorbances at different wavelengths using

    22

  • standard spectrophotorneter (Jennay 6300). In this method, the absorbance

    was measured at 663 and 645 nrn in I cm cells. The concentrations were

    calculated from the following fon-nula according to Harborne 1973):

    a) Total Chlorophyll (mg/1 = 20.2A645+ 8.02A663

    b) Chlorophyll a (mg/1) 12.7A663 - 2.69A645

    b) Chlorophyll b (mg/1) 22.9A645- 4.68A663

    2.2. Absorption coefficients:

    Scanning at spectral region 400 nin - 700 mn was made and the

    absorption spectrum of each sample was read using UV.

    spectrophotometer. The maximum Lambda for each sample was detected

    and compared with the absorption coefficients of both chlorophylls a ad

    b (Stryer, 1992).

    3. Niti-wen Content:

    Nitrogen contents of the studied mosses were estimated using Kjeldahal

    procedure according to (Ryan et al., 1996):

    1. Sample preparation:

    Plant tissue was firstly ground to a powder which passed trough 0.2nu-n

    sieve before digestion.

    11. Sample digestion:

    0.2-Uams of goun

    grams of a catalyst mxture (96%- anhydrous sodium sulphate 035% copper

    23

  • SUlphate and 0.5% selenium dioxide) and 3ml of sulphuric acid were added and

    digestion was proceeded for two hours. The mixture was allowed to cool before

    addition of 300ml water to the digest and the mixture was allowed to cool again.

    Ill. Distillation and titration

    The content of the digestion flask was transferred to te distillation

    apparatus. Another, 50ml conical flask in wch 15ml 42% boric acid ad 2

    drops of bromocresol green methyl red mixed indicator were added, . placed

    tinder te condenser of the distillation apparatus with the tip of the delivery tbe

    below the level of the liquid. 20ml of 40% sodium ydroxide were added

    careftilly down the neck of the distillation apparatus. Moderate heat was applied

    first, and then increased gently for about 7 minutes. The delivery tube was

    washed down and the ami-nonlum. ion was titrated with 0.02N hydrochloric acid

    to the neutral point of the blue colour just disappeared (one drop in excess will

    turn the solution pink).

    IV. Analysis:

    A ftill analysis was carried out without the plant tissue and the volume of the

    standard acid consumed by the traces of nitrogen in the reagent was measured.

    The nitrogen content of each sample then was calculated using the following

    formula:-

    Nitrogen percentage = A - x 14 x I 0 x N

    -4-OOGA-W4--

    Where: A the volume of hydrochloric acid equivalent to the sai-nple titration.

    B the volui-ne of the hydrochloric acid equivalent to the blank

    titration.

    24

  • N = te Non-nality of te used ydrochloric acid.

    Wt = the weigh of the sample

    4) Elemental Analysis:

    Sodiuin, potassium, zc, calciw-n, cobalt, copper, manganese and iron-- -f - - -- -W -d-asing/1--foreach-- a-i-p -- For-these-

    conce-nTraians were e nne s II le

    detenrlinations te dry ashiig procedure was used Perkin Elmer, 1994).

    Method:

    a) Sample Preparation:

    One grain of dried plant tissue was weighed ad placed 'Hi a

    porcelain crucible ad then placed I a cool uffle furnace ad ashed at

    500'C for overnight. Te ash was cooled and dissolved n of 20%

    hydrochloric acid. The solution was filtered tough a acid- washed

    filter paper into 50nil volumetric flask and diluted to the volume with

    deioiuzed water.

    b) Measurement:

    The quantitative estimation was determined by the atoinic

    absorption spectrophotometer (A. A. S. 3 1 0)

    .5. Statistical anal in

    ysis: Cluster analysis was done accord' g to Sokal ad

    Sneath 1963) using the statistical package SPSS.

    0

  • CHAPTER THREE

    RESULTS

    A. Taxonomical Results

    1. Sudan bryoflora (mosses) checklist:

    In this study the actual number of mosses recorded for Sudan

    Republic was worked out from different literature sources, and moss

    specieswe sumi-narized as a checklist of the bryoflora of Sudan,

    because it is the only country Affica that hasn't been covered by a

    proper checklist.

    Table 2 covers all the mosses species with their literature

    references gving the source, addition to the present study check.

    Taxonomic investigations of the studied specimens showed 12

    different species belonging to different families and 10 different

    genera. Compared to the previous studies, 7 of the identified species

    were already reported for Sudan by different authors. However, still

    all the studied genera are known for Sudan except one (Ceratodon

    Bird.). Refer to (Table 3.

    26

  • Table 2): Sudan bryoflora checklist.

    Present Literature reference

    Species list study

    check

    Anacolia laevisphaera Taylor. Townsend 1984)

    Anoectangium aestimm (Hedw.) Mitt

    Anomobryum julaceum var. julaceum (Bird.)

    Schimp.

    Bryum pseudotriquetrum var. bimum (Bird.)

    Lilij.

    Bryum torquescens Bruch ex de Not.

    Didymodon rigidulus var. gracilis Schleich.

    Fabronia abyssinica Muell. Hal.

    Grimmia laevigata Bird. +

    Mielichhoferia clavata Buch&Schimp.

    Philonolis marchica var marchica Hedw.

    Psedocrossidium replicatum var.replicatum F.

    Weber& 0. Mohr

    Torfula bogosica Muell. Hal.

    Tortula sabinae C.C. Towns

    Syntrichia ruralis subsp. Ruralis (Hedw. F

    Weber&O. Mohr.

