VOLUME 37 /// 2014 DURBAN•NATURAL•SCIENCE•MUSEUM•NOVITATES
EDITOR
D.G Allan Curator of Birds Durban Natural Science Museum P.O. Box 4085 Durban 4000, South Africa e-mail: [email protected]
EDITORIAL BOARD
Dr A.J. Armstrong Animal Scientist (Herpetofauna & Invertebrates) Ezemvelo KZN Wildlife
G.B.P. Davies Curator of Birds Ditsong National Museum of Natural History
Dr L. Richards Curator of Mammals e-mail: [email protected]
Prof. P.J. Taylor School of Mathematical and Natural Sciences University of Venda
Dr K.A. Williams Curator of Entomology e-mail: [email protected]
SUBSCRIPTION DETAILS
Librarian Durban Natural Science Museum P.O. Box 4085 Durban 4000, South Africa e-mail:[email protected]
COVER IMAGE
Cape grass lizard Chamaesaura anguina Photo: Johan Marais
Published by the Durban Natural Science Museum 1 EDITORIAL Durban Natural Science Museum Novitates 37
EDITORIAL SOUTH AFRICAN NATURAL SCIENCE MUSEUM JOURNALS - THE PAST 50 YEARS (1964 - 2013)
GOING DOWN SWINGING OR ADJUSTING TO THE RIGHT FIGHTING WEIGHT?
This editorial briefly reviews publication trends in the scientific component (Fig. 1). Four of these are national museums: Ditsong journals published by South African museums that are solely or National Museum of Natural History (Pretoria), National Museum – partially focused on the natural sciences. It is based on an oral Bloemfontein, KwaZulu-Natal Museum (Pietermaritzburg) and Iziko presentation given at the conference of the International Council of South African Museum (Cape Town). Five are provincial museums: Museums – South Africa (ICOM-SA) in August 2014. The conference McGregor Museum (Kimberley) and a cluster of four museums was hosted at the Durban Natural Science Museum Research Centre (making up the ‘Eastern Cape Provincial Museums’) in the Eastern during 26 - 27 August and its theme was “Museum research in South Cape Province: Amatole Museum (King Williams Town), East London Africa – relevance and future”. Museum, Albany Museum (Grahamstown) and Port Elizabeth Museum/ Bayworld. The tenth museum is lodged at local government level: the Durban Natural Science Museum.
South African natural science museum journals Table 1 summarizes details of the various journals that have been produced by the relevant natural science museums in South Africa over time. The Ditsong National Museum of Natural History and its predecessor the ‘Transvaal Museum’ has been responsible for the publication of two formal scientific journals. Its ‘Annals’ began in 1908 and continues to date, albeit with a change in name in 2011. Its more lengthy ‘Memoirs’ began in 1943 and was renamed as ‘Monographs’ in 1983. The most recent edition of the ‘Monographs’ was produced in 2009. The KwaZulu-Natal Museum and its predecessor the ‘Natal Museum’ also produced an ‘Annals’ series that began in 1906 and which continued, with a name change in 1910, until 2000. In the last-mentioned year it was subjected to a further name change and a concentration in focus, as African Invertebrates, that continues to date. The Durban Natural Science Museum and its predecessors the ‘Durban Museum’ and ‘Durban Natural History Museum’ is another institution that has produced a series of ‘Annals’ starting in 1914. This Fig. 1. The 10 museums in South Africa that are either solely devoted to the natural sciences or that have a strong nature-science component: national publication was renamed as the Durban Museum Novitates in 1952 museums: Ditsong National Museum of Natural History (Pretoria), National and again as the Durban Natural Science Museum Novitates in 2008. It Museum – Bloemfontein, KwaZulu-Natal Museum (Pietermaritzburg) and Iziko continues under the latter name to date. South African Museum (Cape Town); provincial museums: McGregor Museum The National Museum - Bloemfontein initiated its ‘Navorsinge’ in (Kimberley) Amatole Museum (King Williams Town), East London Museum, 1952 and these continue to date. This museum also began its series of Albany Museum (Grahamstown) and Port Elizabeth Museum/Bayworld; and local lengthy ‘Memoirs’ in 1960 and these continued until 1989. museums: Durban Natural Science Museum. The Albany Museum produced a series of ‘Records’ from 1903 to 1935. Subsequently the five provincial museums that occurred in the South African natural science museums former ‘Cape Province’ (McGregor, Amatole, East London, Albany There are 10 museums in South Africa that are either solely devoted and Port Elizabeth/Bayworld) were served by the ‘Annals of the Cape to the natural sciences or that have a strong natural-science 2 EDITORIAL Durban Natural Science Museum Novitates 37
Table 1. A summary of the various journals produced by the relevant natural science museums in South Africa over time. See main text for full details.
MUSEUM JOURNAL START END Ditsong National Museum Annals of the Transvaal Museum 1908 2009
of Natural History Annals of the Ditsong National Museum of Natural History 2011 Present
Memoirs of the Transvaal Museum 1943 1979 Monographs of the Transvaal Museum 1983 2009
KwaZulu-Natal Museum Annals of the Natal Government Museum 1906 1909 Annals of the Natal Museum 1910 2000 African Invertebrates 2001 Present
Durban Natural Science Museum Annals of the Durban Museum 1914 1947 Durban Museum Novitates 1952 2007 Durban Natural Science Museum Novitates 2008 Present
National Museum - Bloemfontein Memoirs of the National Museum 1960 1989
Navorsinge van die Nasionale Museum, Bloemfontein 1952 Present
Eastern Cape Provincial Museums:
Albany Museum Records of the Albany Museum 1903 1935
Port Elizabeth Museum/Bayworld
East London Museum
Amatole Museum
(formerly McGregor Museum) Annals of the Cape Provincial Museums (Natural History) 1979 1997 Annals of the Eastern Cape Museums 2000 Present?
Iziko South African Museum Annals of the South African Museum 1898 2004 African Natural History 2005 Present
Provincial Museums (Natural History)’ that began in 1979 and continued Museum, Bloemfontein, others have done so in the past, e.g. the Annals until 1997. In 2000 this series was revived as the ‘Annals of the Eastern of the Natal Museum, and some have never included articles on the Cape Museums’ (now servicing only the Amatole, East London, Albany humanities, e.g. the Durban Natural Science Museum Novitates and its and Port Elizabeth/Bayworld museums), which apparently continues to predecessors. For the purposes of this review, all articles related to date, although the last edition was in 2011. the humanities have been entirely excluded from consideration. The Iziko South African Museum was the first natural science museum in the country to boast a scientific journal. Its ‘Annals’ was Overall contributions by discipline initiated in 1989 and continued until 2004, when it was renamed Over the 50-year period, some 2238 articles covering the natural African Natural History. It continues under this title to date. sciences have appeared in these journals, and these articles span a total of some 62 821 pages (Fig. 2a & b). Clearly, the biggest overall Methology contributors to South African natural science museum journals are The period covered by this review comprises the 50 years 1964 – invertebrate biologists and, to a lesser extent, palaeontologists. These 2013. The methodology at the core of this review was to determine two disciplines have contributed over three-quarters of the total the number of articles published, and the number of pages spanned by in terms of both the number of articles published and the number these articles, in South African natural science museum journals over of pages spanned, with the contribution of invertebrate biologists this time period. The information was further examined at a finer alone amounting to over half of both respective totals. Within the scale related to the various disciplines within the natural sciences broad category of invertebrates, entomology has featured slightly and by the five individual 10-year periods within the overall 50- more prominently than other invertebrate groups. Relevant to the year period of relevance. Some of the relevant journals also include latter group, it might have been valuable to differentiate between coverage of the humanities, e.g. the Navorsinge van die Nasionale contributions addressing finer taxonomic levels within this group, Durban Natural Science Museum Novitates 37 EDITORIAL 3
1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 TOTAL 74 201 74 30 44 TOTAL 3450 8479 3516 1970 1090 Entomology 66533Entomology 552 141 313 209 36 Other invertebrates 31 74 30 7 13 Other invertebrates 1434 3015 1206 404 345 Palaeontology 32 107 37 18 27 Palaeontology 992 4893 1963 1329 678 Figs 2a & b. The number of articles published (2a) and the number of pages spanned byMammalogy these articles (2b) in29101 South African natural science museum journals over Mammalogy 29 209 11 0 31 the 50-year period 1964 – 2013 by individual discipline. The numbers in brackets after theOrnithology name of each discipline22110 represent the number of articles published and the Ornithology 285 135 23 10 0 number of pages spanned for each discipline respectively. Herpetology 13010Herpetology 158 86 0 18 0 perhaps especially between marine and terrestrial forms, but this has 250 9000 not been pursued here. Similarly, it might have been useful to have 8000 differentiated the articles covering entomology into the different 200 7000 TOTAL TOTAL insect orders where relevant but examination at this level of detail 6000 150 Entomology Entomology was not attempted. 5000 Other invertebrates Other invertebrates 4000 Turning to other disciplines, among the vertebrates mammalogy, 100 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003Palaeontology 2004-2013 No. of pages of No. Palaeontology
No. of articles of No. 3000 ornithologyTOTAL and herpetology74 have 201 all featured 74 fairly 30prominently. 44 TOTAL 3450 8479 3516 1970 1090 Mammalogy Mammalogy Ichthyology,Entomology however, 66533is relatively poorly represented and Entomology50 552 141 313 209 36 2000 Ornithology 1000 Ornithology contributionsOther invertebrates from two 31 other natural 74 science 30 disciplines, 7 earth 13 Other invertebrates 1434 3015 1206 404 345 Palaeontology 32 107 37 18 27 Palaeontology0 992 4893 1963 1329Herpetology 678 0 Herpetology sciencesMammalogy and botany, are29101 particularly scant. For this reason, ichthyology, Mammalogy 29 209 11 0 31 earthOrnithology sciences, botany22110 and the disciplines represented by the Ornithology 285 135 23 10 0 ‘miscellaneous’Herpetology category13010 are not considered further in this review. Herpetology 158 86 0 18 0
Trends250 at individual museums over time 9000 8000 Iziko South African Museum 200 7000 The trend at Iziko, represented by the Annals of the SouthTOTAL African TOTAL 6000 150 Entomology Entomology Museum and its successor African Natural History, shows that overall 5000 Other invertebrates Other invertebrates publication peaked markedly in the ten years 1974 – 1983 both in 4000 100 Palaeontology No. of pages of No. Palaeontology termsarticles of No. of the number of articles published and the total number of 3000 pages spanned50 (Fig. 3a & b). Since then there has been aMammalogy monotonic 2000 Mammalogy decrease in the number of pages published and a similarOrnithology pattern is 1000 Ornithology 0 0 evident in the number of articles produced, although theHerpetology latter picked Herpetology up marginally in the final ten years examined. Palaeontology and invertebrates other than insects have predominated as the disciplines of primary interest, with the remaining disciplines under examination here, i.e. entomology, Figs 3a & b. The number of articles published (3a) and the number of pages mammalogy, ornithology and herpetology being poorly represented. spanned by these articles (3b) in the scientific journals published by the Iziko The marked peak in publication in the ten years 1974 – 1983 is South African Museum for each of the five 10-year periods spanning 1964 – driven by publications on palaeontology and invertebrates other than 2013, in total and for each discipline individually. insects, with little evidence of any marked peaks relevant to the other Iziko, this monotonic decrease also applies to the number of articles disciplines. produced. Entomology has predominated as the discipline of primary Eastern Cape Provincial Museums interest, with the remaining disciplines under examination here being The trend at the Eastern Cape Provincial Museums (Amatole, East relatively poorly represented. Similar to Iziko, the peak in publications London, Albany, Port Elizabeth/Bayworld, and partially McGregor), in the ten years 1974 - 1983 is selective in terms of discipline and in represented by the Annals of the Cape Provincial Museums and its this instance is driven solely by publications on entomology. successor Annals of the Eastern Cape Museum, as with Iziko, shows that overall publication peaked in the ten years 1974 – 1983 both in terms National Museum – Bloemfontein of the number of articles published and the total number of pages Publication peaked later at the National Museum – Bloemfontein, spanned (Fig. 4a & b). Again as with Iziko, since then there has been a represented by the Navorsinge van die National Museum, Bloemfontein monotonic decrease in the number of pages published, but, unlike 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 TOTAL 27 39 27 14 5 TOTAL 751 2134 1015 446 120 Entomology 4 23 19 9 4 Entomology 535 1661 647 316 93 Other invertebrates 53011 Other invertebrates 29 24 0 10 27 Palaeontology 10020 Palaeontology 800870 4 EDITORIAL 1964-1973 1974-1983Durban 1984-1993 Natural Science 1994-2003 Museum 2004-2013 Novitates 37 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 Mammalogy 65210 MammalogyTOTAL 8256 181 89 292 116 8 65 0 105 TOTAL 2079 2525 2448 1490 1242 Ornithology 63300 OrnithologyEntomology 69 25 160 35 35 55 0 37 0 41 Entomology 1266 1162 1156 1031 642 Herpetology 55310 HerpetologyOther invertebrates 2855734 108 41 25 0 Other invertebrates 70 32 51 10 25 Palaeontologycompared with Iziko 9 and the 18Eastern Cape, 26 and entomology 18 41 and the Palaeontology 383 598 427 122 334 45 Mammalogy2500 13 16 13 3 6 Mammalogy 283 324 585 291 65 40 Ornithologystudy of other invertebrates,262211 as well as mammalogy, have all featured Ornithology 25 257 127 22 148 35 Herpetologyrelatively2000 strongly at 2 different 9 times. 13 2 2 Herpetology 52 152 102 14 28 TOTAL TOTAL 30 Entomology 1500 Entomology 25 140 3000 Other invertebrates 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 20 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 Other invertebrates Palaeontology 1000120 2500
No. of pages of No. Palaeontology TOTAL 27 39 27 14 5 TOTALarticles of No. 15 751 2134 1015 446 120 Entomology 4 23 19 9 4 Entomology 535 1661 647 316Mammalogy 93 100 TOTAL TOTAL 10 500 Mammalogy 2000 Other invertebrates 53011 Other invertebrates 29 24 0 10 27 Entomology Entomology 5 Ornithology 80 Ornithology Palaeontology 10020 Palaeontology 800870 1500 0 Herpetology 0 Other invertebrates Other invertebrates Mammalogy 65210 Mammalogy 821964-1973 181 1974-1983 292 1984-1993 8 1994-2003 0 2004-2013 60 1964-1973 1974-1983 1984-1993 1994-2003Herpetology 2004-2013 Palaeontology TOTAL 56 89 116 65 105 TOTAL 2079 2525 2448 1490 1242 pages of No. Palaeontology No. of articles of No. 1000 Ornithology 63300 Ornithology 69 160 35 0 0 40 Herpetology 55310 HerpetologyEntomology 28 25 108 35 41 55 25 37 0 41 Entomology 1266 1162 1156 1031Mammalogy 642 Mammalogy Other invertebrates 55734Other invertebrates20 70 32 51 10 25 500 Ornithology Ornithology Palaeontology 9 18 26 18 41 Palaeontology 383 598 427 122 334 45 2500 0 0 Mammalogy 13 16 13 3 6 Mammalogy 283 324 585 291Herpetology 65 Herpetology 40 Ornithology 262211Ornithology 25 257 127 22 148 35 2000 TOTAL Herpetology 2 9 13 2TOTAL 2 Herpetology 52 152 102 14 28 30 Entomology Entomology 25 1500 Other invertebrates 140 3000 20 Other invertebrates 1000 Palaeontology 120 No. of pages of No. Palaeontology No. of articles of No. 15 2500 Mammalogy TOTAL 10 500100 Mammalogy TOTAL 2000 5 Ornithology Entomology 80 Ornithology Entomology 0 Herpetology 0 HerpetologyOther invertebrates 1500 Other invertebrates 60 1964-1973 1974-1983 1984-1993 1994-2003Palaeontology 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 No. of pages of No. Palaeontology
No. of articles of No. 1000 TOTAL 40 12 26 49 39 23 TOTAL 173 1904 2019 979 535 Mammalogy Mammalogy Entomology20 0 6 11 3 1 Entomology500 0 786 701 106 17 Other invertebrates 6 5 11 20Ornithology 13 Other invertebrates 71 100 375 468 323Ornithology PalaeontologyFigs 4a &0 b. The number54612 of articles published (4a) and the numberHerpetology of pages Palaeontology0 72 107 197 10 38Herpetology Mammalogyspanned by these articles 0 (4b) in 5the scientific 11 journals published 7 by 0 the Eastern Mammalogy 0 525 498 167 0 OrnithologyCape Provincial Museums02741 for each of the five 10-year periods spanning 1964 –Ornithology 0 175 172 76 16 Herpetology2013, in total and for 14346each discipline individually. Herpetology 30 211 76 152 141
60 Figs2500 6a & b. The number of articles published (6a) and the number of pages spanned by these articles (6b) in the scientific journals published by the Ditsong 50 National2000 Museum of Natural History for each of the five 10-year periods TOTAL TOTAL 40 spanning 1964 – 2013, in total and for each discipline individually. Entomology 1500 Entomology 30 Other invertebrates Other invertebrates 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 Ditsong National Museum of Natural History 1000
Palaeontology pages of No. Palaeontology TOTAL 12 26 49 39 23 TOTAL articles of No. 20 173 1904 2019 979 535 Like at Bloemfontein, publication also peaked later at Ditsong, Entomology 0 6 11 3 1 Entomology 0 786 701 106Mammalogy 17 Mammalogy 10 represented500 by the Annals of the Transvaal Museum and its successor Other invertebrates 6 5 11 20 13 Other invertebrates 71 100 375 468Ornithology 323 Ornithology Palaeontology 54612Palaeontology 72 107 197 10 38 Annals of the Ditsong National Museum of Natural History, and the 0 Herpetology 0 Herpetology Mammalogy 0 5 11 7 0 Mammalogy 0 525 498 167 0 Memoirs of the Transvaal Museum and its successor the Monographs of Ornithology 02741Ornithology 0 175 172 76 16 the Transvaal Museum, compared with Iziko and the Eastern Cape (Fig. Herpetology 14346Herpetology 30 211 76 152 141 6a & b). This peak occurred in 1984 - 1993, in terms of the number