    27

  • Continue table 2):

    Funaria hygrometrica var. calvescens + Wickens 1976) &Pettet

    (Schwaegr.) Kindb. (1967)

    Lepiobryum pyriforme (Hedw.) Wilson W ickens(1976)

    -L,��do�nnum viti�ulosoides var� timlosoides

    (P. Beauv.) Wijk&Margad

    Polytrichumjuniperinum Hedw. +

    Tortula khartoumensis A.Pettet - Pettet 1967)

    Barbula unguiculata Hedw. -

    Micropoma nilotica (Del.) Lindb. -

    Funaria nutans Mitt. -

    Bryum kilinggraeffii Schimp. -

    Campylopusflaccidus Renauld&Cardot Frahm ( 1 99 5)

    Brachymenium exile (Dozy&Molk.) Ochi 1972)

    Bosh&Sandelac.

    -- Bryun7 capillare var. capillare Hedw.

    Fissidens crispulus Brid. - Pursel ei al (I 992)

    Molendoe sendineriana var. marchica Hedw. - Sollman 1998)

    Thuidium varians We1w. &Duby - Touw 1976)

    Thuidium gratum (P. Beauv.) A. Taeger - cc

    Bryum cellulare Hook - Abou Salama 1998)

    28

  • Continue Table 2

    Entosthodon muhlenbergii (Turner) Fife. Abou Salaina 1998)

    Hyophda laxitexta J. Frochl. Ex -

    Imam&Ghabbour

    Bryum valparalsense Ther - Arts et al (I 995)

    Fissidens pellucidus var. pellucidus Hornsch. - Bruggeinan 1993)

    Fissidens intramarginalus (Hampe) Mitt. - Bruggeman 1997)

    Fissidens sciophyllus Fo. Ulna(Muell. Hal.) - C

    Trachyphllum inflexum (Harv.) A. Gepp. - Buck(1979)

    Henicodium geniculatum (Mitt.) W. R. Buck - Buck(1989)

    Philonotis dregeana (Mull. Hal.) A. Jaeger - Ghanern et al ( 999)

    Philonotisfiontana Hedw. - 4 4

    Pilonotis hastate (Duby) Wijk& Margad - cc

    Gymnostonium aeruginosum Sm. var. - cc

    cochlearifolium Karczmarz

    Pleurochaele squarossa (Bn'd.) Lindb. + K.U. Herbanwn

    Crossidium davidai Catcheside 66

    Funaria nubica C. Mull +

    Gigaspermum mouretii Corb.

    Tortella humilis (Hedw.) Jenn. +

    29

  • Table (3)- Present study list:

    Species Families

    1. Barbula ehrenbergii (Lor.) Fleisch Pottiaceae

    2. Peurochaete squarossa (B'd.) Lmidb. cc

    3. Tortella humilis (Hedw.) Jenn. cc

    4. Pholonotis longisela Bartraini'aceae

    5. Philonotis tenuis (Hedw.) B'd. C

    -&.--Fiinari-a-hygrome-tr-ica-ffew.� �---- -- F-un

    7. Funaria nubica C. Mull. 4 4

    8. Ceratodon purpureus(Hedw.) Brid. Diffichaceae

    9. Grimmia leavigata (Bri'd.) Bnid. Grfimniaceae

    10. Bryum pseudotriquetrum (Brid.) Shwaegr Bryaceae

    I 1. Thuidiumfurfurosum (Broth.) Broth Thuidiaceae

    12. Poly1richumjuniperinum Hedw. Polytrichaceae

    30

  • 2. Present study taxonomic descriptions:

    The followings are the detailed descriptions of the studied speci

    I I u-nens,

    4-iff -aspe-ai-alrefe-refi-ce t-o-ilye--t-a-co-n-o--ica-Iy- important-characters-wh-icIr-

    are also illustrated figures 6-26).

    2.1. Genus: Funaria Hedw., Sp. Muse., 01

    Species autoicous. Leaves lanceolate, concave, crowded at' top of the

    stein, erect or erect-spreading, margin plane or ncurved, denticulate or

    entire, bordered, cells large and thin-walled; capsule on an elongated seta,

    asymi-netrical pyriform, gibbous; neck conspicuous; calyptra flated,

    cucullate, long-beaked; peristome double.

    * Most species are cosmopolitan.

    -4,-Funaria-hygrometrica-Redw-S p.- Muse-: 172� 18R�

    Plants in dense tufts, often large patches, light green to pale

    yellowish green, 310 mm high. Shoot is bulbiforin. Leaf narrow,

    chlorophyllose and with entired bordered margin; costa extends clearly

    to the tip of the leaf, leaf cells are long hexagonal and papillose- alar

    cells are like the rest leaf cells. Capsules: inclined, fijrrowed and

    smooth; Seta is long, smooth and dark brown colour; Pen'stomes

    contiguous (touching but not overlappi or fused) and with cancellate

    (resembling a net work) apical disk; Teeth are entire, sigmold, regular

    and with hyallne tips. (Figs6&7).

    3 1

  • C)

    6. Funaria hygrometrica Hedw.

    a) Whole plantb) Leaf apical cells (x IC)c) Rhizoids at the base of the stem (x4)

    32

  • b)

    a)

    c

    F 7: Funaria hygrometri

    I:m ca Hedw.a) Closed capsule (x IO)b) Closed capsule (x4)c) Opened capsule showing teeth (x IO)

    -33

  • H Funaria nubica-C. Mull.

    -Plants---m--loos-e---tuff-a,-pate green-4-5-nu-n--hWh-Shoot&-,are--

    prostrate with erect branches; Leaf: broad and rolled upward; leaf

    marg n: smooth and thin-walled; Costa absent; leaf cells are spid e-

    shaped; Alar cells are like the rest leaf cells. Capsules: short and

    111clined; Seta are short and brown-, Peristome is single; Teeth are entire

    (Fig. 89).

    2.2. Genus: Ceratodon Brid., Br. Univ., 1826

    Species dioecious. Leaves lanceolate, with revolute margins; cells on

    the whole leaf homogenous, regularly quadrate above, shortly rectangular

    below, nerve strongly excurrent or ending just below the apex. Capsules on a

    _ptupke or_yelb wish ed seta, erect or inclied, when dry sulcate and furrowed.