60 2500 of article published, although the number of pages spanned was marginally higher in the previous ten years 1974 - 1983. The number 50 2000 of articles published decreased in the ten years after 1984 - 1993 TOTAL TOTAL 40 but it increased subsequently during 2004 - 2013, although the total Entomology 1500 Entomology number of pages spanned continued to decrease, i.e. the most recent 30 Other invertebrates Other invertebrates 1000 period has seen an increase in the number of articles produced but
Palaeontology pages of No. Palaeontology
No. of articles of No. 20 the average length of these articles has been relatively short. Mammalogy Mammalogy 10 500 Ornithology Ornithology As with Bloemfontein, the representation by discipline has 0 Herpetology 0 Herpetology been more evenly spread at Ditsong than at Iziko and the Eastern Cape. Entomology has dominated consistently overall but there has also been a fair representation of palaeontology, mammalogy and ornithology. Figs 5a & b. The number of articles published (5a) and the number of pages spanned by these articles (5b) in the scientific journals published by the Durban Natural Science Museum National Museum – Bloemfontein for each of the five 10-year periods spanning The trends shown by the Durban Natural Science Museum scientific 1964 – 2013, in total and for each discipline individually. publications are distinct from those shown by the other museums discussed so far (Fig. 7a & b). The number of articles published by and the Memoirs of the National Museum, than at Iziko and the Eastern the Durban Natural Science Museum during the relevant period, Cape, i.e. in the ten years 1984 – 1993 (Fig. 5a & b). Like these other represented by the Durban Museum Novitates and its successor the institutions though, publication subsequently decreased in terms of Durban Natural Science Museum Novitates, shows a slight peak during both articles printed and the number of pages spanned. the period 1974 - 1983 with a decrease subsequently, although with There has not been the consistent and pronounced domination marked variation. The pattern shown by the number of pages spanned by one or two disciplines in terms of publications at Bloemfontein is quite different, showing a peak in the earliest period (1964 - 1973), 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 TOTAL 49 76 33 66 35 TOTAL 1200 986 510 693 883 Entomology 1 13 2 3 2 Entomology 5 93 44 35 423 Other invertebrates 26161Other invertebrates 14 27 16 38 19 Palaeontology 106133Palaeontology 3 0 94 221 32 Durban Natural Science Museum Novitates 37 EDITORIAL 5 Mammalogy 0051611Mammalogy 0 0 53 162 94 Ornithology 45 51 16 24 17 Ornithology 1178 795 246 196 268 Herpetology 06341Herpetology 0 71 57 41 47
80 KwaZulu-Natal1400 Museum The trends shown by the KwaZulu-Natal Museum scientific 70 1200 60 publications, represented by the Annals of the Natal Museum and its TOTAL 1000 TOTAL 50 Entomology successor African Invertebrates, are also distinct, andEntomology markedly so, from 800 40 Other invertebrates those shown by all the other museums discussed soOther far invertebrates (Fig. 8a & b). 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 600 30 Palaeontology Unlike any of the other museum publications, those of the KwaZulu- No. of pages of No. Palaeontology TOTAL 49 76 33 66 35 TOTAL articles of No. 1200 986 510 693 883 400 Entomology 1 13 2 3 2 Entomology20 5 93 44 35Mammalogy 423 Natal Museum show a monotonic increase in the numberMammalogy of articles 200 Other invertebrates 26161Other invertebrates10 14 27 16 38Ornithology 19 published over the relevant 50-year time period andOrnithology the number of Palaeontology 106133Palaeontology 3 0 94 221 32 0 Herpetology pages0 spanned by these articles also evidences a strongHerpetology increase over Mammalogy 0051611Mammalogy 0 0 53 162 94 time. Ornithology 45 51 16 24 17 Ornithology 1178 795 246 196 268 Herpetology 06341Herpetology 0 71 57 41 47 Two disciplines dominate these publications over the entire relevant time period, entomology and invertebrates other than insects, 80 1400 and the output in terms of these two fields are essentially entirely 70 1200 responsible for the increase in the number of articles published and 60 TOTAL 1000 TOTAL pages spanned over time. Indeed, there is an indication that disciplines 50 Entomology Entomology 800 other than these two have decreased in their representation, except 40 Other invertebrates Other invertebrates perhaps for palaeontology. This is congruent with the focussing of the 600 30 Palaeontology No. of pages of No. Palaeontology
No. of articles of No. journal purely on invertebrate contributions from 2001 onwards and 400 20 Mammalogy Mammalogy as reflected in the change in journal name at that point. 200 10 Ornithology Ornithology 0 Herpetology 0 Herpetology Overall trends over1964-1973 time 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 CombiningTotal the information324 from 539 all the 415museums 363considered 451 here, Total 10730 18571 13222 8816 8606 Ditsong 56 89 116 65 105 Ditsong 2079 2525 2448 1490 1242 showsKwaZulu-Natal that overall the 106 number 108 of articles 116 published 149 in the scientific 239 KwaZulu-Natal 3077 2543 3714 3238 4736 journalsDurban produced by49 these museums 76 peaked 33 during 66 1974 - 1983, 35 Durban 1200 986 510 693 883 Figs 7a & b. The number of articles published (7a) and the number of pages decreasedBloemfontein subsequently 12 over the 26 next two 49 ten-year 39 periods, 23and Bloemfontein 173 1904 2019 979 535 Eastern Cape 27 39 27 14 5 Eastern Cape 751 2134 1015 446 120 spanned by these articles (7b) in the scientific journals published by the Durban recovered slightly in the most recent ten-year period (Fig. 9a & b). Iziko 74 201 74 30 44 Iziko 3450 8479 3516 1970 1090 Natural Science Museum for each of the five 10-year periods spanning 1964 – 2013, in total and for each discipline individually. 600 20000 18000 a subsequent decrease over the following two ten-year periods, 500 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013Total 16000 Total TOTALfollowed by an increase106 over 108the last two 116 ten-year 149 periods. 239 TOTAL 3077 2543 3714 3238 4736 Ditsong 14000 Ditsong Entomology 49 58 62 76 122 Entomology400 1499 1212 1866 1649 2523 In terms of disciplines served, ornithology was overall dominate 12000 KwaZulu-Natal KwaZulu-Natal Otherin terms invertebrates of content 37 but the disciplines 34 48served has 68 become 95more Other invertebrates 795 1045 1548 1481 1964 Palaeontology 6 10 0 1 22 Palaeontology300 213 188 0 35 249 10000 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013Durban Durban Mammalogybalanced over time,12000 with entomology, palaeontology and mammalogyMammalogy 4 38 0 0 0 8000 No. of pages of No. Total 324 539 415 363 451 Total articles of No. 200 10730 18571 13222 8816 8606 Bloemfontein Bloemfontein Ornithologyalso contributingDitsong substantially4000056 at times 89in the last 116 three ten-year 65 105Ornithology Ditsong 100 20790000 2525 2448 1490 1242 6000 Eastern Cape Eastern Cape Herpetology KwaZulu-Natal94640 106 108 116 149 239Herpetology KwaZulu-Natal 466 3077 60 2543 300 3714 73 3238 4736 0 4000 periods examined. 100 Durban 49 76 33 66 35 Durban 1200 986 510 693 883 Iziko Iziko 2000 Bloemfontein 12 26 49 39 23 Bloemfontein 173 1904 2019 979 535 300 5000 Eastern Cape 27 39 27 14 5 Eastern Cape0 751 2134 1015 446 120 0 4500 Iziko 74 201 74 30 44 Iziko 1964-19731974-19831984-19931994-20032004-20133450 8479 3516 1970 1090 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 250 4000 TOTAL TOTAL 3500 200 600 20000 Entomology 3000 Entomology 18000 150 500 Other invertebrates 2500 Other invertebrates 16000 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013Total 2000 Total
Palaeontology pages of No. Palaeontology
TOTAL 106 108 116 149 239 TOTAL articles of No. 3077 2543 3714 3238 4736 14000 100 400 Ditsong 1500 Ditsong Entomology 49 58 62 76 122 Entomology 1499 1212 1866 1649Mammalogy 2523 1000 12000 Mammalogy 50 KwaZulu-Natal KwaZulu-Natal Other invertebrates 37 34 48 68 95 Other invertebrates 795300 1045 1548 1481Ornithology 1964 500 10000 Ornithology Palaeontology 6 10 0 1 22 Palaeontology 213 188 0 35 249 Durban Durban 0 0 8000 No. of pages of No. Herpetology No. of articles of No. 200 Herpetology Bloemfontein Bloemfontein Mammalogy 12000Mammalogy 4 38 0 0 0 6000 Ornithology 40000Ornithology 100 0000Eastern Cape Eastern Cape 4000 Herpetology 94640Herpetology 466100 60 300 73 0 Iziko Iziko 2000 0 0 300 5000 1964-19731974-19831984-19931994-20032004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 4500 250 4000 Figs 9a & b. The number of articles published (9a) and the number of pages TOTAL TOTAL 3500 200 spanned by these articles (9b) in the scientific journals published by the natural Entomology Entomology 3000 science museums of South Africa for each of the five 10-year periods spanning 150 2500 Other invertebrates Other invertebrates 1964 – 2013, in total and for each museum individually. 2000
Palaeontology pages of No. Palaeontology No. of articles of No. 100 1500 Mammalogy 1000 Mammalogy 50 This recent increase, however, has been driven by increases in the Ornithology 500 Ornithology number of articles produced by the KwaZulu-Natal Museum and, to 0 0 Herpetology Herpetology a lesser extent, Ditsong, with the other museums showing relative stagnation or typically ongoing decreases in this regard. Examining the overall number of pages spanned by these articles again reveals a peak during 1974 – 1983 but shows a monotonic decrease thereafter Figs 8a & b. The number of articles published (8a) and the number of pages spanned by these articles (8b) in the scientific journals published by the that almost levels out in the most recent ten-year period. Only the KwaZulu-Natal Museum for each of the five 10-year periods spanning 1964 – KwaZulu-Natal Museum evidences an overall increase in the number 2013, in total and for each discipline individually. of pages published over the relevant time period, although the Durban 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 Entomology 85 141 154 131 173 Entomology 3857 5055 4727 3346 3734 Other invertebrates 86 127 97 105 127 Other invertebrates 2413 4243 3196 2311 2703 Palaeontology 54 139 75 53 95 Palaeontology 1671 5786 2681 1804 1331 6Mammalogy 22 37 32 27 18EDITORIAL Mammalogy 398 1277 1439Durban 628 Natural Science 190 Museum Novitates 37 Ornithology 59 64 29 31 29 Ornithology 1657 1522 603 304 432 Herpetology 18 31 28 16 9 Herpetology 734 688 576 323 216
200 ‘Outside’7000 authors too may be under increasing pressure to publish 180 in6000 international journals. It is also not clear whether a decrease or 160 stagnation in submissions to these journals is responsible for the 5000 140 Entomology Entomology 120 trend, or if the fault lies with decreased or stagnated capacity by Other invertebrates 4000 Other invertebrates 100 these journals to maintain or expand their output. But it is also 3000 80 Palaeontology possible that this general decrease/stagnation in Palaeontologymuseum journal No. of pages of No. 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 articles of No. 60 1964-1973 1974-1983 1984-1993 1994-2003 2004-2013 Mammalogy Mammalogy contributions2000 indeed reflects the same trend in general research Entomology 85 141 154 131 173 Entomology40 3857 5055 4727 3346 3734 Ornithology 1000 Ornithology Other invertebrates 86 127 97 105 127 Other invertebrates20 2413 4243 3196 2311 2703 output from South African natural scientists in museums and/or Herpetology Herpetology Palaeontology 54 139 75 53 95 Palaeontology0 1671 5786 2681 1804 1331 elsewhere0 in the country. Mammalogy 22 37 32 27 18 Mammalogy 398 1277 1439 628 190 Ornithology 59 64 29 31 29 Ornithology 1657 1522 603 304 432 Natural science museum journals have always generally been of Herpetology 18 31 28 16 9 Herpetology 734 688 576 323 216 most importance to invertebrate biologists and palaeontologists, and this remains the case. They have always been of lesser importance 200 7000 to vertebrate biologists, who likely used other publication outlets to 180 6000 160 a greater degree, and their importance to this quarter seems to be 5000 140 Entomology Entomology declining further. It is therefore perhaps not surprizing that the only 120 Other invertebrates 4000 Other invertebrates natural science museum journal showing a consistent and marked 100 increase in output is African Invertebrates. 80 Palaeontology 3000 Palaeontology No. of pages of No. No. of articles of No. 60 Mammalogy 2000 Mammalogy In ending this editorial, it appears worth briefly mentioning some 40 Ornithology 1000 Ornithology aspects directly relevant to decision-making related to the present 20 Herpetology Herpetology and future of natural science museum journals. 0 0 Key advantages to museums producing natural science journals include: 1) providing an outlet for research results in which the museum has an important stake, e.g. when authored by staff employed Figs 10a & b. The number of articles published (10a) and the number of by the museum or personnel otherwise directly associated with the pages spanned by these articles (10b) in all the scientific journals published institution; 2) the provision of an outlet for worthy highly specialized by the natural science museums of South Africa combined for each discipline or lengthy contributions that might struggle to find acceptance individually and for each of the five 10-year periods spanning 1964 – 2013. elsewhere; 3) providing a basis for publication ‘exchange agreements’ Natural Science Museum also shows a pattern of increase in this with other institutions; and 4) enhancing the prestige of the museum. regard over the past three ten-year periods. Key disadvantages include: 1) the challenge of competing with higher- In terms of disciplines, articles on invertebrates and palaeontology impact outside journals; 2) distraction of research-staff time and evidence recent overall increases, while those covering the three effort; and 3) costs. vertebrate fields considered here show decreases (Fig. 10a & b). It is also relevant to briefly compare ‘hard-copy’ printing vs. A similar pattern is evident when examining the number of pages electronic publication through the internet of journals. Advantages spanned by these articles, although palaeontology shows ongoing to printing include: 1) servicing exchange agreements; 2) permanence decreases in this regard similar to the trends shown by the vertebrate and immutability; 3) a more ‘tangible’ output; and 4) a product more groups. easily used in enhancing the prestige of the museum. Advantages to e-publication include: 1) potentially quicker and cheaper production; Conclusions 2) greater flexibility, as constraints such as length, use of colour and Although this review has been based on an examination of both the inclusion of other material, e.g. sound and video files, that can be number of articles published and the number of pages spanned by conveyed by electronic media, fall away; 3) ease of access; 4) many each article, in practice both approaches yielded similar results in researchers, perhaps specially younger researchers, see e-publication most instances. as essential when selecting outlets for their research products; This review suggests an overall trend reflecting a decrease or and 5) less use of physical resources and the energy used in their at least stagnation in the amount of material covering the natural production. Certainly, there would seem no justification for not going sciences published in the scientific journals produced by the museums at least partially digital and increasingly less justification for not going in South Africa that are solely or partially focused on the natural wholly digital in presentation. sciences. The only notable exception in this regard is the KwaZulu- Many of these factors are likely to play a role in influencing the Natal Museum and its journal currently named African Invertebrates. future of South African natural science museum journals. In an era often described as a burgeoning ‘Information Age’, it is of concern that South African natural science museums generally seem David Allan not to be increasing their contribution to global scientific knowledge directly through the channels provided by their respective natural science journals. This is particularly so given that publications in the natural sciences, particularly in the disciplines covered by these museum journals, are often directly relevant to the growing world biodiversity crisis. It cannot be merely assumed that scientific publication output by researchers employed at these museums has necessarily also decreased or stagnated. These researchers may well be publishing elsewhere, especially where under pressure to publish in international journals with high ‘impact factors’. Also, many authors from outside museums also publish in museum journals and the trend may be influenced by factors affecting contributions from this quarter. Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 7
THE CAINOZOIC PALAEONTOLOGY AND STRATIGRAPHY OF KWAZULU-NATAL.
PART 5. THE FALSE BAY CORAL LIMESTONE FORMATION. STRATIGRAPHY AND FAUNA
MICHAEL ROBERT COOPER Durban Natural Science Museum, P.O. Box 4085, Durban 4000, South Africa email: [email protected]
ooper, M.R. 2014. The Cainozoic palaeontology and stratigraphy of KwaZulu-Natal. Part 5. The False Bay Coral Limestone CFormation. Stratigraphy and fauna. Durban Natural Science Museum Novitates 37: 7-24. The False Bay Coral Limestone is the youngest unit of the Uloa Subgroup and correlates with the Port Durnford Formation for which a late Early Pleistocene (Calabrian) age has been established. The molluscan fauna of the formation is described here for the first time. Four extinct species point to the antiquity of the deposit. Eight species are extant but extralimital and thermophilic, typical of tropical Indo-Pacific faunas to the north. Three species are extant in KZN but marginal, being more abundant in warmer waters to the north. Eight species still live along the KZN coast, extending as far south as the Eastern Cape. The fauna is dominated by the bivalves Spiniplicatula muricata (Sowerby), Decatopecten amiculum (Philippi), Mimachlamys sanguinea (Lamarck) and Anadara jacksoni sp. nov., with which are associated extinct Nicaisolopha tridacnaeformis (Cox), Hyotissa virleti (Deshayes) and Megerlina levis Hiller. Also present are extant predominantly tropical species of Barbatia, Saccostrea, Dendrostrea, Anomia, Chama, Fragum, Lutraria, Circe, Gafrarium, Timoclea, Pitar, Gari and Diplodonta, as well as the gastropods Glossaulax, Naticarius, Canarium, Ovatipsa and Pleuroploca.
KEYWORDS: False Bay Coral Limestone Formation, late Early Pleistocene, thermophilic extralimital coral-reef fauna, Mollusca, palaeontology, stratigraphy.
INTRODUCTION current-stable and –unstable attitude. The base of this unit is marked by occasional reworked Cretaceous concretions and subrounded The False Bay Coral Limestone Formation is a thin transgressive intraclasts up to 70 cm in diameter suggesting a minor stratigraphical marine sequence which rests unconformably on Late Cretaceous discontinuity. Onshore exposures of this upper member can be strata. It is exposed intermittently along the western shore of False divided imprecisely into lower and upper subunits. Surface sampling of Bay, Lake St Lucia, between Lister’s Point and Picnic Point (Hobday the lower subunit has yielded the hard substrate-associated epifauna & Orme 1974; Hobday 1976; Cooper & McCarthy 1998) (Fig. 1). At described here, dominated by medium-sized scallops (up to 80 mm Lister’s Point it overlies Late Coniacian (Gauthiericeras margae Zone) long), oysters and arcids (up to 60 mm), with which are associated the strata of the St Lucia Formation with an uneven basal contact which echinoid Laganum, the extinct brachiopod Megerlina laevis Hiller, and dips south at about 5o and, at very low lake level, it is exposed from the mushroom coral Fungia. Molluscs are scarce in the upper subunit, -1.0 m to +3.4 m (SACS 1980). or too comminuted for recognition, but there are fairly large (<50 At the base of the formation is some 30 cm of compact cm) randomly-orientated blocks of colonial coral (Favia, Favites and a homogeneous coarse-grained biocalcarenite which has been piped number of unidentified forms) as well as occasional upright bioherms down into the underlying Cretaceous by large irregular burrows. (it cannot positively be shown they are in situ and their orientation Although these burrows have been identified as Ophiomorpha (Hobday may be accidental). An upward increase in comminuted bioclasts 1976) they are too large for callianasid mud prawns, the makers points to marine regression, progressively-increasing water turbulence of this ichnogenus. Offshore exposures at the stratotype, revealed (wave action) and shoaling of the depositional environment. Much only at very-low lake level, are dominated almost exclusively by the of the upper subunit comprises coarse-grained bioclastic washover bivalve Spiniplicatula, many orientated subhorizontally and some with debris from an offshore barrier reef. conjoined valves. This may be an in situ thanatocoenosis. but the bulk of the material has been transported locally by wave and storm AGE OF THE FORMATION activity. This basal member is followed by some 4 m of hard, well-lithified, The False Bay Coral Limestone was deposited by a +5-8 m highstand unbedded, vuggy, bioclastic calcarenite with randomly-orientated of sea level which, initially, was dated to the Later Pleistocene coarse shell debris which includes frequent whole shells in both (Eemian) (Hobday & Orme 1974; SACS 1980). However the formation 8 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
has long been correlated with the Port Durnford Formation further Kilburn 2001). However a rich Mio-Pliocene pectinoid fauna occurs to the south (Orme 1973; Hobday & Orme 1974; Cooper 1991; in the Uloa Subgroup (Cooper in prep.) with Mirapecten, Mimachlamys Cooper & McCarthy 1998), for which an Early Pleistocene age has and Minnivola conspicuous, but very different from the living faunas. since been established for the vertebrate fossils (Hendey & Cooke 1985; Hendey & Volman 1986). This age is corroborated by the 87Sr/86Sr signature obtained from a shell of Paratapes textile (Gmelin) (B. Eglington unpublished data) which translates to an age of 1.7 Ma (McKenzie et al. 1990), i.e. late Early Pleistocene (Early Calabrian). Biostratigraphically this corresponds approximately with nannofossil zone NN19 (Liu et al. 1996) and oxygen-isotope stage 63 (Lisiecki 2005). The compositional disparity between the coeval False Bay and Port Durnford faunas is attributed to facies difference, the Port Durnford Formation having been deposited in a muddy back-barrier lagoon in which the mud crab (Scylla serrata) abound (Cooper & Kensley 1992). Since a Late Pleistocene (Eemian, oxygen-isotope substage 5e, =126 ka) age is favoured for the undoubtedly younger Mduku Formation which occurs further inland along the Mzinene River (Cooper 1999), the False Bay Coral Limestone is late Early Pleistocene. Further support for this antiquity is provided by three extinct species and the occurrence of Nicaisolopha tridacnaeformis (Cox), which otherwise is known only from the Pliocene of East Africa; Hyotissa virleti (Deshayes) ranges from Middle Miocene to Early Pleistocene (Cox 1927, 1930), but may not be distinct from living H . hyotis (Linné).
DEPOSITIONAL ENVIRONMENT
Present-day False Bay (Fig. 1) is a silt trap for the vast amounts of sediment pumped annually into Lake St Lucia by the Mzinene, Nyalazi and Hluhluwe Rivers. For large parts of the year its turbid waters are brown with silt, mud and suspended organic debris. Freshwater catfish Clarias gariepensis are an important component of the ichthyofauna. This environment is deleterious to reef formation, colonial corals favouring clear well-oxygenated marine water. Thus for colonial corals to thrive in False Bay, the present-day entry of silt and mud had to be inhibited. This could occur only if reed-choked marshes and papyrus swamps existed at river entry points, filtering out suspended fines. Thus it seems certain the Muzi swamp is a small relict of a once- great Early Pleistocene swampland analogous to the Bangweulu of Zambia. At this time water salinity in False Bay would have fluctuated seasonally and hence significantly the dominant corals in the fauna, Favia and Favites, are euryhaline taxa. The present-day tropical Red Sea, in which salinities vary between 36-41‰ (average seawater is Fig. 1 Locality map. ~35‰), provides a close modern analogue. The abundance of the scallops Decatopecten amiculum and Mimachlamys sanguinea points FAUNAL AFFINITIES to a carpet of the marine grass Thalassodendron ciliatum (Dijkstra & Kilburn 2001), indicating deposition in the lower intertidal to The faunal affinities of the False Bay Coral Limestone are clearly with upper sublittoral zone with water depths <17 m. Thus the False Bay the tropical Indo-Pacific, living elements of which enter northern Coral Limestone was deposited in a shallow marine embayment of KZN in the coral reefs at Sodwana Bay. Of the species recorded four warm clear tropical water. The absence of patellids and trochids are extinct: among many others testify to a predominantly soft substrate, mainly Nicaisolopha tridacnaeformis (Cox) bioclastic sand, and the scarcity of rocky outcrops. Hyotissa virleti (Deshayes) Intriguingly, given the diversity of pectinoids off the coast of Anadara jacksoni sp. nov. (endemic) northern KawZulu-Natal (Dijkstra & Kilburn 2001), its shallow- Megerlina levis Hiller (endemic) marine Pleistocene deposits are impoverished. Barnard (1964) reported Decatopecten plica (Linné) from the Harbour Formation of Eight species are extralimital, found only in tropical warm waters to Durban, and Cooper (1999) figured Volachlamys fultoni (Sowerby) from the north as far as the Red Sea: the Port Durnford Formation. To this may be added the two species Dendrostrea folium (Linné) described here from the False Bay Coral Limestone. This suggests the Spiniplicatula muricata (Sowerby) modern pectinoid fauna is a product of recent invasion from other Fragum nivale (Reeve) faunal provinces, in particular the Indo-West Pacific (Dijkstra & Timoclea costellifera (Adams & Reeve) Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 9
Fig. 2. A. Barbatia foliata (Forskal), DNSM-PQZ5867. B-C. Anadara jacksoni sp. nov. B, a paratype, DNSM-PQZ5859; C, the holotype, DNSM-PQZ5858. D-E. Mimachlamys sanguinea (Linné). D, DNSM-PQZ5870; E, DNSM-PQZ5910. F-H. Decatopecten amiculum (Philippi). F, DNSM-PQZ5860; G, DNSM-PQZ5908; H, DNSM- PQZ5902. I-K. Saccostrea cucullata (Born), I-J, DNSM-PQZ5868, K, DNSM-PQZ5872. L. Spiniplicatula muricata (Sowerby). DNSM-PQZ5851. 10 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
Pitarina hebraea (Lamarck) Barbatia foliata (Forsskål) Kilburn & Rippey 1982: 157, pl. 35, fig. 9, Gafrarium pectinatum pectinatum (Linné) text-fig. 105; Kilburn 1983: 516, figs 4-5; Drivas & Jay 1988: 135, pl. 52; Diplodonta aff. subrotundata Issel Springsteen & Leobrera 1986: 294, pl. 84; Bosch et al. 1995: 207, fig. Pleuroploca trapezium (Linné) 1911; Paulay 1996: 21.
Three species prefer tropical warm waters to the north, but extend Material into northern KwaZulu-Natal: A single specimen, DNSM-PQZ5867. Decatopecten amiculum (Philippi) Mimachlamys sanguinea (Linné) Measurements Gari maculosa (Lamarck) L H T AL DNSM-PQZ5867 46 29.6 (0.64) 12.1 (0.26) 12 (0.26) Eight species occur along the entire coast of KZN, extending into the Eastern Cape: Description Naticarius alapapilionis (Röding) Medium sized, inequilateral, compressed, elliptical, with anteriorly- Ovatipsa chinensis (Gmelin) positioned low umbones, rounded anterior and posterior margins, and Canarium mutabilis (Swainson) uniform height posteriorly. Flanks ornamented with fine radial ribs Glossaulax didyma (Röding) which strengthen posteriorly. Chama limbula Lamarck Barbatia foliata (Forsskål) Discussion Anomia acheus Gray The specimen displays the features of highly variable and widespread Saccostrea cucullata (Born) Indo-Pacific B. foliata, an extant species which also occurs in the Harbour Formation of Durban (Barnard 1964), and extends as SYSTEMATIC PALAEONTOLOGY far south as Port Alfred. Barbatia lacerata (Bruguière) which is recorded from the Pleistocene of Kenya (Cox 1930) is much broader All the material described here is housed in the palaeontological posteriorly and has a trapezoidal outline. collections of the Durban Natural Science Museum (DNSM). Abbreviations for measurements are as follows: L = length, Lmax Subfamily Anadarinae Reinhart, 1935 = maximum length, H = height, T = thickness (of a single valve), AL Genus Anadara Gray, 1847 = anterior length (length of valve in front of beaks), l = length of Type species: Arca antiquata Linné, 1758; by original designation. structures other than valve, h = height of structures other than valve; percentages relative to shell length are in brackets. Some of the Anadara jacksoni sp. nov. identifications are based on information provided by the following Fig. 2B-C, 7A-C websites: www.conchylinet.com www.conchologistsofamerica.org Etymology www.conchology.be www.coralhub.info jacksoni – for Garnet and Toni Jackson, former wardens at False Bay www.elrincondelmalacologo.com www.en.wikipedia.org Park, for assistance, friendship and hospitality over very many years. www.femorale.com/shellphotos www.idscaro.net www.jaxshells.org www.gastropods.com Material www.malacs.chez www.mollusca.co.nz The holotype, DNSM-PQZ5858, and 15 more-or-less fragmentary www.marinespecies.org www.molluskonline.com paratypes 5859, 6937-6950. www.pectensite.com www.reeffrontiers.com www.saltcorner.com www.scallop.nl/classification Measurements www.seashellhub.com www.seashells.com L H T AL www.topseashells.com www.vianetconchology.com DNSM-PQZ5858 56 45.2 (0.81) 23.8 (0.43) 19.0 (0.34) www.worldwideconchology.com DNSM-PQZ6937 c.58 49.2 (0.85) 25.5 (0.44) 20.5 (0.35)
Class BIVALVIA Linné, 1758 Description Superfamily ARCOIDEA Lamarck, 1809 Moderately large (Lmax = 58 mm), thick shelled, trapezoidal, longer Family Arcidae Lamarck, 1809 than high, with prominent well-rounded umbones, strongly-incurved Subfamily Arcinae Lamarck, 1809 prosogyrous beaks, inflated valves, prominent well-rounded umbonal Genus Barbatia Gray, 1842 ridge and broad flat trigonal corselet. Type species. Arca barbata Linné, 1758; by the subsequent designation In lateral profile the steep anterior margin is gently convex, of Gray (1857). passing imperceptibly into the broadly-convex ventral margin. The almost-straight posterior margin is obliquely truncate and Barbatia foliata (Forsskål, 1775) the posterodorsal margin almost straight. The corselet is almost Fig. 2A perpendicular to the flank and barely visible in lateral view. The moderately narrow (w/l = 0.15-0.23) and finely-striated Arca nivea Chemnitz, Krauss 1848: 16; Turton 1932: 217; Nardini 1937: ligamental area extends the entire dorsal length of the hinge. At both 241, pl. 15, fig. 15; Braga 1952: 104, pl. 8, fig. 7. ends there are 2-3 very short ligamental grooves. The dentition is taxodont, with a straight graded series of about 44 teeth and sockets Arca (Barbatia) helblingii Bruguière, Barnard 1964: 374; Boshoff 1965: which increase in size and inclination laterally. 109, pl. 1, fig. 2; Moura 1971: 53, pl. 9, fig. 2. The convex flanks are ornamented with 25-26 flat-topped radial Barbatia helblingi (Bruguiere) Hassan 1994: 249. ribs which increase in width ventrally and are separated by very Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 11
narrow interspaces; 6-7 similar ribs ornament the corselet. Many Material of the posterior flank ribs show a faint median depression without Numerous specimens, including DNSM-PQZ5860, 5899, 5902 and actually being grooved, and all are crenulated by commarginal growth 6972-6994. striae which increase in strength ventrally. Where not eroded the ribs are finely lirate. Measurements L H T DNSM-PQZ5860 50.2 50 (1.00) 11.8 (0.24) DNSM-PQZ5899 47.5 49 (1.03) 11.0 (0.23) DNSM-PQZ5902 31.0 32.0 (1.03) 6.0 (0.19)
Description Moderately large (Lmax = 50.2 mm), suborbicular, biconvex, equivalve, equilateral, slightly higher than long, with weakly-inflated valves and central umbones. The anterolateral and posterolateral margins are shallowly-concave, forming an apical angle of 80-85o, and the ventral margin is strongly convex. There are 5-8 low gently-convex ribs which broaden ventrally and are generally wider than the shallow interspaces, but much narrower on DNSM-PQZ5860. The entire shell surface is finely lirate, with up to 10 lirae per rib at midlength. The lirae split into finer threads close to the ventral margin in mature shells. The small auricles are of similar size, that to the posterior slightly larger, and are ornamented with up to 11 fine radial lirae. There is no byssal notch. Fig. 3 A typical example of Anadara antiquata (Linné), for comparison with A . jacksoni sp. nov. Discussion Suborbicular outline and numerous broad low rounded ribs serve Discussion to distinguish D. amiculum from D. plica (Linné) and D. radula (Linné). The presence of a few very short ligamental grooves at either end of The former is reported from the Kenya (Cox 1930). This is a tropical the cardinal area places A. jacksoni sp. nov. in the “antiquata complex” Indo-Pacific species whose southern limit in northernmost KwaZulu- which “... needs careful study using modern methods” (Kilburn 1983: Natal (Kosi Bay) where it is recorded from 40-50 m depth (Dijkstra & 536). Anadara antiquata (Linné) (Fig. 3) itself is a widely-reported Indo- Kilburn 2001). Pacific species which ranges from Miocene to Recent (Cox 1930). It differs from A. jacksoni sp. nov. in having a more-inclined corselet which Subfamily Chlamydinae von Teppner, 1922 is clearly visible in lateral view, so that the outline is more trapezoidal, Tribe Mimachlamydini Waller, 1993 and in having wider interspaces between ribs. These features also Genus Mimachlamys Iredale, 1929 serve to distinguish A. chalcantha (Reeve), A. trapezia (Deshayes) and Type species: Pecten asperrimus Lamarck, 1819; by original designation. A. maculosa (Reeve). Anadara uropigimelana (Bory) has undivided ribs and is a subquadratic species, as is A. natalensis (Krauss); the latter Mimachlamys sanguinea (Linné, 1758) is reported from the Early Pleistocene Harbour Formation (Barnard Fig. 2D-E 1964). Anadara birleyana Standen is wedge shaped with a long straight posterior margin. Anadara erythraeoensis (Philippi), which is present in Chlamys senatorius Gmelin, Nardini 1937: 233, pl. 14, fig. 4; Cox 1927: the Port Durnford Formation, is posteriorly produced and elongate- 45, 75, pl. 7, figs 1-3, pl. 15, fig. 3, pl. 17, fig. 10; 1929a: 108, 152, pl. 13, trapezoidal in outline, A. ehrenbergi (Dunker) has subcentral umbones fig. 21; 1929b: 191; 1939: 76, pl. 1, fig. 14; Braga 1952: 135; Barnard 1964: and widely-spaced undivided ribs and A. demiri (Piani) is a small 430; Boshoff 1965: 135; Hassan 1994: 250. tropical species (L=20-25 mm) with oval outline and widely-spaced ribs which become nodate towards the margins. Pliocene Scapharca Chlamys senatoria Gmelin, Cox 1930: 122, 152; 1931: 6; 1936: 54, pl. 5, morrumbenensis Cox (1939: 81, pl. 2, fig. 19) is a short squat species fig. 18, pl. 6, fig. 9; Eames, 1950: 145; Eames & Cox 1956: 14, 40 (cum. with only about 16 strong rounded radial ribs. synon.); Bosch et al. 1995: 230, fig. 1003.
Superfamily PECTINOIDEA Wilkes, 1810 Mimachlamys sanguinea (Linné) Dijkstra & Knudsen 1998: 83, pl. 4, fig. Family Pectinidae Wilkes, 1810 16 (cum synon.); Dijkstra & Kilburn 2001: 305, figs 44-45 (cum synon.). Subfamily Pectininae Wilkes, 1810 Tribe Decatopectinini Waller, 1986 Material Genus Decatopecten Rüppel, 1839 Numerous specimens, including DNSM-PQZ5870, 5910 and 6924-36. Type species. Ostrea plica Linné, 1758; by monotypy. Measurements Decatopecten amiculum (Philippi, 1851) L H T Fig. 2F-H, 7D DNSM-PQZ6924 75.0 78.0 (1.04) 4.0 (0.05) DNSM-PQZ5910 59.5 67.6 (1.14) 4.5 (0.08) Decatopecten amiculum (Philippi) Abbott & Dance 1982: 307; Bernard DNSM-PQZ5870 46.0 50.1 (1.09) 3.0 (0.07) et al. 1993: 51; Poutiers 1998: 205; Dijkstra & Knudsen 1998: 55, pl. 5, fig. 25, pl. 9, figs 42–43 (cum. synon.); Dijkstra & Kilburn 2001: 276, figs Description 10-11.. Medium sized (Lmax = 75 mm), orbicular, equilateral, strongly 12 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
Fig. 4 A,C. Nicaisolopha tridacnaeformis (Cox). A, DNSM-PQZ5861; C, DNSM-PQZ5901. B. Dendrostrea folium (Linné). DNSM-PQZ5898. D-E. Chama limbula Lamarck. DNSM-PQZ5869, F-G. Pitarina hebraea (Lamarck), DNSM-PQZ5878. H-J. Fragum nivale (Reeve), DNSM-PQZ5875. K-N. Circe scripta (Linné). K-L, DNSM-PQZ5862; M-N, DNSM-PQZ5897. O-P. Gafrarium pectinatum (Linné). O, DNSM-PQZ5903; P, DNSM-PQZ5877. Q-T. Gari maculosa (Lamarck). Q-R, DNSM-PQZ5863; S-T, DNSM-PQZ5905. U-V. Lutraria aff. australis Deshayes, DNSM-PQZ5866. W-X. Timoclea costellifera (Adams & Reeve). W, DNSM-PQZ5907; X, DNSM-PQZ7011. Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 13
compressed, auriculate with unequal ears, deep byssal notch and an the first with 7 simple strong rounded ribs, and the second with a apical angle of 85-102o. The flanks are ornamented by 24-26 strong strongly convex left valve, large attachment area and unequal irregular simple round-topped radial ribs which are slightly narrower than ribs. Plicatula exigua Cox (1930: 126) is yet another small East African the interspaces. But for small patches of well-spaced transverse Pliocene species; it is trigonally-ovate, with about 10 simple acute ribs scales to rib crests microsculpture is not preserved. The auricles are radiating from the umbo. ornamented with radial costellae, six on the posterior ear. Superfamily ANOMIOIDEA Rafinesque, 1815 Discussion Family Anomiidae Rafinesque, 1815 The False Bay material is typical of this common and widespread Genus Anomia Linné, 1758 tropical Indo-Pacific species which ranges from Middle Oligocene Type species: Anomia ephippium Linné, 1758; by the subsequent to Recent (Cox 1939); vagrant specimens have been recorded as far designation of Schmidt (1818). south as southern KwaZulu-Natal. Anomia achaeus Gray, 1850 Family Plicatulidae Watson, 1930 Fig. 5C-D Genus Spiniplicatula Habe, 1977 Type species: Plicatula muricata Sowerby, 1873; by original designation. Anomia ephippium Linné, Barnard 1964: 442; Boshoff 1965: 139.