    Peristome with basal membrane. Teeth papillose, deeply divided.

    Ceratodonpurpure (Hedw.) Brid., Br. Univ., 1826.

    Plant is erect (1-5) mm. high, green. Leaf. lanceolate and plane;

    Leaf margin is toothed and thin walled; costa is present, ending below

    the apex; leaf cells are elongated; Alar cells are swollen and

    transparent. Capsule: erect smooth and strumulose (having a small

    swelling on one side); Seta is long, erect and purpureous to red-purple;

    Calyptra is smooth and mitrate; Teeth are papillose. (Figs: 10, II, 12).

    34

  • --k

    ONUSAA

    a

    C

    b

    d

    I o, 8: Funaria nubic C Mull.a) Reproductive buds and adventitious rhizolds (x4)

    c) li%) Whole plant>( e) Leaf apical cells (x4O)

    d) Vegetative buds (x IO)

    35

  • 0

    0

    b)

    C)

    Fig. 9'Fitnaria nubica C. Mull.

    a) Inclined capsule (x4).b) Teeth and spores (x IO)c) Teeth and spores (x IO)

    36

  • I

    a) iI

    I , I

    II

    I

    II

    I

    j.''p l-Ceratodonpurpureus(Hedw.)Brid.IIna) Leaf apical cells (x4O prn)b) Whole plant

    37

  • a)

    V, Ceratodon purpureits(Hedw-) Brid.a) Opened capsule (x IO)b) Opened capsule (x4)c) M itrate calyptra (x IO)

    C)

    38

  • mal

    a)

    b)

    C)

    HP 12:Ceratodonpurpureza(Hedw.) Brid.a) Reproductive bd WO) -Ab) Vegctati ve bud (x I )

    go

    O Actinomorphic sem (XI )

    39

  • Syn. Dicranum purpure um

    ��e'cie arecosinopontan.

    2.1Genus:,Tortella Lim'ber., Laubm., 1888

    Species dioicous; Stem erect. simple or branched. Leaves erect.

    erect�spreading or patulous and often recurved when moist, sually

    crisped and curled or incurved and contorted Wn dry. linear-

    lanceolate, acurninate or mucronate; margins plane or incurved, below

    bordered of a row sinooth, elongate cells; niic stow, cii(III i the

    apex or excurrent; cells in upper part of leaf rounded-quadrate or

    hexagonal, strongly papillose. Capsule erect, straight or slightly curved-,

    Calyptra cucullate or mtrate; Peristorne teeth from a very low bsal

    tube, filiform, pupillose. Plants small or up to cm high drk, iht-

    green or often yellowish green.

    Tortella humifiv. (Hedw.) Jcnn,

    Plant is erect, in loose green wfis, 3 cm. high. Lcaf narrow ad

    rolled upward; in is entire and thin-walled; Ca�ta is present

    and stop clearly near the tip of leaf, leaf cells are smooth and

    isodiameffic; Alar cells are small rectangular. Capsule: erect and

    smooth Seta is long, smooth and brown; Calytptra is smooth and

    mitrate; Peristome is single; Teeth are entire, irregular and ciliate.

    (Figs 3 4.

    40

  • a)

    RP

    Fi R. I 3 Tortella humilis (Hedw.) Jenn.a) Leaf apicai cells (x4O)b) Zygom orgp i c stem x I )c) Whole plant

    4

  • eim-AOM,,

    d)

    4 el�

    a')

    all

    C),

    F 14: Toriella humilis (Hedw.) Jenn.

    a) Opened capsule (x IO)b) Whole capsule (x3.2)c) C apsul e teeth (x I d) Mitrate calyptra (x4)e) Opened capsule (x4)

    42

  • 2.4. Genus: Pleurocliaete Lindb. In M. K. V. A. Forch., 1864

    Species dioicous;j florescence allary. Leaves from sheathing base,_,

    lihceplAte" serrate. at a6ex squarrose; nerve strong, excurrent or percurredt;

    cells in the upper pt of leaf small', strongly papi to t t

    border of several rows of thin-walled, yaline cells. Stems ith w

    central strand. Seta elongate; Capsules erect, cylindrical; Calyptra cucullate�

    Peristome with low membrane,, Teeth long filiforin, papillose.

    Pleuroehoetesquarros (Brid.)Lindb.InOfv.K.V.A.Forch,1824.

    Plant is green, prostrate or decumbent with erect branches. Leaf

    lanceolate and rolled upward, serrate at apex; Leaf margin is entire, smooth

    and thin-walled; Costa is present; Leaf cells are swollen and transparent

    (Fig. 5).

    Syn.: Barbula squarossa Brid., Br Univ., 1826

    Torfulasquarrosa D. Not. In Mem. Accad 138

    Parbbula iorfulosa Br. Ew-., 1842

    Torlellasquarossa_Llmpr., Laubrn.. 1888

    43

  • A

    d)--'w77W7

    ho. 15: Plettrochaete suarossa (BrId.) Lindb.

    a) Leaf apical cells (x4O)b) Reproductive bud xlO)c) Whole plantd) Adventitious rhizolds (0)

    44

  • 2.5. Genus: Barbuld fedw., Sp. Musc. 1 15, 1 801

    Plant: dioicous or,! possibly sometimes rhizautoicous forining dense.,.

    cushions' elowish brown o dark green. Stem short to elongate, 23.5 c;

    hyaline. Leaves incurved to weekly spreading, oen contorted 6r7I"iSted`

    about stem, occasionally catenulate when dry, spreading when moist; igulte'

    or broadly lanceolate to long-triangular, adaxial surface usually deeply

    grooved along costa,; margins sometimes narrowly decurrent, occasionally

    plane throughout entire or weakly denticukite ear apex, ,apex rounded to

    obtusely acute, usually mucronate. Costa percurrent to shortly excurrent.