Spiniplicatula muricata (Sowerby, 1873) Anomia achaeus Gray, Kilburn & Rippey 1982: 172, pl. 39, fig. 1; Oliver Fig. 2L 1992: 85; Bosch et al. 1995: 234, fig. 1017.
Plicatula muricata (Sowerby) Bernard et al. 1993: 56. Material Three specimens, DNSM-PQZ5871 and 7041-42. Material Numerous specimens, including DNSM-PTQ5843-5861 and 6995-6997. Measurements L H T AL Measurements DNSM-PQZ5871 c.54.0 45.0 (0.83) 13.0 (0.24) c.22 (0.41) L H T AL DNSM-PQZ5849 49.0 57.0 (1.16) 12.1 (0.25) 28.4 (0.58) Description DNSM-PQZ5851 49.5 58.5 (1.18) 11.0 (0.22) 22.0 (0.44) Relatively large, irregularly-subcircular, weakly-convex with thin test DNMSM-PQZ5852 48.0 50.0 (1.04) 13.2 (0.28) 27.5 (0.57) and feeble pustules in indistinct widely-spaced radial rows. DNSM-PQZ5847 41.3 48.7 (1.18) 10.0 (0.24) 21.0 (0.51) Discussion Description The specimens conform to variable A. achaeus, the only extant Indo- Relatively large (Lmax = 53 mm), robust, equilateral to markedly West Pacific species; geographically it ranges as far south as Algoa inequilateral, higher than long, trigonally-ovate to oval, generally Bay (Eastern Cape). Anomia ephippium Linné, recorded from South curved and posteriorly produced, with weakly-inflated valves. Africa in error (Barnard 1964), is restricted to the northeast Atlantic, Ornament comprises 7-9 irregular radial plicae which, in middle to Mediterranean and Black Sea. later growth, branch 2-3 times so that there are up to 36 plications to the commissure. Plicae are about as wide as the interspaces but pinch- Superfamily OSTREOIDEA Rafinesque, 1815 and-swell irregularly, and are ornamented with short hyote spines and Family Ostreidae Rafinesque, 1815 pustules. Dentition is isodont, with a simple tooth on either side of Subfamily Crassostreinae Torigoe, 1981 the triangular ligament pit. Tribe Striostreini Harry, 1985 Genus Saccostrea Dollfuss & Dautzenberg, 1920 Discussion Type species: Ostrea sacellus Dujardin, 1835, = O. cucullata Born, 1778; This bivalve overwhelmingly dominates offshore exposes of the False by monotypy. Bay Coral Limestone to virtual exclusion of all else, pointing to depth- related ecological partitioning of the fauna. It closely resembles extant Saccostrea cucullata (Born, 1778) Indonesian S. muricata although the latter species is smaller (Lmax = Fig. 2I-K 32 mm). Plicatula squamosissima Smith (Kilburn & Rippey 1982: 170, pl. 38, fig. 9) is a smaller (L<30 mm) more finely-ribbed species with Ostrea cucullata Born, Krauss 1848: 30; Cox, 1929a: 154; 1929b: 178; numerous fine divaricating riblets. The False Bay material resembles 1931: 6; 1939: 97; Nardini 1937: 226, pl. 12, figs 4-5; Braga 1952: 96, pl. S. horrida (Dunker) but the latter is a much-smaller northwest Pacific 7, fig. 5; Franca 1960: 91, pl. 23, figs 1-2, pl. 24, fig. 1; Hassan 1994: 250. and Chinese species which has fewer ribs. Large size, robust shell, subtrigonal outline, with radiating dichotomizing plicae immediately Crassostrea cuccullata (Born) Barnard 1962: 446; Boshoff 1965: 141, pl. distinguishes the fossil material from P. plicata (Linné) (Nardini 1937: 12, fig. 2; Kilburn & Rippey 1982: 169, pl. 38, fig. 4, text-fig. 134. 228, pl. 13, figs 2-5) which also has fewer non-squamose plicae. Plicatula imbricata Menke and P. gibbosa Lamarck are both widespread Saccostrea cucullata (Born) Stenzel 1971: N1134, fig. J106; Harry, 1985: Indo-Pacific species which range from Miocene to Recent (Cox 150; Bosch et al. 1995: 228, fig. 996; Siddiqui & Ahmed 2002: 115, Fig. 1930); both differ from False Bay material in having rounder outlines 5B-C; Lam & Morton 2004: 20, figs 4D-F). and fewer non-dichotomizing plicae. Plicatula ramosa Lamarck and P. simplex Gould are trigonal species with fewer simple radiating plicae. Material Pliocene P. woodae Cox (1930: 125, pl. 14, figs 20-23) and P. mombasana Four specimens, DNSM-PQZ5868, 5871-72 and 6953-6955. Cox (1930: 126, pl. 14, figs 16-18) are both small orbicular species; 14 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
Measurements Material L H T One specimen, DNSM-PQZ5898. DNSM-PQZ5868 25 38 (1.52) 14.8 (0.59) Measurements L H T Description DNSM-PQZ5898 44.3 40 (0.90) 8.2 (0.19) Medium sized, irregularly elongate oval to subrectangular, much higher than wide, with thick solid test, strongly-convex left valve, Description deep umbonal cavity and moderately large attachment area. There are Small (Lmax = 44.3 mm), suborbicular, with large attachment area, irregular wavy radial ribs, a coarsely-plicate commissure and a few pit shallow left valve, very shallow umbonal cavity, short ligament area, chomata in the umbonal area. lack of commissural shelf, restricted chomata, crescentic adductor scar, seven simple round-topped radiating plicae and lustrous Discussion pustulose interior. This is a highly-variable and common Indo-Pacific species which ranges from Miocene to Recent (Cox 1930) and has been recorded Discussion from as far south as Algoa Bay (Eastern Cape). This is a widespread tropical Indo-Pacific species which ranges from Early Miocene to Recent (Cox 1930) and hgas been recorded from Subfamily Liostreinae Malchus, 1990 as far south as southern Moçambique Elongate form and narrow Tribe Flemingostreini Stenzel, 1971 attachment area (to mangrove roots) distinguishes D. frons (Linné). Genus Nicaisolopha Vyalov, 1936 Type species: Ostrea nicaisei Coquand, 1862; by original designation. Family Gryphaeidae Vyalov, 1936 Subfamily Pycnodonteinae Stenzel, 1959 Nicaisolopha tridacnaeformis (Cox, 1927) Tribe Hyotissini Harry, 1985 Fig. 4A,C Genus Hyotissa Stenzel, 1971 Type species: Mytilus hyotis Linné, 1758; by original designation. Alectryonia aff. crista-galli (Linné) Newton 1900: 549, pl. 12, figs 5-6. Hyotissa virleti (Deshayes, 1833) Ostrea tridacnaeformis Cox 1927: 71, pl. 15, fig. 1; 1929: 181; 1930: 128, Fig. 5A-B pl. 14, fig. 25. Ostrea virleti Deshayes, Cox 1927: 66, pl. 11, pl. 16, fig. 1; 1929b: 177; Material 1930: 127, 155, pl. 15, figs 9-10. Thirteen specimens, DNSM-PQZ5861, 5901 and 6998-7008. Ostrea fuchsiana Newton 1899: 205, pl. 9, figs 1-2). Measurements L H T Ostrea (Alectryonia) virleti Deshayes, Douglas 1927: 6, pl. 1, fig. 3, pl. 2, DNSM-PQZ5861 50.0 47.0 (0.94) 15 (0.30) figs 1-2.
Description Ostrea (Lopha) virleti Deshayes, Cox 1936: 49, pl. 7, fig. 2. Medium sized, irregularly ovate, thin shelled, weakly convex, about as long as high, with 2-5 round-crested simple radial plicae with Hyotissa virleti (Deshayes) El-Hedeny 2005: 719, pl. 4, fig. F. prominent squamae near the ventral margin. Material Discussion Four specimens, DNSM-PQZ7013 and 7021-24. Nicaisolopha tridacnaeformis (Cox) is recorded from the Pliocene of Zanzibar, the Red Sea, Iran and the Mekran coast. Superficially similar Measurements Lopha cristagalli (Linné) has up to nine sharp-crested plicae and L H T AL claspers. DNSM-PQZ7013 c.205 c.245 (1.20) 65 (0.32) 93 (0.45)
Subfamily Lophinae Vyalov, 1936 Description Tribe Lophini Vyalov, 1936 Very large, heavy, robust, curved, much higher than long with a broad Genus Dendrostrea Swainson, 1838 (70 mm) ligamental area, no umbonal cavity, very thick shell (up to Type species: Ostrea folium Swainson, 1838; by the subsequent 69 mm umbonally and 50 mm at distal end of major plicae), circular designation of Herrmannsen, 1847. adductor scar and vesicular shell structure. At the ventral commissure there are three major plicae, and 10 much smaller ones along the Dendrostrea folium (Linné, 1758) anterior margin. Fig. 4B Discussion Ostrea folium Linné, Cox 1930: 109, pl. 13, figs 28-29. Hyotissa virleti was described from the Middle Miocene of Egypt and is common in the Pliocene, but ranges into the Pleistocene of Kenya Lopha folium (Linné) Stenzel 1971: N1157, fig. J47. (Cox 1930). It is distinguished by its very thick shell and numerous Dendrostrea folium Swainson, Harry 1985: 137, fig. 18; Lam & Morton occasionally bifurcating plicae. Differences from other species of 2004: 22, figs 5A-C; Taylor & Glover 2004: 262. Hyotissa are noted by Cox (1927). Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 15
Fig. 5 A-B. Hyotissa virleti (Deshayes), DNSM-PQZ7013, x0.5; C-D. Anomia achaeus Gray, C. DNSM-PQZ5871, D. DNSM-PQZ5900. 16 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
Fig. 6 A-B. Pleuroploca trapezium (Linné), DNSM-PQZ5911; C-D. Canarium mutabilis (Swainson), C. DNSM-PQZ5880, D, DNSM-PQZ5912; E-F. Ovatipsa chinensis violacea (Rous), DNSM-PQZ5882; G-I. Naticarius alapapilionis (Röding), G. DNSM-PQZ5881, H-I. DNSM-PQZ5883; J. Glossaulax didyma (Röding), DNSM-PQZ5884. Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 17
Discussion Superfamily CHAMOIDEA Lamarck, 1809 This tropical Indo-Pacific species ranges from Tanzania northwards. Family Chamidae Lamarck, 1809 Fragum fragum (Linnaeus) Cox 1929: 159; Prashad 1932: 274; Barnard Genus Chama Linné, 1758 1964: 494; Kira 1962: 154, pl. 55, fig. 14; Moura 1971: 16, pl. 10, fig. Type species: Chama lazarus Linné, 1758; by ICZN Opinion 484. 7; www.conchology.be) differs from the present specimen in having a trapezoidal outline, more-numerous ribs and an arcuate umbonal Chama limbula Lamarck, 1809 ridge. Fragum bannoi (Otsuka) is a trapezoidal species with deeper Fig. 4D-E wider interspaces and nodose ribs, F. mundum (Reeve) has a steeper anterior slope, flat-topped ribs which are scaled anteriorly and Chama nivalis Newton (non Reeve) 1900: 556. posteriorly, and very narrow interspaces, and F. sueziense (Issel) is subcircular. Fragum carinatum (Lynge) is an Indonesian species and F. Chama limbula Lamarck, Cox 1930: 161; Kilburn & Rippey 1982: 174, pl. hemicardium Linné is an Australasian species. Fragum retusum (Linné), F. 28, fig. 7. subretusum (Sowerby) and F. inedo (Linné) are all Indo-Pacific species of the subgenus Lunulicardia. Quaternary Fragum tomlini Cox (1930: 159, Material pl. 15, fig. 12) is subquadratic and has a rounded umbonal ridge. Three specimens, DNSM-PQZ5869 and 6951-52. Superfamily MACTROIDEA Lamarck, 1809 Measurements Family Mactridae Lamarck, 1809 L H T AL Subfamily Lutrariinae Adams & Adams, 1856 DNSM-PQZ5869 38.5 44.3 (1.15) 21.0 (0.55) 20.0 (0.52) Genus Lutraria da Costa, 1778 Type species: Mya lutraria Linné, 1758; by tautonomy. Description Relatively large, thick-shelled, robust, strongly-convex left valve Lutraria aff. australis Deshayes, 1855 with spirally-incoiled umbo, eroded external surface with coarse Fig. 4U-V commarginal rugae and smooth internal margin Compare: Lutraria australis Deshayes, Bosch et al. 1995: 248, fig. 1099. Discussion This is a common and widespread Pleistocene to Recent Indo-Pacific Material species (Cox 1930) recorded from as far south as the Eastern Cape Three fragmentary specimens, including DNSM-PQZ5866. (Transkei). Krauss’s (1848) record of Chama gryphoides Linné from Quaternary deposits of KwaZulu-Natal probably pertain to this Description species. Chama lazarus Linné, C. asperella Lamarck and C. ratoi Boshoff Moderately large (Lmax ~110 mm), markedly inequilateral, with all are smaller species with crenulated internal margins and radial compressed valves (W/L = ~0.19), low broad umbo positioned rows of spines to the external valve surface. anteriorly, incurved orthogyrous beaks, almost straight antero- and posterodorsal margins, distinct umbonal ridge with deeply-excavate Superfamily CARDIOIDEA Lamarck, 1809 very narrow escutcheon and broader lunule. Shell surface with Family Cardiidae Lamarck, 1809 commarginal growth rugae only. The anterior end of the shell is Subfamily Fraginae Stewart, 1930 not preserved, but there is a small posterior gape. Internal features Genus Fragum Röding, 1798 unknown. Type species: Cardium fragum Linné, 1758; by tautonomy. Discussion Fragum nivale (Reeve, 1845) All the material is too fragmentary for positive identification but Fig. 4H-J the affinities of the fauna invite comparison with L. australis and L . rhynchaena Jonas The presence of a post-umbonal cavity allies it to Fragum nivale (Reeve) www.femorale.com/shellphotos; www. L. inhacaensis Boshoff (1965: 175, pl. 8, fig. 2, pl. 14, figs 4-5) but in marinespecies.org that species the anterior and posterior hinge borders are raised into a crest. Both L. oblonga (Gmelin) and L. philippinarum Reeve have Material concave posterodorsal margins, and that of L. lutraria is convex. Lutraria A single specimen, DNSM-PQZ5875. angustior Philippi, recorded by Barnard (1964: 522, fig. 29a, c) from South Africa, is a northeast Atlantic species which ranges south only Measurements as far as Portugal, and his material is now placed in L. capensis Reeve, a L H T AL cold-water south Atlantic species. DNSM-PQZ5875 25.2 32 (1.23) 10.0 (0.40) 8.7 (0.35) Superfamily LUCINOIDEA Fleming, 1828 Description Family Ungulinidae Adams & Adams, 1857 Small, subtrigonal, higher than long, with moderately-inflated valves, Genus Diplodonta Bronn, 1831 tiny pointed umbones and incurved orthogyrous beaks. There is a Type species: Venus lupinus Brocchi, 1814 (= Tellina rotundata Montagu, strong sharply-rounded umbonal ridge which separates the flank from 1803); a flat triangular corselet which is at an acute angle to the former. by the subsequent designation of Hermannsen, 1846. Ornament comprises flat-topped radial ribs which broaden ventrally and are wider than the punctate interspaces; there are at least 10 to Diplodonta aff. subrotundata Issel, 1869 the flank and eight on the corselet. The ribs are crenulated by growth striae but there is no evidence of tuberculation or scales. Compare: Diplodonta subrotundata Issel, Bosch et al. 1995: 237, fig. 1039. 18 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
Material Gafrarium pectinatum (Linné, 1758) Three specimens, DNSM-PQZ5873-74 and 5906. Fig. 4O-P, 7F
Measurements Gafrarium pectinatum (Gmelin) Prashad 1932: 228; Cox 1939: 84, 102; L H T AL Barnard 1964: 502; Moura 1971: 60, pl. 11, fig. 2; Kilburn & Rippey DNSM-PQZ5906 20.0 20.2 (1.01) 7.9 (0.40) 5.8 (0.29) 1982: 192; Abbott & Dance 1988: 354; Salvat & Rives 1983: 375; Drivas & Jay 1988: 146, pl. 58; Bosch et al. 1995: 268, fig. 1200. Description Small, suborbicular, thin-shelled, weakly inflated, almost equilateral Gafrarium aduncum Franca 1960: 94, pl. 26, fig. 2. with small low anteriorly-positioned umbo, tiny incurved beak, very narrow lanceolate escutcheon, no lunule, lack of ornament and two Gafrarium pectinatum (Linné) Drivas & Jay, 1988: 146, pl. 58; Salvat & oblique cardinal teeth. Reeves, 1991: 146, pl. 40; Hassan 1994: 251.