    Perichaetia terminal. Seta 05-2.5 cm.Capsule: stegocarpus. Peristome teeth

    filamentous to narrowly triangular, usually strongly twisted. Calyptr

    cucullate.

    Barbuld erenh (Lor.) Fleisch.

    Plant is prostrate with erect branches, 1-3) cm. igh in dense

    dark green cushions. Leaf broad and slightly curved; leaf margin is smooth

    and thm'-walled; leaf cells are isodiametric; alar cells are swollen and

    transparent; Costa is present and stops clearly near the leaf tip. (Fig: 16).

    45

  • Ar j

    C)

    Fig. 16: Barbula ehrenbergii (Lor.) Fleisch()

    a) Whole plantb) Leaf apical cells (x4O).c) Comites (stiffeners) support the stern (x25)d) Adventitious rhizolds (x4).

    '16

  • 2.6.Genus: Philonotiv Brid.

    Plant dioicous usully,.,erect, sometimes densely caespitose and rhizoid-

    �tomentose-below, Leaveserect secund, ovate-lanceolate, blunt]

    toothed, papillose; nervc strong, often ek ii&Rf,

    ven 1-nicular towards the leaf apex, gradually shorter and wider downward,,'at'

    the base loosely rectangular. Capsule globose striate, when dry furrowed, with

    long seta. Calyptra, mitrate. Peristorne double and badly developed

    I Yhilonods longiye (Michx.) Britt.

    Plant is prostrate with erect branches, 12.5 cm. high in dense

    dark green tufts. Leaf narrow and rolled upward; leaf margin is smooth and

    thin-walled; leaf cells are elongated; alar cells are small and rectangular; leaf

    base stops at the stem; Cost is present (Fig: 17).

    11. Philonotiv tenuiv (Hedw.) Bird.

    Plant is prostrate with erect branches, 1-1.5 cm. igh in loose

    tufts. Stem is brown. Leaf. narrow and rolled upward; leaf margin is tin-

    walled; leaf cells are elongated; alar cells are swollen and transparent; Costa

    is present and stops clearly near the tip (Fig: 18).

    2. 7. Genus: Polytrichum Hedw.,Sy. Musc. 1801

    Species dioicous. Leaves from broad, erect, sheathing gradually

    narrowed into a linear or narrowly ovate-lanceolate, erect spreading; margin

    47

  • a

    !�A, A A-�

    all

    d)C) XN

    A."

    1:j.IT Pholonotis longisela

    a) Whole plantb) Lal'apical cells (x4O)c) Transverse section through stem (x25)d) Young sporophyte covered by perichaetium (x IO)

    49

  • ;�7

    b)

    C) 4'

    Ilk

    v

    g:P

    Philonolis lennis (-ledw.) Brid.

    a) Whole planth) Leaf apical cells (x4O')c) Transverse section showing stern and alar cells (.x25,d) Adventitious rhizolds (x4)

    4 9

  • in. upper part of the leaf narrowly or broadly incurved, dentate or etire; nerve

    4;;DAUTQW4AUqI=IV' 1USIM

    Seta smooth, strong, engate- Capsule erect or.inclined, short or elongate,,-.

    Calatra large, strongly hairyihr6bghput;.Peristome single.

    Pol um iumperinum Hed 1801

    ytrich W. . Muse.

    Plant is erect, 0.5-1.0 cm. high in loose pale reen mats. Leaf-

    filiform and plane; leaf margin is smooth; and thin-walled; leaf cells are

    elongated and smooth; Costa is present; alar cells are swollen and transparent.

    (Fig: 19).

    2. 8. Genus Bryum Hedw., So. Muse., 1801

    Species dioicous, or synoicous; inflorescence terminal, bud-like

    or dish-shaped. Leaves lanceolate, often smaller and distant below, larger and

    crowded in stem apex; margin smooth or denticulate in upper part, border

    usually distinct, but sometimes indistinct or nearly absent, cells in tipper part

    of leaf hexagonal to rhomboidal. Seta mostly reddish, elongate; Capsule erect

    or horizontal to pendulous; Calyptra mitrate; Peristome double

    Bryum pseudo2iquetrum (Hedw.) Shwaegr.

    Plant is prostate with erect branches, 15-2.5 cm. high in dense

    green tufts. Leaf broadly lanceolate and rolled upward; leaf margin is

    bordered; Costa is present; leaf cells are smooth and spindle shaped; alar cells

    50

  • 71C)

    d)

    1-.,o. 9. Polytrichumftiniperinum Hedw.

    a) Whole plant (x4)b) Whole plantc) Leaf apical cells (x4O)d) Transverse section through the stem (x25)

  • are greatly swollen and transparent. Capsule: smooth, erect with brown, seta;

    2j GrimmiaHedw., Sp., Musc..1801..P

    Species autoicous or dioicous, acrocarptis. Leaves lanceolate.

    mostly acunninate and with a long hyaline hair-point, rarely otti§ ad

    without hair-point nerve round or flat at back, percurren; cells in tipper part

    of leaf rounded-quadrate, stnooth or slightly sinose, at margin oen a border

    of hyaline cells. Seta sort or clongatc, straight or arcuate; Capsules sually

    symmetrical, rarely gibbous at base, sometti-nes sulcate when dry and cmptl;

    Peristorne single. Teeth reddish, entire or nTegularly divided and perforated to

    about te iniddle; Calyptra smooth, itrate or calculate, usually long-beaked.

    Grininiia laevigah�.(Brid) Brid., Br. Univ.,1826.