Discussion Material The material is too poorly preserved for positive identification but, Three specimens, DNSM-PQZ5877, 5903 and 5909. given the composition of the associated fauna, may belong here. Differences from other African species are noted by Kilburn (1997). Measurements Cox (1930) recorded this species from the Pleistocene of Kenya. L H T AL DNSM-PQZ5903 37.7 29.9 (0.79) 8.6 (0.23) 15.5 (0.41) Superfamily VENEROIDEA Rafinesque, 1815 Family Veneridae Rafinesque, 1815 Description Subfamily Circinae Dall, 1896 Small (Lmax = 38 mm), inequilateral, oval, compressed, longer than Genus Circe Schumacher, 1817 high, with low umbones positioned. anteriorly. There is a narrow Type species: Circe violacea Schumacher, 1817 (= Venus scripta Linné); lanceolate lunule circumscribed by a groove, and a very narrow ill- by monotypy. defined escutcheon with raised margins. Ornament comprises fine rounded nodate radial ribs, wider than the interspaces, splitting into Circe scripta (Linné, 1758) finer riblets distally. Posteriorly the radial ribs are superimposed with Fig. 4K-N a series of curved ribs to produce a cancellate ornament.
Circe scripta (Linnaeus) Cox 1930: 156; Prashad 1932: 233; Braga 1952: Discussion 111, pl. 10, fig. 5; Barnard 1964: 501; Bosch et al. 1995: 267, fig. 1197. The False Bay fossils display the essential characters of this Gafrarium scripta (Linnaeus) Franca 1960: 95, pl. 26, fig. 4; Taylor & widespread and common tropical Indo-Pacific species which ranges Glover 2004: 263. from southern Moçambique northwards and is first recorded from the Pliocene (Cox 1939). Medium size and transverse rows of nodes Material to the ribs suggest reference to the nominate race; the southern A number of specimens, including DNSM-PQZ5862, 5897 and 6956- subspecies G. p. alfredense Barsch (Kilburn & Rippey 1982, pl. 44, fig. 1) 6971. is smaller (<26 mm) and without nodes. Gafrarium tumidum (Röding) is more-coarsely ornamented. Gafrarium divaricatum (Gmelin) is Measurements smaller, rounder and more compressed, with a broader rounder umbo L H T AL and without nodes to the ribs; it is recorded from the Pleistocene of DNSM-PQZ5862 45.5 40 (0.89) 6.5 (0.14) 15 (0.30) Kenya (Cox 1930). DNSM-PQZ6956 43.0 44.0 (1.02) 7.5 (0.17) 15.5 (0.36) Subfamily Pitarinae Stewart, 1930 Genus Pitar Römer, 1857 Description Subgenus Pitarina Jukes-Browne, 1913 Medium sized (Lmax = 46 mm), subcircular, srongly compressed, with Type species: Cytherea citrina Lamarck, 1818; by original designation. anteriorly-positioned tiny umbones and subtruncate posterior margin. The flanks are ornamented with strong, fine concentric riblets which Pitarina hebraea (Lamarck, 1818) are broader than the interspaces. Fig. 4F-G
Discussion Cytherea hebraea Lamarck 1818: 578. This is a common and widespread tropical Indo-Pacific species which ranges from southern Moçambique northwards and first appears in Pitar hebraeus (Lamarck) Cox 1930: 129, pl. 15, fig. 8; Kilburn & Rippey the Pliocene (Cox 1930). Circe jucunda Smith, C. nana Melvill and C . 1982: 217. salamensis Jaeckel & Thiele all have radial costellae, and C. corrugata (Dilwyn) is more inflated, with coarser concentric ornament. Circe non Meretrix hebraea (Lamarck) Franca 1960: 578, 95, pl. 26, fig. 6 (= P. mombasana Cox (1930: 157, pl. 15, figs 6-7) is a Comus. abbreviatus). non Pitaria hebraea (Lamarck) Boshoff 1965: 164, pl. 10, fig. 4 (= P. Genus Gafrarium Röding, 1798 abbreviatus). Type species: Venus pectinata Linné, 1758; by the subsequent designation of Dall (1902). non Pitaria hebrea (Lamarck) Moura 1968: 49, pl. 10, fig. 2; 1971: 60, pl. 11, fig. 3a, b (= P. abbreviatus). Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 19
Fig. 7. Reconstructions of some of the fossils described herein, to better illustrate their characters. A-C. Anadara jacksoni sp. nov. D. Decatopecten amiculum (Philippi). E. Naticarius alapapilionis (Röding). F. Gafrarium pectinatum (Linné). G-H. Canarium mutabilis (Swainson). I. Ovatipsa chinensis violacea (Rous). J. Gari maculosa (Lamarck). K. Timoclea costellifera (Adams & Reeve). 20 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
Pitar hebraea (Lamarck) Bosch et al. 1995: 270, fig. 1211. Discussion The False Bay fossils compare well with living T. costellifera. Another Material Indo-Pacific species, T. marica (Linné), differs in being trigonally-ovate, Two specimens, DNSM-PQZ5876 and 5878. more narrowly-rounded at both ends, with more-prominent umbones and divaricating ribs on the posterior flank; Cox (1930) recorded Measurements this species from the Pleistocene of Kenya. Timoclea arakana (Nevill) L H T AL is shorter and squatter, with a pointed posterior and strengthened DNSM-PQZ5878 20 16.2 (0.81) 8 (0.40) 5 (0.25) radial ribbing both anteriorly and posteriorly; T. lavrani Fischer-Piette is higher, the ribs not obviously nodate, and it has a subtruncate Description posterior margin and smooth unornamented patch on the anterior Small, subquadratic, longer than high, rather inflated, with prominent flank. Timoclea subnodulosa (Hanley) has a subtrigonal profile and anteriorly-positioned incurved umbones. The anterior and posterior divaricating costae. margins are broadly rounded. and there is a relatively large cordate lunule which is raised and tent like. Ornament comprises growth Superfamily TELLINOIDEA Blainville, 1814 striae only. Internal features unknown. Family Psammobiidae Fleming, 1828 Subfamily Psammobiinae Fleming, 1828 Discussion Genus Gari Schumacher, 1817 A widespread living Indo-Pacific species which ranges as far south Type species: Gari vulgaris Schumacher, 1817, = Solen ameythystus of northern Moçambique (Kilburn & Rippey 1982). The specimen Wood, 1815; by original designation. figured by Cox (1930) from the Pliocene of Kenya is more-produced posteriorly. Extant P. abbreviatus (Krauss) (Kilburn & Rippey 1982: Gari maculosa (Lamarck, 1818) 194, pl. 45, fig. 1, text-fig. 207) which lives off the coast of KwaZulu- Fig. 4Q-T, 7J Natal is larger, ovate, with subcentral umbones and a sunken lunule. Pitar madecassinus (Fischer-Piette & Delmas) is trigonally ovate, with a Psammobia maculosa Lamarck, Dautzenberg & Fischer 1914: 215, pl. 6, weak umbonal ridge and angulated posterior. Pitar affinis (Gmelin), of figs 1-3. which. P. prora Lamprell & Kilburn (1999) is a junior synonym (Huber 2010), is shorter and subtrigonal. Pitar noguchi Habe has prominent Gari maculosa (Lamarck) Prashad 1932: 298; Kilburn & Rippey 1982: umbones, a more-concave anterodorsal margin and slightly broader 190; Taylor & Glover 2004: 263; Bosch et al. 1995: 260, fig. 1163; Taylor more-upright posterior margin. Pitar kathiewayae Lamprell & Kilburn & Glover 2004: 263. is trigonally ovate, more inflated, and with high umbones positioned further from the anterior (AL = 32-36%). non Psammobia maculosa Lamarck, Barnard 1964: 533 ( = Gari depressa). Subfamily Chioniinae Frizzell, 1936 Genus Timoclea Brown, 1827 Material Type species: Venus ovata Pennant, 1777; by monotypy. Three specimens, DNSM-PQZ5863, 5905 and 7010.
Timoclea costellifera (Adams & Reeve, 1850) Measurements Fig. 4W-X, 7K L H T AL DNSM-PQZ5863 46.7 25 (0.54) 7 (0.15) 19 (0.41) Timoclea costellifera (Adams & Reeve) Springsteen & Leobrera 1986: DNSM-PQZ5905 48.9 22.8 (0.51) 6.0 (0.12) 21 (0.43) 305, pl. 86; Bosch et al. 1995: 266, fig. 1194. Description Material Medium sized (Lmax = 50 mm), inequilateral, subelliptical, much Three specimens, DNSM-PQZ5907 and 7011-12. longer than high (H/L = 0.52-0.54), with low insignificant subcentral umbones (40-43% of shell length from anterior margin), and weakly- Measurements inflated valves (W/H = 0.21-0.29). The anterodorsal margin is almost L H T AL straight, the anterior margin well rounded, the broad ventral margin DNSM-PQZ7011 25.5 20.0 (0.78) 6.7 (0.26) 10.0 (0.39) gently convex, the broad respiratory margin is obliquely subtruncate, and the posterodorsal margin almost straight. The dorsal commissure Description shows the weak lateral flexure characteristic of the genus. A weak Small, inequilateral, subovate, slightly longer than high, with small umbonal ridge separated the flank from a trigonal posterior area. rounded subcentral umbones, weakly-inflated valves, broadly-rounded The flanks are ornamented with fine oblique costellae which cut at anterior margin and somewhat narrower posterior margin. There is low angle across growth striae and are wider than the interspaces. a prominent swollen raised unornamented lanceolate lunule. Flank Posteriorly these costellae become irregular and may join, being ornament is cancellate, with about 28 strong narrow concentric replaced by coarser more-irregular growth wrinkles which parallel costellae, narrower than the interspaces, superimposed with about the respiratory margin and ornament the posterior area. 30 radial costellae which broaden ventrally and anteriorly but not posteriorly, and are wider than the interspaces. The junction Discussion between the two sets of ribs is marked by a prominent node, and The False Bay material displays the essential characters of G. maculosa the interspaces by a pit. Although the extreme ventral margin is (Lamarck), a widespread Indo-Pacific species which extends as far not preserved in any of the specimens, there is no indication of south as Durban. Gari anomala (Deshayes) is smaller, with a vertically divarication. truncate respiratory margin and lacks growth wrinkles. Gari elongata Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 21
Lamarck lacks oblique costellae and growth wrinkles, as do G . 1995: 73, fig. 247. amethysta (Wood), G. depressa (Pennant), G. convexa (Reeve) and G . hodgei Willan. Gari pennata (Deshayes) is a much smaller species with Material coarse divaricating ribs anteriorly. A single specimen, DNSM-PQZ5882.
Class GASTROPODA Cuvier, 1797 Measurements Superfamily STROMBOIDEA Rafinesque, 1815 L W T Family Strombidae Rafinesque, 1815 DNSM-PQZ5882 35 21.6 (0.62) 17.1 (0.49) Genus Canarium Schumacher, 1817 Type species: Strombus urceus Linné, 1758; by original designation. Description Small, ovoid, base longer than wide, with a moderately convex dorsum. Canarium mutabilis (Swainson, 1821) Left margin more convex than right, straightening somewhat anteriorly. Fig. 6C-D, 7G-H Anterior end narrow. Apertural slit convexoconcave, of more-or-less the same width along the length of the shell (maximum width 14% of Strombus floridus Lamarck, Braga 1952: 85, pl. 5, fig. 2. shell width), remaining relatively narrow anteriorly and not reflexed at the siphonal canal. Long fine columellar teeth (~20), wider than Strombus mutabilis Swainson, Barnard 1964: 65; Kilburn & Rippey 1972: the interspaces, extend well onto the columellar callus, remaining 58, pl. 12, fig. 13; Kensley 1973: 117, fig. 398. relatively uniform along the length of the shell, and outnumber the 15 relatively coarse labral teeth. Fossula not displayed. Strombus (Canarium) mutabilis Swainson, Ladd 1972: 58, pl. 15, figs 10- 15; Bosch et al. 1995: 61, fig. 200. Discussion The False Bay fossil is a good match for living eastern African material Material which is assigned to the subspecies violacea (Rous). The nominate Two specimens, DNSM-PQZ5912 and 7016. subspecies ranges as far south as Jeffrey’s Bay.
Description Superfamily NATICOIDEA Forbes, 1838 Small, biconical, longer than wide, with moderately high spire (38% of Family Naticidae Forbes, 1838 length), acute apical angle, angulated shoulder with 14 strong radially- Subfamily Naticinae Forbes, 1838 elongate tubercles per whorl, and a narrow, flat, steeply-inclined Genus Naticarius Duméril, 1805 adapical face. The upper one-third of the body whorl has flat parallel Type species: Nerita canrena Linné, 1758; by monotypy. flanks below which they converge strongly to the siphonal canal. The convergent lower part of the body whorl is ornamented with thin Naticarius alapapilionis (Röding, 1798) distant spiral grooves, the adoral eight of which are closely spaced Fig. 6G-I, 7E and deeply impressed. There is a broad smooth columella callus to the inner lip and a thin broken outer lip. Natica alapapilionis (Röding) Kilburn 1976: 843, fig. 6 (cum. synon.); Kilburn & Rippey 1982: 70, pl. 16, fig. 1. Discussion A common and widespread Indo-Pacific species which ranges from Natica (Naticarius) alapapillionis (Röding) Bosch et al. 1995: 87, fig. 323. Jeffrey’s Bay northwards, with a stratigraphic range of Pliocene to Recent (Ladd 1972). Similar Dolomena plicata (Röding), recorded from Material the Late Pliocene to Recent of the Indo-Pacific province (Cox 1930), Two specimens, DNSM-PQZ5881 and 5883. has a higher spire with slightly concave sides and 25 or more axial ribs. Other strombids recorded fossil from southern and eastern Measurements Africa include Tridentarius dentatus (Linné) (= Strombus tridentatus L W T Gmelin) from the Pleistocene of Zanzibar (Cox 1927: 86, pl. 18, fig. 5) DNSM-PQZ5881 33 28.5 (0.86) 18 (0.55) and Gibberulus gibberulus (Linné) (Cox 1930: 138) from the Pleistocene of Kenya. The latter also occurs in the Isipingo Formation of Durban, Description a unit correlated with the False Bay Coral Limestone (Cooper & Liu Small, globular, longer than wide, moderately convex, with an elongate 2008). base, broadly-convex dorsum and low obtuse spire whose sutures are obscured by callus. The semicircular aperture has a long straight Superfamily CYPRAEOIDEA Rafinesque, 1815 smooth slightly-inclined inner lip and unornamented thin convex outer Family Cypraeidae Rafinesque, 1815 lip. The narrowly-lunate umbilicus is entered by an oblique chord. Subfamily Erroneinae Schilder, 1927 Ornament is lacking save for growth striae. Tribe Bistolidini Meyer, 2003 Genus Ovatipsa Iredale, 1931 Discussion Type species. Cypraea chinensis Gmelin, 1791; by original designation. Naticarius alapapilionis (Röding) is a widespread Indo-Pacific species which also has been assigned to the subgenus Glyphepithema. In South Ovatipsa chinensis (Gmelin, 1791) Africa it ranges as far south as the Transkei of the Eastern Cape. Fig. 6E-F, 7I Natica cernica Jousseaume is smaller, shorter and squatter, with a more-inclined inner lip, and N. arachnoidea (Gmelin) is shorter still. Cypraea chinensis Gmelin, Barnard 1964: 47; Kilburn & Rippey 1972: 68, Naticarius onca (Röding) (Cernohorsky 1971: 186, figs 33-35) is a pl. 15, fig. 6; Kensley 1973: 343, fig. 347; Oliver 1980:92, fig.; Bosch et al. widespread Indo-Pacific species which is wider than long (W/H = 22 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
0.93-1.05), with a plugged umbilicus and higher spire. Naticarius A single specimen, DNSM-PQZ5911. orientalis (Gmelin) (Kilburn 1976: 844, fig. 6 (right)) is a shorter squatter species, also wider than high, with a wide-open umbilicus, Description higher spire, rounder outer lip and pointed anterior. Naticarius Large (>150 mm high), robust, fusiform, high spired, with 7-8 swollen hebraea (Martyn) is a shorter squatter species with oblique inner tubercles to the angulated shoulder. lip and gaping umbilicus. The fossil material differs from N. maurus in having a curved inner lip. Discussion This is a common and widespread Indo-Pacific species. It is abundant Subfamily Polinicinae Gray, 1847 in Recent (Flandrian) sediments of southern Moçambique (Moura Genus Glossaulax Pilsbry, 1929 1966, 1968, 1970, 1971). Type species: Natica reclusianus Deshayes, 1839; by original designation. ACKNOWLEDGEMENTS
Glossaulax didyma (Röding, 1798) I should like to thank Mrs. E.H. Greyling for assistance both in the Fig. 6J field and laboratory, and Dr Martin Zuschin (Vienna) for a helpful review of the manuscript. Natica lamarckii Réclus, Krauss 1848: 91. REFERENCES Natica didyma (Röding) Turton 1932: 159. ABBOTT, R.T. & DANCE, S.P. 1982. Compendium of seashells. Dutton. Natica (Neverita) didyma (Bolton) Cox 1939: 96. BARNARD, K.H. 1963. Contributions to the knowledge of South African marine Mollusca. Part Ill. Gastropoda: Prosobranchiata: Natica ampla Philippi, Braga 1952: 89, pl. 5, fig. 10. Taenioglossa. Annals of the South African Museum 47(1-2): 1-199. BARNARD, K.H. 1964. Contributions to the knowledge of South Polynices ampla (Philippi) Franca 1960: 62, pl. 4, fig. 7. African marine Mollusca. Part V. Lamellibranchiata. Annals of the South African Museum 47 (3): 361-593. Polynices didyma (Bolten) Barnard 1963: 63. BARTSCH, P. 1915. Report on the Turton Collection of South African marine mollusks, with additional notes on other South African Polynices didyma (Bolton) Kensley 1973: 98, fig. 331. shells contained in the United States National Museum. Bulletin of the Smithsonian Institution, United States National Museum 91: 1-305. Polinices (Glossaulax) didyma (Röding) Kilburn 1976: 860. BERNARD, F.R., CAI, Y.Y. & MORTON, B. 1993. Catalogue of the living marine bivalve molluscs of China. Hong Kong University Press: Hong Polinices didyma (Röding) Kilburn & Rippey 1982: 71, pl. 16, fig. 5. Kong. BOSCH, D.T., DANCE, S.P., MOOLENBEECK, R.G. & OLIVER, P.G. Neverita (Glossaulax) didyma (Röding) Bosch et al. 1995: 86, fig. 315. 1995. Seashells of eastern Arabia. Motivate Publishing: Dubai. BOSCHOFF, P.H. 1965. Pelecypoda of Inhaca Island, Moçambique. Material Memórias do Instituto de Investigaçao Cientifica de Moçambique, Série A single specimen, DNSM-PQZ5884. A (Ciências biologicas) 7: 65-206. BRAGA, J.M. 1952. Materiais para estudo da fauna malacológica de Discussion Moçambique. Trahalho da Missuo Zoologica de Moçambique 7(3): The fossil is poorly preserved, but large size (W = 39 mm), very 64-125. low spire and thick test all suggest reference to this widespread and CERNOHORSKY, W.O. 1971. The family Naticidae in the Fiji Islands. common Indo-Pacific species which ranges from Miocene to Recent Records of the Aukland Institute and Museum 8: 169-207. (Cox 1939) and extends as far south as Still Bay.. COOPER, M.R. 1991. Quaternary Mollusca from the Port Durnford Formation. South African Journal of Geology 93(3): 538-546. Superfamily BUCCINOIDEA Rafinesque, 1815 COOPER, M.R. 1999. The Cainozoic palaeontology and stratigraphy Family Fasciolariidae Rafinesque, 1815 of KwaZulu-Natal. Part 3. The Mduku Formation. Stratigraphy and Genus Pleuroploca Fischer, 1884 fauna. Durban Museum Novitates 24: 48-56. Type species: Fasciolaria trapezium Linné, 1758; by original designation. COOPER, M.R. & LIU, K. 2006. The Cainozoic palaeontology and stratigraphy of KwaZulu-Natal. Part 4. The post-Karoo geology of Pleuroploca trapezium (Linné, 1758) the Durban area, with special reference to the Isipingo Formation. Fig. 6A-B Durban Museum Novitates 21: 1-23. COOPER, M.R. & MCCARTHY, M.J. 1998. The Cainozoic Fasciolaria trapezium Linnaeus, Braga 1952: 73, pl. 2, fig. 1; Franca 1960: palaeontology and stratigraphy of KwaZulu-Natal. Part 2. 79, pl. 13, fig. 2; Kensley 1973: 164, fig. 611. Stratigraphy of the Uloa Subgroup. Durban Museum Novitates 23: Fasciolaria (Pleuroploca) trapezium (Lineu) Moura 1966: 43, pl. 6, fig. 2; 3-28. 1968: 42, pl. 8, fig. 1; 1969: 26, pl. 9, fig. 6a, b; 1970: 79, pl. 6, fig. 2; 1971: 43, pl. 7, fig. 3. Pleuroploca trapezium (Linné) Abbott & Dance 1982: 182; Springsteen COX, L.R. 1927. Neogene and Quaternary Mollusca from the & Leobrera 1986: 176, pl. 47; Drivas & Jay 1987: 84, pl. 27; Bosch et al. Zanzibar Protectorate: 13-102. In: STOCKLEY, G. M., ed., Report 1995: 136, fig. 577. on the palaeontology of the Zanzibar Protectorate. Government of Zanzibar: Dar es Salaam. Material COX, L.R. 1929. Moluscos miocénicos, pliocénicos e post pliocénicos Durban Natural Science Museum Novitates 37 FALSE BAY CORAL LIMESTONE FORMATION 23
de Moçambique. Boletim Serviços de Industria, Minas e Geologia, coastal caves in the Cape Province, South Africa. Quaternary Colónia de Moçambique 3: 65-102 (Portuguese), 103-163 (English). Research 25: 189-198. COX, L.R. 1929b. Notes on the post-Miocene Ostreidae and HOBDAY, D.K. & ORME, A.R. 1974. The Port Durnford Formation: Pectinidae of the Red Sea region, with remarks on the geological a major Pleistocene barrier-lagoon complex along the Zululand significance of their distribution. Proceedings of the malacological coast. Transactions of the geological Society of South Africa 77(2): Society 18(4): 165-209. 141-149. COX, L.R. 1930. Reports on geological collections from the coastlands HOBDAY, D.K. 1976. Quaternary sedimentation and development of of Kenya made by Miss M. McKinnon-Wood. V. Miocene, Pliocene the lagoonal complex, Lake St Lucia, Zululand. Annals of the South and post-Pliocene Mollusca. Monograph of the Hunterian Museum, African Museum 71: 93-113. Glasgow 4: 103-163. HUBER, M. 2010. Compendium of bivalves. A full-colour guide to 3,300 COX, L.R. 1931. The geology of the Farsan Islands, Oizan and Kamaran of the world’s marine bivalves. A status on Bivalvia after 250 years of Islands, Red Sea. Geological Magazine 68(1): 1-13. research. Conchbooks: Hackenheim.. COX, L.R. 1936. Fossil Mollusca from southern Persia (Iran) and KENSLEY, B.F. 1973. Seashells of southern Africa. Gastropods. Maskew Bahrein Island. Palaeontologia indica, Memoirs of the geological Survey Millar: Cape Town. of India, n.s., 22(2): 1-69. KILBURN, R.N. 1976. A revision of the Naticidae of southern Africa COX, L.R.1939. Depósitos terciários e post-pliocenos do distrito de and Moçambique (Mollusca). Annals of the Natal Museum 22(3): Inhambane. Estudo paleontologico de moluscos. Boletim Serviços 829-884. de Industria, Minas e Geologia, Colónia de Moçambique 3: 21-58 (65- KILBURN, R.N. 1983. The Recent Arcidae (Mollusca: Bivalvia) of 103). southern Africa and Mozambique. Annals of the Natal Museum COX, L.R. et al. 1969. Treatise on invertebrate paleontology. Part N, 25(2): 511-548. Volume 2 (of 3), Mollusca 6, Bivalvia: N670-690. Geological Society KILBURN, R.N. 1997. The family Ungulinidae in Southern Africa and of America, University of Kansas Press: Lawrence. Mozambique. Annals of the Natal Museum 37: 267-286. DANCE, S.P. & EAMES, F.E. 1966. New molluscs from the Recent KILBURN, R.N. & RIPPEY, E. 1982. Sea shells of southern Africa. Hammar Formation of south-east Iraq. Proceedings of the Macmillan South Africa: Johannesburg. malacological Society of London 37: 35-43. KIRA, T. 1962. Shells of the Western Pacific in colour. Hoikushu Publishing DAUTZNBERG, P. & FISCHER, H. 1914. Sur quelques types de garidés Company: Osaka. de la collection de Lamarck existant au museum. Journal de KRAUSS, F. 1848. Die südafrikanischen Mollusken. Ein Beitrag zur Conchyliologie 611: 215-228. Kenntniss der Mollusken des Kap- und Natallandes und zur DAUTZENBERG, P. 1929. Mollusques testacés marins de geographischen Verbreitung derselben, mit Beschreibung und Abbildung Madgascar. Faune des Colonies Françaises 3, Société d’editions der neuen Arten. Ebner & Seubert: Stuttgart. Géographiques, Maritimes et Coloniales: Paris. LADD, H.S. 1972. Cenozoic fossil mollusks from western Pacific DIJKSTRA, H.H. & KILBURN, R.N. 2001. The family Pectinidae in South islands; Gastropoda (Turritellidae through Strombidae). Professional Africa and Mozambique (Mollusca: Bivalvia: Pectinoidea). African Paper, United States geological Survey 532: 1-79. Invertebrates 42: 263–321. LAMARCK, J.P.B. 1818-19. Histoire naturelle des animaux sans vertèbres. DIJKSTRA, H.H. & KNUDSEN, J. 1998. Some Pectinoidea (Mollusca: 5-7. Paris. Bivalvia: Propeamussiidae: Pectinidae) of the Red Sea. Molluscan LAM, K & MORTON, B. 2004. The oysters of Hong Kong (Bivalvia: Research 19: 43–104. Ostreidae and Gryphaeidae). The Raffles Bulletin of Zoology 51(1): DRIVAS, J. & JAY, M. 1988.Coquillages de la Réunion et de l’ile Maurice. 11-28. Les Editions du Pacifique: Singapore. LAMPRELL, K. & KILBURN, R.N. 1999.The genera Lioconcha and Pitar DUNKER, W. 1877. Mollusca nonnulla nova maris Japonici. in South Africa and Mozambique, with descriptions of three new Malakozoologische Blätter 24: 67-75. species (Mollusca: Bivalvia: Veneridae). Vita Marina 46(1-2.): 19-41. DUNKER, W. 1882. Index molluscorum maris Japonici. Theodor Fischer: LISIECKI, L.E. 2005. A Pliocene-Pleistocene stack of 57 globally Cassel. distributed benthic 18O records. Paleoceanography 20. EAMES, F.E. 1950. Notes on some Caenozoic molluscan species from the doi:10.1029/2004PA001071 Far East. Proceedings of the malacological Society 28(4-5): 145-155. LIU, L., MAIORANO, P. & ZHAO, X. 1996. Pliocene-Pleistocene EAMES, F.E. & COX, L.R. 1956. Some Tertiary Pectinacea from East calcareous nannofossils froim the Iberian abyssal plain. In: Africa, Persia and the Mediterranean region. Proceedings of the WHITMARSH, R.B., SAWYER, D.S., KLAUS, A. & MASSON, D.G. malacological Society of London 32: 1-69. (eds), Proceedings of the Ocean Drilling Program, Scientific Results 149: FRANCA, M. DE L. PAES DA. 1960. Sobre uma colecção malacológica 147-164. recolhida na Ilha da Inhaca (Moçambique). Memórias da Junta de MCCARTHY, M.J. 1988. Late-Gondwana to Recent geology of the Investigações do Ultramar 15: 43-102. Natal north coast. Excursion Guide to geocongress ’88: 1-45. HARRY, H.W. 1985. Synopsis of the supraspecific classification of living Geological Society of South Africa: Durban. oysters (Bivalvia: Gryphaeidae and Ostreidae). The Veliger 28: 121- MOURA, A.R. 1966. Gastropodes dos recifes corais elevados de 158. Messouta e Matibane (Moçambique). Revista dos Estudos Gerais HASAN, A.K. 1994. A taxonomic review of the bivalve and gastropod Universitarios de Moçambique (2) 3: 15-54. mollusc fauna along the Saudi intertidal zone of the Arabian Gulf. MOURA A.R. 1972. Contribuçao para o conhecimento dos moluscos Special Issue, Symposium on the Red Sea marine environment, Jeddah, subfosseis do Relanzapo (Nacala – Moçambique). Revista de 1994: 245-253. Ciencias Biologicas, Lourenço Marques (A) 5: 9-66. HENDEY, Q.B. & COOKE, H.B.S. 1985. Kolpochoerus paiceae MOURA A.R. 1976. Contribuçao para o conhecimento da fauna (Mammalia: Suidae) from Skurweberg, near Saldanha, South Africa, malacologica subfossil de Bartolomeo Dias (Inhassoro – and its palaeoenvironmental implications. Annals of the South African Moçambique). Memórias de Instituto de Investigação Científica de Museum 97(2): 9-56. Moçambique 12: 41-87. HENDEY, Q.B. & VOLMAN, T.P. 1986. Last interglacial sea levels and MOURA, A.R. 1968. Molluscos des grés costeiras da Ilha de Inhaca 24 FALSE BAY CORAL LIMESTONE FORMATION Durban Natural Science Museum Novitates 37
(Moçambique). Revista Ciencias Biologicos, Lourenco Marques (A) 1: SACS 1980. Stratigraphy of South Africa. Part 1: Lithostratigraphy of 17-52. the Republic of South Africa, South West Africa/Namibia and the MOURA, A.R. 1969. Ilha de Inhaca, Moçambique. Trabalhos da Institut republics of Bophuthatswana, Transkei and Venda. Handbook of the de Investigações Cientifica do Moçambique 23: 1-20. geological Survey of South Africa 8: 1-690. MOURA, A.R. 1970. Contribução para o conhecimento dos molluscos SALVAT, B. & REEVES, C. 1983. Coquillages de Polynésie. Les Editions du subfosseis do Mucoque (Vilanculos). Revista Ciencias Biologicos, Pacifique: Honalulu. Lourenco Marques (A) 3: 55-96. SALVAT, B. & REEVES, C. 1991. Shells of Tahiti. 4 th ed. Les Editions du MOURA, A.R. 1971. Contribução para o conhecimento dos molluscos Pacifique: Honalulu. subfosseis do Relanzapo (Nacala – Moçambique). Revista Ciencias SIDDIQUI, G. & AHMED, M. 2002. Oyster species of the subtropical Biologicos, Lourenco Marques (A) 4: 9-66. coast of Pakistan (northern Arabian Sea). Indian Journal of Marine NARDINI, S. 1937. Molluschi della spiagge emerse del mar Rosso e Science 31(2): 108-118. dell’Oceano Indiano. I. Lamellibranchi. Palaeontographica italica 37: SPRINGSTEEN, F.J. & LEOBRERA, F.M. 1986. Shells of the Philippines. 225-298. Carfel Shell Museum: Manila. OLIVER, A.P.H. 1975. The Hamlyn guide to shells of the World. Hamlyn: SPRY, J.F. 1964. The sea shells of Dar es Salaam: Part 2: Pelecypoda N.Y., Sidney, Toronto. (Bivalves). Tanganyika Notes and Records 63: 1-52. OLIVER, P.G. 1992. Bivalved seashells of the Red Sea. Verlag Christa MURUGAN, A. & VIJAYALAKSHIMI, S. 2010. New distributional Hemmer, National Museum of Wales. records of Hyotissa hyotis (Linnaeus, 1758): Gryphaeidae from ORME, A.R. 1973. Barrier and lagoon systems along the Zululand Mandapam area – southeast coast of India. World Journal of Fish coast, South Africa: 181-217. In: COATES, D.R. (ed.), Coastal and Marine Sciences 2(1): 42-43. geomorphology. State University of New York: N. Y. TAYLOR, J.D. & GLOVER, E.A. 2003. Diversity and distribution of PARKINSON, B. J. 1982. The shell resources of Fiji. South Pacific subtidal benthic molluscs from the Dampier Archipelago, Western Commission: Noumea. Australia; results of the 1999 dredge survey (DA2/99). Records of PAULAY, G. 1996. New records and synonymies of Hawaian bivalves the Western Australian Museum, Supplement 66: 247-291. (Mollusca). Occasional Papers of the Bishop Museum 45: 18-29. THIELE, J. & JAECKEL, S. 1931. Muscheln der Deutschen Tiefsee POUTIERS, J.M. 1998 Bivalves. Acephala, Lamellibranchia, Pelecypoda: Expedition. Wissenschafliche Ergebnisse der deutschen Tiefsee- 123 - 362. In: CARPENTER, K.E. & NIEM, V.H. 1998. FAO species Expedition auf des Dampfer Valdivia 21: 161-268. identification guide for fishery purposes. The living marine resources of TURTON, W.H. 1932. The marine shells of Port Alfred, South Africa. the Western Central Pacific. Volume 1. Seaweeds, corals, bivalves, and Oxford University Press: Oxford. gastropods. Rome, FAO WALLER, T.R. 1993. The evolution of “Chlamys” (Mollusca: Bivalvia: PRASHAD, B. 1932. The Lamellibranchia of the Siboga Expedition. Pectinidae) in the tropical western Atlantic and eastern Pacific. III. Systematic part 2. Pelecypoda (exclusive of the Pectinidae). American malacological Bulletin 10(2): 195-249. Monograph of the Siboga Expedition 53(C): 1-354. WALLS, J.G. 1980. Conchs, tibias and harps. T.F.H. Publications: Hong RIPPINGGALE, O.H. & MCMICHAEL, D.F. 1961. Queensland and Great Kong. Barrier Reef shells. Jacaranda Press: Brisbane. Durban Natural Science Museum Novitates 37 DRUG RESPONSE OF CHRYSOMYA CHLOROPYGA 25
EFFECTS OF HYDROCORTISONE AND SODIUM METHOHEXITAL ON GROWTH RATE OF CHRYSOMYA CHLOROPYGA WEIDEMANN (DIPTERA: CALLIPHORIDAE): DEVELOPMENTAL AND BEHAVIOURAL INDICATIONS OF PRESENCE OF DRUGS
K.A. WILLIAMS* & M.H. VILLET Department of Zoology and Entomology, Rhodes University, Grahamstown 6140, South Africa *Current address: Entomology Department, Durban Natural Science Museum, P.O. Box 4085, Durban 4000, South Africa *Author for correspondence: [email protected]
illiams, K.A. & Villet, M.H. 2014. Effects of hydrocortisone and sodium methohexital on growth rate of Chrysomya chloropyga Wweidemann (Diptera: Calliphoridae): developmental and behavioural indications of presence of drugs. Durban Natural Science Museum Novitates 37: 25-29. Larvae of the blowfly Chrysomya chloropyga Weidemann were reared on chicken liver dosed with either a steroid or a barbiturate at concentrations that would be half the median lethal dose (MLD), the MLD and twice the MLD for humans. No significant differences were observed between dosages in terms of durations of any stage of development or total development or mortality for larvae exposed to either drug. These results show a different trend to those obtained using Sarcophaga tibialis Macquart, suggesting that extrapolations from one taxonomic family or species to another are unreliable. Activity levels of larvae exposed to the barbiturate and the steroid were significantly increased from the control on Days 1 and 2 of the experiments, indicating stimulation experienced by the larvae in the presence of the drugs. Uncharacteristic behaviours of maggots may thus indicate the presence of drugs, providing toxicological investigations with a new tool in the form of behavioural bioassays.
KEYWORDS: behaviour, behavioural bioassay, Calliphoridae, Chrysomya chloropyga, development, post mortem intervals.