    Plant is prostrate with erect branches, 1-1.5 cin igh in loose

    dark green tufts. Leaf broadly lanceolate and rolled pward; leaf argi is

    toothed; leaf cells are isodirnetric; alar cells are like te rest cells, Costa is

    extends clearly beyond te leaf tip as an awn. (Fig. 23&24).

    Syn.: Grinnnia campewris Burcb. In Hook, Muse. Exot., 1820

    G. leucophaea-Grev. In Wern. Trans., 1822

    Campylopus laevigaluv Brid., Mant. Muse., 1819.

    52

  • �JW

    A"

    V

    (Brid.) Shwacgr

    Fig. 20: Bryumpseudoiriquetrum IIa) Teeth (x4O)b) Whole plantc) Capsule 0)

    53

  • b)

    44

    C)

    4

    aegr2 i3ri-ton seudotriquetrum

    a) Vegctative and reproductive buds (x4),b) Vegetative bud (x I 0)c) Adventitious rhizoids (x IO)

    54

  • At

    a)b)

    7777777rrrM1""'11

    C)

    d)

    F I g. M-yunt pseudotriquetrum (Brid.) Shwaegr

    a) Alar cells (00)b) Alar cells WO)c) Transverse section through the stern (x25)d) Leaf apical cells (00)

    55

  • a)

    Grimmia kj',"-�',igata (Brid.) Brid.Fig. 23:a) Whole plant showing adventitious rhizolds (0)

    b) Whole plant

    56

  • 4,14

    "W"

    C)

    Fig. 24: Grimmia leavigata (Brid.) Brid.

    I - I Ia) Awn and apical leaf cells (x4O)b) Rolled leaves (x4)c) Transverse section through the stern (x2.5)

    57

  • 2. 1 0. Genus: 7tidium P Bru�h t al, Bryol. Eur. 5Fasc. 51) 1852.

    gre

    brown nats. Stems creeping,--to arclied-ascending, lanceolate or filifon�,!V

    -leaves, are more or, less tan lar with a heart-sha e base ad long,

    minate 1as'I I 11acetl strong midrib ends in ti F t"I �6- I "

    P Ali e, exadte e e because.!,-,Io

    one papilla. leaf margin usually revolute, papillos , s rrate abov

    the projecting cell ends. Costa: short, weak, ending below apex. Cells Uniform

    throughout, rounded to oblong, exagonal tick- walled. Capsule inclined to

    horizontal, crved, cylindrical, asymnictrici], sooth. Calyptra: cticidlate,

    Peristorne single.

    77tui(fium furfurosum (Broth.) Broth.

    Plant prostrate with erect branches 15-2 cm. ig i loose pale

    green 1nats. Leaf broad. and plane; leaf inargin is toothed ad thin-walled;

    leaf cells are elongated; afar cells are greatly swollen and transparent; Costa is

    present. (Fig.25&26).

    Syn.: 7huidiopsis carralensis Herzog &Schwabe 1939,T fuifurosuln Herzog

    &Schwabe 1939, T. masqfuerae Brotherus 1924 and Thuidit1117 corralensis

    Dusen 1903

    58

  • C)

    X J:ie. 25 Thuidiumfirfurosum (Broth.) Broth,�)

    a) Whole plantb) Leaf apical cells (x4O)c) Transverse section through the stem (x25)

    59

  • pro Ith� oliwi I �-Iq�IVTW III rV:'flj-

    A

    NO

    r

    Oa)

    Ae,

    4LW

    b)

    F 6:Thuidit(m furfurosum (Broth.) Broth

    a) Young sporophyte covered by perichaetlum (x4)b) adventitious rhizoids (x4)

    60

  • Key to the studied genera:

    A. Capsule with double Penstomes%

    5. QaJyptra -infl

    BB, Calyptra mitrateo

    C. Capsule globose, striate, finTowed when &y.....Philonottv

    CC. Capsule erect or horizontal to pendulous ....... Bryum

    AA. Capsule with single penistome -

    D. Teeth strongly twisted, flamentous and entit'e....Barbilla

    DID. Teeth erectfiliform and papillose-

    E. Leaves lanceolate, serrate at apex ....... Pleurochaete

    EE. leaves lanceolate, entire at apex�,

    F. Seta purple or yellowish red ........ Ceratodon

    FF. Seta brown

    G. Capsule asymmetrical ............... Auidium

    GG. Capsule symmetrical

    H. Calyptra large elongate, strongly hairy throughotit... okiricum

    HH.Calyptra itrate and entire,;

    1. Leaves mostly acun-linate with long hyaline hair-point, often

    u�-'-4k sinose margin ... Grimmia

    11. Leaves linear-lanceolate with si-nooth margins ........ Tortella

    61

  • B. Chemotaxonomical Results

    LI Elemental analysis

    The following are the results of the e'leme�'Ianaysmb:"t:6.I

    studied mosses with special reference to their localities:

    1. Sodium content:

    The results of this analysis show that, the studied mosses are

    approximately similar in their sodium content. The recorded datawe fe,

    show the content range (I 2-18.5 mg/l ) where 12 mg/l,,recorded for

    Polytricum junip&rnum collected from Togi and 18.5 mO, recorded

    for Philonotis tenuis collected from Golol (Table 4 and Fig.27

    2. Potassium content:

    The potassium content of the studied mosses reflects a

    relatively homogenous entity: Barbula ehrenbergii, Pleurochae-le

    squarossa and Tortella humilis which are grouped as Nyrtete

    collection. However, minimum and maximum contents (I 700 and

    62

  • vmm, Ot M,." 411-I& 17, 1e

    ..... .. ...(O co --j 0cn J�lcr

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    CO OD OD U7 CA) CO OD M

    63

  • CDW -4 Cn 4,-W

    co

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    sodium content ng/l

    Tof i

    -4

    CA

    6 4

  • 57.25 mg/l) were recorded for two different species from onegi-'(Tabid-5-arid rg

    -6oI'1[6&fom'groap,'*t ans'To Ti 2S)

    in t eir "Zirit

    The studied mosses are extremely valn'abl h

    contents. The results show maximum zinc content of about 30.175

    mg/l. recorded for Tortella humilis collected from Nyrtete, and

    minimum zinc content of about 4975 mg/l recorded for Girimmia

    laavigata collected from Beldong. Also, wit',in the same collection

    group there are clear variations between different species (Table 6

    and Fig. 29)

    4. Cobalt content:

    cobalt content show the range between

    0.000 and 0.050 rngA which is recorded for four different species

    fi-om two different collection groups: Grimmia lewvigata collected

    fi-om Beldong and Bryum psedotriquetrum, huidium fitrfurosum

    and Polytricum juniperinum collected from Togi, and 0.050 mg/]

    recorded for Philonotis longiseta collected from Golol. However,

    65

  • N) C) --4 ) n w N) CDCO 001 crlo,�i co 0 p �n -n CD

    al

    Q)IC..

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    66

  • CDI<

    p o -n -n CDQ) cn

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    -A d-, s, Z C),Cz

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    Potassium contcnt mg/]tlj w .11, �-A \

    C:) C: c c c c)

    41- intn.

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    (O OD -4 0) (31PO 0 CD

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    68

  • . . . CDCD OD 0) Y -06 CA) N)C)0

    r-4-

    0S -� l

    CDOCO :r (D CDCD E C: (1)CD

    Z :3 cb CDZ :A CL Z3

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    Zinc content mg/I.

    0 Cn C> Ln

    10,

    --4

    co

    co fffiC

    m0.

    69

  • Variation within the species from the same collection group is ver

    5.-Conver content:

    I Copper content show the range between 0044

    mo as minimum record and 0 109 mg/] as maximum records for

    the two species.- Grimmia laevigata and Tortella humilis

    respectively. Also, the analysis shows the maximum records for

    Togi group compared to the others (Table and Fig. 3

    6.-Manganese content:

    Manganese content of the studied mosses rangeA

    between 0637 mg/I recorded for Tortella humilis and 217 mg/I

    recorded for Bryum psedoriquetrum which are different species

    from different localities. Also, it is observed that the three different

    species: Bryum pseudotriquetrum, Thuidium furfurosum and

    Polytricumjuniperinum which are grouped as Togi collection group

    70

  • co 00 -4 0) (31 4�6 Cj N)N) CD

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    Cobalt content mg/I

    O 0 0 0 0 O0 0

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    72

  • 10 00 -4 0) (it ok0 M cr

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    00 C" Ul 00 CA) -4

    73

  • CD 0 4 ) CTI J�k W N)

    CD IZ (D, .1eZ3 t:-, --. , 4.1-,. z,� I a)'.%0.1, I D0-0� :Z. 1. 0; - CZ

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    74

  • show the maximum records compared to the other four groups

    -Conte: t.-

    The iron content of the studied mosses shows minimum

    content of a about 18.525 m/l recorded for Tortella humilis

    collected from Nvrtete and maximum content of about 46.575 m/l

    recorded for Thuidium fitrfurosum collected from Tog]. GenerallyI

    the studied mosses are extremely variable in teir iron content even

    those collected from the sanne localit,%.. (Table I and FP-33).

    11. Qualitative analysis of the chloroplast pigments:

    Acetone extracts of' chloroplast pigimnts were separated on

    Whatman filter paper no. I using petroleurn ether-acetone-n-

    propanol solvent system resulted in a number of spots. Table (I I)

    and Fig. 34) show the chromatographic properties of the separated

    compounds.

    75

  • . .. ..... ...

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    76

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    m

    O M

    is. 'Amm

    77

  • C) (O 0 4 ) Cy - (A) cr

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  • Table I : Chromatographic properties of the separated compounds.0 i

    Plant species compoun value

    J.Bai-hula ehrenbergii 1 0.16

    2 �0.36;

    3 0.77

    2. Plew-ocheatesquarossa 1 0.22

    2 0.44

    3 0.79

    3. Toriella humilis 1 0.14

    2 0.28

    Philonotis longiew 1 0.64

    2 0.86

    Philonolis lenuis 1 0.31

    2 0.53

    3 0.73

    4 0.86

    80

  • 4&i Z�winwo,�,,,fable VT,

    Separated Rf.

    Plant species compound value

    Grimmia leavigata 1 0.52

    2 0.74

    Bryum pseudofriquetruin

    71iuidiwnfiu�furosum 1 0.43

    2 0.84

    Poly1ricunijuniperinum 1 0.35

    2 0.57

    3 0.74

    4 0.98

    8 1

  • - ---------

    J

    1. Barbu thmememrr-'i 2.Pleurochaele 5Mqr-rO.3a 3. Torlella limits

    AP

    4.,,Philonotis loupLse 5. hilonotis_Lenuis ometrica

    6 Funaria hy&�-..

    Fig. 34- Paper chromatograrns of acetone extracts. Solvent systern:petroleurn ether acetone n-propanol ( 90 : 10 45, v/v).Viewed in day

    light.

    82

  • Finaria nubica 8. Ceralodon purpureus 9. Grimmia laevigata

    IO. Bryum pseudotriquetrum IL Thuidiumfurfurosum 12.Polytrichumfitniperinum

    Contlnu'c Fg. 34

    83

  • M. Quantitative analysis of the chlorophylls:

    ,A). Chlorophyll con&ntrations:

    The chlorophyll contents of the studied mosses sh6w;' difY&dAfI.

    concentrations for both chlorophylls a and b. Studied mosses are

    greatly varied in their chlorophylls which record maxin-iUrn total

    content of about 40.815 rng/l for the species Philonotis lenuis and

    minimum total content of about 6822 mg/I for the species Funaria

    hygrometrica. Table(12) and Fig. 35) show the concentrations of

    the total chlorophyll, chlorophyll a and chlorophyll b for each

    species.