INTRODUCTION de Carvalho 2010; Rivers & Dahlem 2014). Medico-criminal forensic entomologists use larval development to estimate the duration The use of insects, especially maggots, as toxicological material for of the postmortem interval (PMI) in criminal investigations (Catts the detection of drugs and toxins has received substantial attention 1992; Rivers & Dahlem 2014). When estimating a PMI, it is assumed (Beyer et al. 1980; Catts & Goff 1992; Wilson et al. 1993; Goff & Lord that during the first few weeks of decay, insects (especially flies) will 1994, 2010; Tracqui et al. 2004), as have the effects of these substances develop at predicable rates under the prevailing climatic conditions. on the development rate of dipteran larvae (Goff & Lord 1994; Lopes This may not always be true if drugs (or their metabolites) are present 26 DRUG RESPONSE OF CHRYSOMYA CHLOROPYGA Durban Natural Science Museum Novitates 37
Fig. 1. Median age (with range and interquartile range) at which larval feeding, larvae wandering and the pupariation stage end in Chrysomya chloropyga when exposed to four levels of (a) hydrocortisone or (b) sodium methohexital contamination in their larval diet. No significant differences between concentrations were found for any event (Table 2).
(Goff & Lord 1994, 2010; Lopes de Carvalho 2010). MATERIALS AND METHODS This study examined the effects of sodium methohexital (a barbiturate) and hydrocortisone (a steroid) on the developmental A culture of C. chloropyga was bred from wild flies trapped on carrion rate of the blowfly Chrysomya chloropyga Weidemann. Our primary in Grahamstown, South Africa. The culture was kept in a constant reason for using these drugs was that sodium methohexital is a environment room at 25°C with a photoperiod of 12L: 12D hours. metabolic depressant, while hydrocortisone is a metabolic stimulant, The flies were fed on sugar, milk powder and water ad libitum and and they may therefore have pronounced and opposite effects on fresh chicken liver was provided prior to the start of the experiment development (Musvasva et al. 2001). Furthermore, previous work on as a protein meal for the flies. The flies were allowed to lay eggs on the same drugs (Musvasva et al. 2001) used a flesh fly (Sarcophagidae) the liver and when eggs were discovered they were removed with a and it would be useful to establish if different species of fly respond small portion of liver. When they hatched, the first instar larvae were to the same drugs similarly. Barbiturates are potentially dependence- transferred to the experimental jars. producing drugs and are therefore likely to be abused, and can Three concentrations (3 μl/g, 6 μl/g and 12 μl/g) of occur in large quantities in a body (Foye 1976). Steroids are used hydrocortisone were prepared using 5 ml of 0.9% saline and 50 g of to treat a variety of common inflammatory conditions, but are not chopped chicken liver, and placed in 150 ml glass bottles. Solutions of usually abused. Chrysomya chloropyga was chosen because it is a sodium methohexital (50 μg/g, 100 μg/g and 200 μg/g) were also made blowfly (Calliphoridae), and one of the first and commonest insects using 5 ml of 0.9% saline and 50 g of chopped chicken liver, and placed to colonise dead bodies in southern Africa, which gives it forensic in 150 ml glass bottles. These concentrations were half of, equal to, significance. or double the median lethal dose (MLD) for mammals (S. Daya, pers.
Fig. 2. Mean locomotion rates (with standard error and standard deviation) of C. chloropyga larvae exposed to increasing concentrations of (a) hydrocortisone and (b) sodium methohexital. Durban Natural Science Museum Novitates 37 DRUG RESPONSE OF CHRYSOMYA CHLOROPYGA 27
Fig. 3. Mean activity levels (with standard error and standard deviation) of C. chloropyga larvae exposed to increasing concentrations of (a) hydrocortisone and (b) sodium methohexital.
Table 1. Mean (and standard deviation) of survival rate (%) of Chrysomya chloropyga larvae exposed to increasing concentrations of hydrocortisone and sodium methohexital. Differences between concentrations in each row were not statistically significant (Kruskal-Wallis tests, all p > 0.39). MLD = median lethal dose.
DRUG CONCENTRATION CONTROL 0.5 MLD MLD 2 MLD Hydrocortisone Feeding phase 100 ± 0.0 100 ± 0.0 100 ± 0.0 100 ± 0.0 Wandering phase 100 ± 0.0 100 ± 0.0 100 ± 0.0 100 ± 0.0 Pupal phase 94 ± 9.0 88 ± 12.5 96 ± 6.1 93 ± 6.2 Total 94 ± 9.0 88 ± 12.5 96 ± 6.1 93 ± 6.2
Sodium methohexital Feeding phase 100 ± 0.0 100 ± 0.0 100 ± 0.0 100 ± 0.0 Wandering phase 100 ± 0.0 100 ± 0.0 96 ± 8.9 100 ± 0.0 Pupal phase 94 ± 9.0 89 ± 13.6 75 ± 35.3 59 ± 46.9 Total 94 ± 9.0 89 ± 13.6 74 ± 37.1 59 ± 46.9
comm; Musvasva et al. 2001). It is probable that breakdown products Eclosing flies were provided with liver for feeding and oviposition. of the drugs formed during the course of the experiment due to the Due to significant heteroscedacity (Levene’s test, p < 0.05) or presence of bacteria, enzymes in the chicken liver, and the metabolism relatively small numbers of replicates (n = 5) in a nested sample of the larvae (Goff & Lord 1994). Five replicates of each concentration design, all analyses involved Kruskal-Wallis tests performed on for both drugs were used. Five replicates containing only 5 ml of 0.9% means of replicates. Median tests were used to identify the cause of saline in 50 g of chicken liver were made up in 150 ml glass bottles to significant ANOVA results. act as controls. To preclude the effects of competition and metabolic heating on RESULTS growth (Goodbrod & Goff 1990; Davies & Ratcliffe 1994), only ten larvae were transferred to each bottle, which was then placed in a 2 l No significant differences in survivorship through the larval, wandering tub containing a layer of sand to allow larvae to burrow and pupariate. or puparial phases or in total survivorship for either drug were The tubs were covered with fine netting secured by an elastic band detected (Table 1) despite trends in the data indicating higher to prevent larvae from escaping. The larvae were kept in a controlled mortalities in the sodium methohexital at higher concentrations. No environment room at 25.9° C (SD = 1.69° C). The sand was sieved morphological defects were observed in the adult flies that emerged daily and when larvae were found, they were placed separately in Petri and they were all fecund and fertile, laying eggs that hatched. dishes to complete metamorphosis. The activity levels of larvae on Although trends to faster larval development at higher the second and third day were recorded in two ways: as the number drug concentrations were found (Fig. 1), no significant effects of of larvae that were active on the sides of the container rather than hydrocortisone or sodium methohexital (and/or their possible in the liver; and as the number of locomotory contractions in a five breakdown products (Goff & Lord 1994) were detected in any stage second period. The dates of migration, pupariation, and eclosion were of development (Table 2). recorded for each larva. Deaths at each of these stages were noted. Activity levels, measured as the rate of locomotory contractions, 28 DRUG RESPONSE OF CHRYSOMYA CHLOROPYGA Durban Natural Science Museum Novitates 37
Table 2. Kruskal-Wallis tests of differences in duration of development of Chrysomya chloropyga larvae exposed to increasing concentrations of hydrocortisone and sodium methohexital in their diets.
DRUG DEVELOPMENTAL STAGE H P-VALUE Hydrocortisone Larval feeding stage 7.586 0.0554 Larval wandering stage 0.833 0.8415 Pupariation 1.995 0.5735 Total development 3.217 0.3593
Sodium methohexital Larval feeding stage 6.996 0.0720 Larval wandering stage 0.271 0.9654 Pupariation 2.325 0.5078 Total development 0.660 0.8825 showed no significant differences in the hydrocortisone treatments No significant effects were detected in the length of the puparial
(H3, 60 = 7.148253, p = 0.0673, Fig. 2a), but a significant trend of developmental stage for either drug. By comparison, in S. tibialis the increasing activity with increasing drug concentrations was detected duration of the puparial stage was significantly shorter only at the in the presence of sodium methohexital (H3, 59 = 23.83089, p = 0.0000, lowest concentration of both drugs (Musvasva 2001). The pattern in C . Fig. 2b). However, significantly more immature larvae left the liver chloropyga may be because maggots are able to eliminate, metabolise and wandered around the jar when levels of hydrocortisone were or sequester a wide variety of drugs (Wilson et al. 1993; Sadler et high (H3, 20 = 9.052674, p = 0.0286, Fig. 3a). There was also a trend al. 1995, 1997; Bourel et al. 2001). If the drugs were eliminated or for more larvae to leave the liver as levels of sodium methohexital metabolised before pupariation, they may then not have an effect on increased, but it was not significant (H3, 20 = 7.548892, p = 0.0563, Fig. the developmental rate of pupae. We did not have the opportunity to 3b). assay the puparial remains for drugs, and so cannot tell if they were sequestered in the larval cuticle. However, barbiturates are not always DISCUSSION recoverable from larval tissue (Sandler et al. 1997), and no predictable pattern in this characteristic has been detected (Sandler et al. 1995, This study makes two important findings. First, it confirms that the 1997). same drugs produce different results in different fly species, and that The effects of sodium methohexital and hydrocortisone on larval extrapolations from these results to other families of flies may be developmental rates of C. chloropyga were different to the effects of unreliable. Second, this study provides an example of the potential of the same drugs on S. tibialis (Musvasva et al. 2001). A similar pattern behaviour to provide a bioassay for the presence of drugs or their has been found using heroin and its metabolite morphine: Goff et al. metabolites. (1991) reported an accelerated growth rate in the larval stage of the fleshfly S. peregrine Robineau-Desvoidy in response to cocaine, while Developmental responses to drugs Bourel et al. (1999), reported a retarded growth rate in the blowfly The larval and puparial stages in C. chloropyga were reported by Prins Lucilia sericata Meigen in response to morphine. In another study (1982) as being 162 – 230 and 188 – 204 h, respectively, at a room using L. sericata and codeine, the development rates were shown temperature of 22 - 25° C. In another study the larval stage was to be accelerated and two metabolites of codeine, morphine and approximately 140 hours and the pupal stage 250 hours at 25° C norcodeine, were detected in the larvae (Kharbouche et al. 2008). (Richards et al. 2009). These values are comparable with the control The design of the experiment and the use of live animals or dead results obtained in this experiment, the average temperature of animal tissue as the food source for these experiments may also which was slightly higher (25.9° C). Survivorship at the larval and affect the results due to the presence or absence of metabolites. The pupal stages did not differ significantly from the control. These two issues of whether these trends are widespread, and whether they details imply that the experiment was not confounded by anomalous have a phylogenetic component, need to be clarified. developmental conditions. Development was unaffected by the presence of either Behavioural responses to drugs hydrocortisone or sodium methohexital, even though steroids like Levels of activity of the maggots, as measured by the rate of hydrocortisone are metabolic stimulants (Foye 1976). Contrarily, locomotory contractions, increased significantly with an increase in both drugs significantly delayed larval development by a few hours the concentration of sodium methohexital (Fig. 2b). This agitation at lower concentrations in Sarcophaga tibialis Macquart, but less parallels the number of immature maggots that were counted so at higher concentrations (Musvasva et al. 2001). On the other wandering outside the food (Fig. 3b). On the second day of the hand, barbiturates are metabolic depressants acting on the neural experiment, the number of maggots outside the food was significantly and muscular system in mammals (Foye 1976). Thus the expected different from the control, but by the third day it was no longer effect would be a reduction in the developmental rate. Sodium significant. This suggests that either the sodium methohexital had methohexital’s effects of increasing locomotion rate (Fig. 2) and degraded and that its metabolites were no longer a stimulant to wandering (Fig. 3) were thus not anticipated. It appears that the the maggots, or that the maggots had acclimated to the levels of effects of the drugs are ameliorated by the changes in behaviour of the drug (or both). This response was also observed in the highest the larvae, and we hypothesise that they are leaving the liver to avoid concentration of hydrocortisone, although it was less marked. the drugs. While an effect on behaviour was not persistent in this example, Durban Natural Science Museum Novitates 37 DRUG RESPONSE OF CHRYSOMYA CHLOROPYGA 29
forensic scientists should be aware that unusual necrophage behaviour rate of Boettcherisca peregrina (Diptera, Sarcophagidae) and the may indicate the presence of drugs. The development of behavioural implications of this effect on estimation of postmortem intervals bioassays for drugs would provide forensic science with an additional using arthropod development patterns. Journal of Forensic Sciences tool. 36: 537-542. GOODBROD, J.R. & GOFF, M.L. 1990. Effects of larval population ACKNOWLEDGMENTS density on rates of development and interactions between two species of Chrysomya (Diptera: Calliphoridae) in laboratory We are indebted to Nicola Lunt for technical assistance; Santy Daya culture. Journal of Medical Entomology 27: 339-343. for pharmalogical assistance; Thomas Pape for nomenclatural advice; KHARBOUCHE, H., AUGSBURGER, M., CHERIX, D., SPORKERT, F., Nikite Muller for toxicological advice; and Rhodes University and the GIROUD, C., WYSS, C., CHAMPOD, C. & MANGIN, P. 2008. National Research Foundation for funding. Any opinion, findings and Codeine accumulation and elimination in larvae, pupae, and imago conclusions, or recommendations expressed in this material are those of the blowfly Lucilia sericata and effects on its development. of the authors and the NRF does not accept any liability in that regard. International Journal of Legal Medicine 122: 205-211. LOPES DE CARVALHO, L.M. 2010. Toxicology and forensic REFERENCES entomology. In: Amendt, J., Campobasso, C.P., Goff, M.L. & Grassberger, M. (eds), Current concepts in forensic entomology, p p. BEYER, J.C., ENOS, W.F. & STAJIE, M. 1980. Drug identification through 163-178. Springer: Heidelberg. analysis of maggots. Journal of Forensic Sciences 25: 411-412. MUSVASVA, E., WILLIAMS, K.A., MULLER, W.J. & VILLET, M.H. 2001. BOUREL, B., HEDOUIN, V., MARTIN-BOUYER, L., BECART, A., Preliminary observations on the effects of hydrocortisone and TOURNEL, G., DEVEAUX, M., PHARM, D. & GOSSET, D. 1999. sodium methohexital on development of Sarcophaga (Curranea) Effects of morphine in decomposing bodies on the development tibialis Macquart (Diptera: Sarcophagidae), and implications of of Lucilia sericata (Diptera: Calliphoridae). Journal of Forensic estimating post mortem interval. Forensic Science International 120: Sciences 44: 354-358. 37-41. BOUREL, B., FLEURISSE, L., HÉDOUIN, V., CAILLIEZ, J.C., CREUSY, PRINS, A.J. 1982. Morphological and biological notes on six South C., GOSSET, D. & GOFF, M.L. 2001. Immunohistochemical African blow-flies (Diptera, Calliphoridae) and their immature contribution to the study of morphine metabolism stages. Annals of the South African Museum 90: 201-217. in Calliphoridae larvae and implications in forensic RICHARDS, C.S., CROUS, K.L. & VILLET, M.H. 2009. Models of entomotoxicology. Journal of Forensic Sciences 46: 596-599. development for the blowfly sister species Chrysomya chloropyga CATTS, E.P. 1992. Problems in estimating the postmortem interval in and C. putoria. Medical and Veterinary Entomology 23:56-61. death investigations. Journal of Agricultural Entomology 9: 245-255. RIVERS, D.B. & DAHLEM, G.A. 2014. The science of forensic entomology. CATTS, E.P. & GOFF, M.L. 1992. Forensic entomology in criminal Wiley-Blackwell: New York. 392 pp. investigations. Annual Review of Entomology 37: 253-272. SADLER, D.W., FUKE, C., COURT, F. & POUNDER, D.J. 1995. Drug DAVIES, L. & RATCLIFFE, G.G. 1994. Development rates of some accumulation and elimination in Calliphora vicina larvae. Forensic pre-adult stages in blowflies with reference to low temperatures. Science International 71: 191-197. Medical and Veterinary Entomology 8: 245-254. SADLER, D.W., SENEVIRATNE, C. & POUNDER, D.J. 1997. Effects of FOYE, W.O. 1976. Principles of medicinal chemistry. Lea and Febiger: 3,4-methelenedioxymethamphetamine in decomposing tissues Philadelphia. 489 pp. on the development of Parasarcophaga ruficornis (Diptera: GOFF, M.L. & LORD, W.D. 1994. Entomotoxicology: A new area for Sarcophagidae) and detection of the drug in postmortem blood, forensic investigation. American Journal of Forensic Medicine and liver tissue, larvae and pupae. Journal of Forensic Sciences 42: 1212- Pathology 15: 51-57. 1213. GOFF M.L. & LORD W.D., 2010. Entomotoxicology: insects as TRACQUI, A., KEYSER-TRACQUI, C., KINTZ, P. & LUDES, B. 2004. toxicological indicators and the impact of drugs and toxins on Entomotoxicology for the forensic toxicologist: much ado about insect development. In: BYRD, J.H. & CASTNER, J.L. (eds), Forensic nothing? International Journal of Legal Medicine 118: 194-196. entomology: the utility of arthropods in legal investigations, pp. 427- WILSON, Z., HUBBARD, S. & POUNDER, D.J. 1993. Drug analysis 436. CRC Press: London. in fly larvae. American Journal of Forensic Medicine and Pathology GOFF, M.L. BROWN, W.A., HEWADIKARAM, K.A. & OMORI, A.I. 1991. 14:118-120. Effect of heroin in decomposing tissues on the development 30 OBSERVATIONS ON AFRICAN GRASS LIZARDS Durban Natural Science Museum Novitates 37
OBSERVATIONS ON THE AFRICAN GRASS LIZARDS CHAMAESAURA FITZINGER (REPTILIA: SAURIA: CORDYLIDAE) IN SWAZILAND, WITH EMPHASIS ON FIRE IMPACTS ON POPULATIONS IN MALOLOTJA NATURE RESERVE
RICHARD C. BOYCOTT P.O. Box 5245, Mbabane, Swaziland e-mail: [email protected]
oycott, R.C. 2015. Observations on the African grass lizards Chamaesaura Fitzinger (Reptilia: Sauria: Cordylidae) in Swaziland Bwith emphasis on fire impacts on populations in Malolotja Nature Reserve. Durban Natural Science Museum Novitates 37: 30-39. The distribution, habitat, size and sexual dimorphism of Chamaesaura in Swaziland are described. The effect of fire as an ecological factor on Chamaesaura populations is discussed with emphasis on the populations in Malolotja Nature Reserve, in western Swaziland. Evidence is submitted that could possibly support the hypothesis that the grass-dwelling Chamaesaura represent a group of fire-adapted lizards. It is suggested that appropriate fire management of the grassland habitats where these species occur is likely to enhance their survival.