    B). Absorption coefficients:

    Absorption spectra of acetone extracts for te studied

    species are demonstrated in Fig. 36).

    94

  • INSmliy,is, cont e'n't-,,ofthcstttdie&m'(S�vs.

    Cincinnati, Total' h`y,14iN0cmO*mchlo h' 11 a mz4

    mg/I

    1. Barbula ehrenbergii 14.228 9.215 5.016

    2. Pleurochaete squarossa 25.229 16.897 8.408

    3. Tortella humilis 11.168 7.282 3.880

    4. Philonotis longesita 24.353 16.069 8.330

    5. Philonotis tenuis 40,815 15.808 25.020

    6 unaria hygrometrica 6.882 4.005 2.517

    7. Funaria nubica 19.567 12.745 6.826

    8. Ceratodon purpureus 10.862 7.143 3.720

    9. Grimmia laevigata 13.315 8.170 5.118

    1 0. Bryum pseudotiquetrum 10.638 6.581 4.058

    11. Thuidimfurfurosum 18.742 11.891 6.497

    12. Polytricumjuniperinum 12.829 11.646 j1.185

    85

  • chloroohylls contents

    W 0 Cn Lon

    J-:1f

    ehrembergil

    2.Pleurochaetssquarossa

    3.Tortella humilis

    4.Philonotis longesb

    C.25. Philonotis tenuis.

    6.Funariahygrometrica =r

    7.FunaHa nubicam

    B.Ceratodon 0purpureus

    m

    9.Gdmmia aevigatamCL

    =1

    pseu tum 0mCA

    II.Thuidimfurfurosum ...

    12.Polytdcumjumpennum

    0 0 --I

    O:r0

    33

  • WAV al

    &Orhl�la

    2.Pleurochaele, Mgr-Osa

    3. Torfello, humilis

    14NI 7c-

    4. Philonotis Ion i-sela

    36.0-12) QN. Absorption spectra ofchlorophylls a and b.X- xis Wvelength (nm) Yaxis- Absorpt'on oell"cients (m l)

    8.7

  • 6 tMaria h� e�lnca

    I5. Philontfi-S Initially

    . O

    14

    Saint v

    4.00 50. 6oo 700

    8. Ceratodon purpureus

    see

    7.fv!�aria 17"hica

    4�

    9. Grimmia leoxigala

    0- 600

    300 1100 500

    colitilILIC 36.

    88

  • W14, ��,l

    ONI

    lon 660

    10. Bryumpseudotriquetrum

    3-

    II. AuidiumArfurosum

    3� too*

    12.Polyfrichumjuniperinun?

    Contillue Fig. 36-

    89

  • IV. NitroLyen content

    The minimum and, the maxim .um. percentage nitrogen woreen 7Zorte

    1(L&-v���x d' ��,'�"hiii)iiftswte

    Uto k" d te moSseTh6�' nitrogen content range; d

    3.22%.Gololl, Beldong and Togi collection groups show the low est

    variation within species in their nitrogen content,- Beldong

    collection group shows the lowest record compared to the two other

    groups. Table 13) and Fig. 37) show the percentage nitrogen for

    the studied mosses.

    90

  • co 00 -4 0) Cil 0h CA) N) crCD

    Book,zlli 70 -o -1 IRWINY :Y -a v

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    91

  • (D OD -4 a) Cri W N)

    pot it ii4i 00 �4 Z3 CD Z Zk a (bz -'j T--- zz: ZL CD

    (b -Q) CJ) 0 0 CD

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    (D 23U)

    Nitrogen

    7L(A CA) (n

    CL

    0

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    92

  • C. Numerical taxonomy

    The data of the previous quantitative investigations of sodiukn,.

    m- t6w t COPMr"i1----rnR71gvffeSe!I C Mvp

    nitrogen which is computed for area under curve and tonIP16fe'

    linkage dendrogram. resulted into 10 different groups. The following

    is the arrangement of the groups starting firom the group that

    includes the most closely related individuals (Fig.38)-

    Group ()- a) Barbida ehrenberg,,

    b) Thwdiumfitrfitrosuni

    c) Grimmia lbwi��qata

    Group (2)- a) F unaria nubica

    b) Polytricumjuniperinum

    Group 3): a) Tortella humilis

    b) Ceratodon purpureus

    93

  • Group 4): a) Thuidiumfurfurosuni

    O"Philonotis longiseta

    Group (5): a) Funaria nubica

    b) Polyti-icunijuniperinum

    c).fftnaria hygrometrica

    Group (6): a) Pleurochaelesquarossa

    b) Philonotis tenuis

    Group (7): a) Pleurochaute squarossa

    b) Philonotis tenuis

    c) Grimmia Imvigata

    Group (8)- a) Philonotis longiseta

    b) PleurochaWe squarossa

    94

  • c) Philonolis lenuis

    --d�--Fitnaria-mtbica-

    01�tricumjuniperinum

    Group (9)' a) Tortella humilis

    b) Ceralodon Furpo-rens

    c) Brytim psedotriquetrum

    Group 10)- a) Ph4lonotis tenitis

    b) Funaria nubica

    c) Polytricumjuniperinum

    d) Funaria hygrometi-ica

    c) Tortella humilis

    f) Ceratodonpc4rvc-lreus

    95

  • A.

    pipT - . ........

    Opost �F.�

    t.W A

    4.?hi-1jr_nod.v Ion isela -Ihdonofi-� mis -Yrica6. Funaria hygome

    7. rinqrLy mibica 8. Ceralodon urpurens 9. Grimmia7aevigala

    Ak "NJA

    IO. Brympseudwriepielnim 1. 7huidiumfurfurosum

    Rescaled Distance Cluster Combine

    C A 0 5 10 15 20 25Label Num ----------------------------------- ---------------

    9

    4

    2

    5

    12 T3

    PI

    10Area Linder CUrve and complete linkage dendrop-rarn Co r

    111C StUdled mosses.

    96

  • CHAPTER FOUR

    ffiscu-SSION"

    the .M.Wial-rimpmUnce of JebehiMarraas-i habitat --for-btybph �Iftsildy!

    Darfiir State stems from the fact that this high volcanic massif gives rise.t6:..

    better soils, lower temperatures, and higher rainfall tan the rest of the region

    of west Darfur. Moreover, the degree of slopes. soil characteristics. rainfall

    conditions, and the nature of erosion arc closcIv related to te requirements of

    these plants.

    The selected sites of te studN.- in Jebel Marra are verv rich in mosses ill

    terms of densitv., he fijlly dentified entities are 12 7 of which are already

    In Sudan by other athors. And for Jebel Marra only 4 species are

    quoted in the previous studies, these are: Funaria hygrometrica, Grimmia

    leavigara, Bryum pseudotriquelruni, and folytrichwn juniperininn. This

    foundation is probably possible because both of t precious authors:

    (Wickens, 1976) and (Townsend, 1984) did ot qote te sarne taxa ad

    Townsend, 1984) stated that, there will be further species to be od in

    Jebel Marra. The rest 3 species: Pleurochaele squarrosa, Torlella humilis,

    and Funaria nubica were reported for Erkowit which of slightly sli-nilar

    environment to Jebel Marra.

    97

  • The genus Funaria wich is recorded for Jbel Marra, Erkowit,

    .'..Khattoum.,.,and-iebeT..T$�- seems to.-Ibe--the�most;- idelv-distributed-Ionei Xkas;;

    remark confirins the description of many consulted floras for this genus*sd

    widespread.

    (;igwspermium sp. which is identified for Erkowit in 1994 bv WtliAfn�

    R.Buck, the curator of bophytes of the New York Botanic Garden is

    reported as of a particularly interest since it is a rare Mediterranean species

    known from Morocco. Palestine, Spain, and te Canary islands.

    The newlv recorded species: Barbula eherenhergii, Philonoliv lenuis,

    Philonolis longiseta, Ceratodon purpureus and Auidiumfiu�furoswn are from

    the alreadv known genera except Ceraiodon which is reported for Sudan for

    the first time.

    More four entities which were collected by Wickens 1976 bt not

    identified to the specific level can be regarded for Sudan tese are:

    Brachymenium sp, Bryumsp, Derpanocladus sp ad Physcomitriumsp.

    It can be concluded tat, in addition to the newlv identified entities, this

    research gathers the scattered information about the bryoflora checklist of

    Sudan Republic in an easy followed foundation giving a base to which

    researchers can add or subtract as required.

    98

  • .Mosse�s-:17-v-es-tig-aEcT,1'6r- eir.-eleffiFifa ..-coiistituents..,were..,wiaimtediia&4-';1

    the dorninant species in pach study site, and as it is followed in this stud

    TrIost..,,,methods n heavy 'metals monitoring employ bryophyte§ as

    bioaccumLilators and invol%--e sample collection followed by laboratory

    analysis technique to detect actual levels. Piipo 1982) employed epiphytic

    bryophytes as climatic indicators in Eastern Fennoscandia. Also, Kansanen

    and Venetvaara 1991) compared the capability of two mosses, an epiphytic

    lichen, pine bark samples, pine necdle litters, earthworins, and moths in

    assessing airborne chromium and nickel dust near a ferrochrome and stainless

    steel works in Finland. The two mosses and the lichens were the most

    effective biornonitors at low and moderate depositions

    Part of the result of this study also agrees with that of Mukhe 'ee and

    Nuorteva. 1994) who found that the best indicators for zinc and aluminum are

    mosses and epiphytic lichens.

    The records of the levels of te investigated elements will bettbase for

    future comparisons so as to estimate the atmospheric pollution degree in Jebel

    Marra as it is followed now in many countries such as the Republic of Karelia

    (Fedorets, 1998).

    The statistical analysis for all chemical investigations grouped different

    species from different localities as closely related individuals. This important

    99

  • result confirms the fact tat mosses are good bioaccumulators particularly for

    retil cono�ntrations-!�refiect.-- -deposit.-i-on46witlmiv44.t4v*.v-

    complication of additional-!uptake via root system. (Longton, 1980). Then th&,-

    chemical constituents of the ddibihant species in each site reflect the chemical,,

    constituents of the soil and air in Jebel Marra. However, within the studie 'd

    sites, Golol shows two chemically related individuals: Philonotis 'Ienu'is"a'6d

    Funaria hygromeirica coming in one group. Also, tis study grouped

    (Funaria nubica and 1"unaria hygroinefrica) and (Philonolis Winds and

    Philonotis longisela) each two in one group wich is taxonomically tnie.

    As a conclusion: since Philonotis tenuis and Peurocheale squarossa)

    were grouped together 3 tirries and (7"huidium fiu�furosurn and Grimmia

    leavigata) were grouped together 2 tirries then (Nyrtete and Golol) ad (Togi

    and Beldong) are of relatively the same chemical constituents ptting i 1-nind

    that Togi and Beldong are the highest study sites whereas Nyrtete and Golol

    are the lowest study sites.

    The chloroplast pigments. specially the clorophyll content of the

    studied mosses, reflect considerable variations that can partly help in

    describing them. Taxonomically the studied mosses were described either as

    light or dark green in most of the consulted relevant floras and chemically,

    mosses described as light green recorded the low chlorophyll content and tat

    described as dark green recorded the high clorophyll content. So it can be

    concluded that this part of te chernical analysis not only on line with but.

    100

  • also confirms morphological descriptions and also agrees with authors of the

    t6yf9dw Iota,

    101

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