KEYWORDS: biology, conservation, Chamaesaura, distribution, ecology, fire, management, habitat, Swaziland.
INTRODUCTION of appropriate fire management. The highly specialized African grass lizards (Chamaesaura) represent a group of animals ideally suited to a The African grass lizards of the genus Chamaesaura occur in the study of the impact of fire on their population numbers and Malolotja grassland, savanna and fynbos biomes of eastern and southern Africa Nature Reserve represents the ideal field laboratory for such a study. (Broadley & Howell 1991; Branch 1998; Spawls et al. 2004; Bates It is believed that in Swaziland, and in adjacent South Africa, two 2014). These serpentiform lizards are highly specialized and well main factors, namely forestry and fire, are affecting the conservation adapted to live, feed and breed in grassland habitats. Five species are status of these lizards. The severity of these threats is very apparent currently recognized (Bates 2014) of which three occur in Swaziland in Swaziland and has resulted in C. aenea and C. anguina being including C. aenea, C. anguina and C. macrolepis (Boycott 1992a, 1992b) regionally classified as Near-Threatened (Monadjem et al. 2003). within the following vegetation types Barberton and Kangwane The observations reported on in the present paper were Montane Grasslands, Sour Bushveld and Mixed Bushveld (Dobson & made primarily in Malolotja Nature Reserve, in western Swaziland. Lotter 2004). Additional information was obtained from elsewhere in the region. In southern Africa fire is a natural ecological process in highveld Malolotja Nature Reserve is located on the edge of the southern grassland and savanna, as it is in the savanna regions of eastern Africa. African escarpment, in south-eastern Africa between latitudes 25° Although the natural occurrence of fires continues, it is the frequency 58´ S and 26° 15´ S and between longitudes 31° 00´ E and 31° 10´ of fire that has been altered by man. While certain plant and animal E. Altitude within the reserve ranges from 640 m in the Nkomati species may be adapted to the natural fire regime, it is debatable valley to 1800 m a.s.l. on Ngwenya Mountain (Fig. 1). With decreasing whether these plants and animals are able to adapt to the man- altitude there is a noticeable increase in both the tallness of the altered fire regime in highveld grassland, without the intervention grasses and in the woodland element. The average annual rainfall, Durban Natural Science Museum Novitates 37 OBSERVATIONS ON AFRICAN GRASS LIZARDS 31
which falls between September and April inclusive, is, at the highest DISTRIBUTION AND HABITAT altitudes, approximately 1200 mm, although in wet years this may be as high as 1600 mm. The lower regions of the reserve receive Chamaesaura aenea approximately 890 mm per year. The summer climate may be There is a single historical sight record of C. aenea from the 1970’s described as warm, wet and humid, with considerable thunderstorm from a locality 5 km south of Forbes Reef (James Culverwell pers. activity. The winter climate as cold and dry, with occasional frosts comm.). No recent collections have been made at this locality and it appears as if C. aenea, based on current knowledge, is restricted to Malolotja Nature Reserve. Chamaesaura aenea is endemic to South Africa, Lesotho and Swaziland (Boycott 1992a). In Swaziland the species is restricted to the temperate highveld region in the west of the country. It occurs in short montane grassland, above 1500 m, which falls within the Barberton and Kangwane Montane Grassland vegetation type (Dobson & Lotter 2004). In Malolotja Nature Reserve, C. aenea and C. anguina occur sympatrically in an area of approximately 2 km2, at 1500 m to 1540 m (Table 1), between the main entrance gate and the Nkomati Viewpoint (VP) road turnoff. In the reserve C. aenea has not been recorded below 1500 m. The area of sympatry between the two species is likely to extend through higher altitudes in the reserve as there is suitable habitat above 1540 m, with the highest point, Ngwenya Mountain (1837 m), falling well within the altitudinal range of both species as recorded by Jacobsen (1989) and Bourquin (2003).
Fig. 2. Chamaesaura aenea, a species endemic to Swaziland and South Africa, and ‘Near-Threatened’ in Swaziland (photo. J. van Wyk).
Fig. 1. Map of Malolotja Nature Reserve in western Swaziland. The locations of the sample sites are shown. Key: solid lines – tourist roads, dotted lines – management tracks and hiking trails, o – viewpoints, NVR1 – Nkomati Viewpoint road sample site 1, NVR2 – Nkomati Viewpoint road sample site 2, LVR1 – Logwaja Viewpoint road sample site 1. and very infrequent snowfalls. While most of the reserve is subject to a temperate climate, the Nkomati valley represents a subtropical corridor extending westwards through the reserve into adjacent South Africa. As a consequence of the range in altitude and the associated differences in micro-climates and habitat types, the herpetofauna of Malolotja Nature Reserve is composed of a rich diversity in which both temperate and subtropical elements occur. The herpetofauna comprises 91 forms, including 25 amphibians, one chelonian, 29 lizards and 36 snakes (Boycott 1992a, 1992b, 1996, Boycott & Parker 2003). Malolotja’s reptiles represent 59 % of Swaziland’s recorded species. Fig. 3. Distribution of Chamaesaura aenea in Swaziland. Over a period of several years, all three species of Chamaesaura occurring in southern Africa, C. aenea, C. anguina and C. macrolepis, Chamaesaura anguina were found to occur in short grassland, tall grassland and savanna Chamaesaura anguina has been recorded from just four localities woodland habitats within the nature reserve. At some localities outside Malolotja Nature Reserve, three of which are quite old. sympatry between C. aenea and C. anguina, and between C. anguina and These include a KwaZulu-Natal Museum specimen from Mbabane C. macrolepis, was recorded. It was also discovered that the ranges of collected in 1939 and recorded by FitzSimons (1943), and a Ditsong C. aenea and C. macrolepis do not overlap. 32 OBSERVATIONS ON AFRICAN GRASS LIZARDS Durban Natural Science Museum Novitates 37
Table 1. Localities and altitude for Chamaesaura in Malolotja Nature Reserve.
Locality Co-ordinates Altitude (m)
C. aenea Senior Warden’s house ca. 26°08’36”S; 31°08’22”E. 1 510 Between Majolomba Picnic Site and Nkomati VP turnoff ca. 26°08’00”S; 31°07’35”E. 1 510 Tourist Office and Main Gate ca. 26°08’47”S; 31°08’16”E. 1 520 Majolomba Picnic Site ca. 26°08’00”S; 31°07’37”E. 1 510 Log Cabins ca. 26°08’23”S; 31°08’13”E. 1 520 Water tanks above Log Cabins ca. 26°08’22”S; 31°08’17”E. 1 540 Logwaja VP road near Nkomati VP turnoff ca. 26°07’57”S; 31°07’47”E. 1 500
range: 1 500 m to 1 540 m
C. anguina Senior Warden’s house ca. 26°08'36"S; 31°08'22"E. 1 510 Warden’s house ca. 26°08'21"S; 31°07'00"E. 1 435 Nkomati VP road (various sites) ca. 26°07'29"S; 31°07'32"E. 1 370 – 1 400 Between Campsite and workshop ca. 26°08'29"S; 31°07'45"E. 1 465 Logwaja VP road (various sites) ca. 26°08'00"S; 31°07'20"E. 1 460 - 1 520 Nkomati VP turnoff ca. 26°07'57"S; 31°07'37"E. 1 500 Majolomba River crossing ca. 26°07’20”S; 31°07’35”E. 1 380 Malanti area (various sites) ca. 26°06’52”S; 31°08’06”E. 1 220 - 1 260 Ngwenya Mountain ca. 26°11’28”S; 31°02’20”E. 1 620
range: 1 220 m to 1 620 m
C. macrolepis Malolotja jeep track (various sites) ca. 26°05'00"S; 31°07'11"E. 920 – 1 140 Nkomati VP road (various sites) ca. 26°07'29"S; 31°07'32"E. 1 240 – 1 400 Nkomati South trail ca. 26°03'57"S; 31°07'58"E. 1 000
range: 920 m to 1 400 m
National Museum of Natural History specimen from Manzini (listed between 1220 m and 1440 m (Table 1). In the reserve C. anguina has as Bremersdorp) collected in 1956. Subsequently the species has been not been found to occur below 1220 m. recorded from Black Mbuluzi Falls, 5 km north of Mbabane in 1977 (James Culverwell pers. comm.), while the most recent record of the species from outside Malolotja was collected by the present writer, in a patch of grassland on the Sibebe Rock massif, approximately 7 km north-east of Mbabane in January 2003. Chamaesaura anguina is endemic to South Africa and Swaziland (Boycott 1992a). It occurs in the temperate highveld region and in the transition zone between temperate grassland and subtropical savanna in western Swaziland. It occurs in tall, medium and short grassland habitats between 1220 m and 1620 m, which fall within Sour and Mixed Bushveld and Barberton and Kangwane Montane Grassland vegetation types (Dobson & Lotter 2004). In Malolotja Nature Reserve C. anguina and C. macrolepis occur sympatrically along the Fig. 4. Chamaesaura anguina, a species endemic to Swaziland and South Africa, Nkomati Viewpoint (VP) road over a distance of approximately 6 km, and ‘Near-Threatened’ in Swaziland (photo. J. Marais).
Table 2. Snout-vent length in Chamaesaura.
Species FitzSimons 1943 Jacobsen 1989 Branch 1998 Swaziland material C. aenea 134.0 mm 117.0 mm 137.0 mm 125.0 mm C. anguina 132.0 mm 140.0 mm 140.0 mm 139.0 mm C. macrolepis 166.0 mm 166.0 mm 178.0 mm Durban Natural Science Museum Novitates 37 OBSERVATIONS ON AFRICAN GRASS LIZARDS 33
Table 3. Snout-vent length as a % of total length in two species of Chamaesaura from Malolotja.
Chamaesaura anguina Chamaesaura macrolepis sex range % n= sex range % n= ♂♂ 20.8 – 23.4 7 ♂♂ 19.8 – 20.3 3 ♀♀ 23.4 – 30.1 3 ♀♀ 21.5 – 22.7 6 All 20.8 – 30.1 10 19.8 – 22.7 9
subtropical Swaziland and marginally in the temperate highveld region in the west of the country. It occurs in tall to medium grassland between 920 m and 1400 m but primarily in tall wooded grassland which falls within the Sour and Mixed Bushveld vegetation type (Dobson & Lotter 2004). In the reserve C. macrolepis has not been found to occur above 1400 m.
Fig. 7. Chamaesaura macrolepis occurs in Swaziland, South Africa and Zimbabwe (photo. R.C. Boycott). Fig. 5. Distribution of Chamaesaura anguina in Swaziland.
Fig. 6. Habitat of Chamaesaura anguina and C. aenea in Malolotja Nature Reserve (photo. R.C. Boycott).
Chamaesaura macrolepis Apart from the Malolotja Nature Reserve records reported on in the present paper, C. macrolepis has been recorded from five localities Fig. 8. Distribution of Chamaesaura macrolepis in Swaziland. in Swaziland, one of which is based on a sight record from Tshaneni (Barry Washington pers. comm.). A more recent historical record is a sighting of Chamaesaura, made in February 1993, from Nyamane School, approximately 5 km south of Nhlangano (Michael Staresinic pers. comm.). This could confirm FitzSimons’ (1943) earlier record of C. macrolepis from this locality, listed previously as Goedgegun. Nyamane School is located at 1000 m, which falls within the altitudinal range recorded for both C. macrolepis and C. anguina in Swaziland. However, as the specific identity of the lizard could not be established the record remains unconfirmed. The most recent record of the species from outside Malolotja Nature Reserve was secured by the present writer from near Mnyokane in December 2000. Chamaesaura macrolepis is endemic to South Africa and Swaziland (Boycott 1992a) with a relict population in the Chimanimani District of Zimbabwe (Broadley 1988). Sympatry between C. macrolepis and C. anguina in Malolotja Nature Reserve has been dealt with in Fig. 9. Habitat of Chamaesaura macrolepis in Malolotja Nature Reserve (photo. the previous account. Chamaesaura macrolepis occurs throughout R.C. Boycott). Note the firebreak placement on one side of the track. 34 OBSERVATIONS ON AFRICAN GRASS LIZARDS Durban Natural Science Museum Novitates 37
The recorded localities of Chamaesaura in Malolotja Nature when adpressed to the side of the body is closer to the vent than in Reserve provide an interesting pattern of habitat selection, in respect males. In males the limb joint is virtually in line with the vent and the of altitude and vegetation type, by the three species. It has been tip of the limb extends well beyond the vent. shown that C. anguina shares parts of the habitat with both C. aenea and C. macrolepis, while the areas of occurrence of C. aenea and C . FIRE AS AN ECOLOGICAL FACTOR macrolepis do not overlap (Table 1). Known voucher specimens (n=37) of the three species of As early as the 1980’s it was suggested that the paucity of these Chamaesaura from Swaziland are listed together with their localities lizards in the former Transvaal province of South Africa was directly and quarter-degree-grid cells in Appendix 1. as a result of the frequency of grassland burning (Jacobsen 1989). In eastern Africa it was reported that in burnt areas populations have SIZE AND SEXUAL DIMORPHISM been known to crash (Spawls et al. 2004). Large areas of Afromontane grassland in western Swaziland have Size been transformed into agricultural land and extensive areas are The genus contains highly specialised serpentiform lizards with very under alien forestry plantations, with some areas having been planted long and slender tails. The total length of large specimens can exceed to pine and gum in the early 1950’s. Grassland areas are invariably 500 mm (Appendix 2). Of the three species of Chamaesaura occurring burned every year to the detriment of Chamaesaura populations. Two in southern Africa, C. macrolepis attains the greater snout-vent length. herpetological surveys have been conducted by the writer in montane FitzSimons (1943) and Branch (1998) record a maximum snout-vent grassland areas in Mondi and Sappi plantations located in the Piggs length of 166.0 mm. The largest C. macrolepis from Swaziland is an Peak and Mbabane/Bhunya areas respectively. During these surveys no adult female (DM1685) with a snout-vent length of 178.0 mm (Table Chamaesaura specimens were found. The impact of annual burning of 2). The largest C. anguina recorded from Swaziland (Boycott 1990), an firebreaks in these regions cannot be discounted as one of the major adult female (TM67797), has a snout-vent length of 139.0 mm which factors responsible for the apparent disappearance of Chamaesaura falls within the maximum size of 140.0 mm given by Jacobsen (1989) populations from these areas. and Branch (1998). However, during the field study in Malolotja Fire is a natural ecological event in highveld grassland, as lightning Nature Reserve a captured and released adult female with a snout- strikes cause grass fires during the spring and summer. Although the vent length of 145.0 mm exceeded this (Appendix 2). The largest C . natural occurrence of fires continues, it is the frequency of fire that aenea from Swaziland is an adult female (DM1698) with a snout-vent has been altered by man. While certain plants and animals may be length of 125.0 mm which falls within the maximum size of 134.0 adapted to the natural fire regime, it is of concern whether or not mm and 137.0 mm given by FitzSimons (1943) and Branch (1998) they are able to adapt to the man-altered fire regime that is now respectively. practiced in highveld grassland. Most fire-burning programmes in highveld grassland commence as soon as the grass is dry enough to Sexual dimorphism burn, which invariably is during the winter months, in May or June. The sex of individual lizards of the two species encountered during Natural lightning-strike fires occur towards the end of winter, in the field study, namely C. anguina and C. macrolepis, was determined August and September, and can occur as late as February (pers. by the gravid condition of females and the length of the hindlimbs in obs). While the majority of hibernating reptiles might escape winter males and females. Road casualty specimens collected prior to the fires by remaining under cover beneath rocks, in rock crevices, investigation were dissected and lizards with larger size and larger underground or in termitaria, Chamaesaura hibernates in grass girth were found to be female, with some containing developing tussocks and consequently is more vulnerable to fires. embryos. These specimens are housed in the Durban Natural Science When fires burn into the wind, the actual belt of fire is seldom Museum and the Ditsong National Museum of Natural History wider than 50 cm, depending on the thickness of the grass, which collections. enables lizards to escape the fire and survive. Conversely, when the Colour-pattern differences were noted and invariably adult fire is burning with the wind, a far wider belt of fire is generated and females were more brightly marked than adult males and immatures. this could prevent escape and prove fatal to the lizards. Rocky ridges, In some cases femoral pores could be seen on the hindlimbs but this rock outcrops and wooded or forested river courses in highveld did not prove to be a useful field character. It is also impossible to grassland serve as natural firebreaks in their habitat. As fires burn into detect, unlike in many other lizard species, any swelling at the base or through rocky areas the fire can die due to a lack of combustible of the tail of males to indicate the presence of reproductive organs. material in these areas, and similarly, a wooded stream course or There appears to be a difference in the size and shape of the shields located posterior to the vent, with those in males being larger and Table 4. Malolotja fire-burning compartments. more flared than in females. However, this requires further study. Compartment Size Twenty-five percent of the lizards collected during the present study had lost their original tails and displayed re-generation or Office and Main Camp 2.5 X 1.0 = 2.5 km² truncations (Appendix 2). In a sample of 19 adult specimens believed Malolotja Vlei 2.5 X 1.5 = 3.8 km² to have their original tails intact, the snout-vent length represented Upper Majolomba catchment 1.0 X 1.5 = 1.5 km² 19.8 % to 30.1 % of their total length (Table 3). Although the Lower Majolomba catchment 3.0 X 1.5 = 4.5 km² sample is small there is some evidence of sexual dimorphism in this Mbulukudvu catchment 3.0 X 4.0 = 12.0 km² characteristic, with males showing a slightly shorter snout-vent length Nkomati valley 3.0 X 3.0 = 9.0 km² in relation to total length than females. Tjomoloti catchment 1.0 X 2.0 = 2.0 km² Of the features mentioned, the length of the hindlimbs proved far Ngwenya 1.5 X 6.0 = 9.0 km² more reliable in determining the sex of individuals. This is best noted She Mine 4.5 X 3.5 = 15.7 km² by the position of the vent in relation to the adpressed hindlimb. In both males and females the tip of the hindlimb extends beyond the Maphandakazi catchment 4.5 X 3.0 = 13.5 km² vent. With known gravid female specimens, the tip of the hindlimb Durban Natural Science Museum Novitates 37 OBSERVATIONS ON AFRICAN GRASS LIZARDS 35
Table 5. Chamaesaura numbers observed at sample sites in July 1997.
Sample Last Age of Number of Species Sex ratio site burned grassland Lizards breakdown (♂:♀:imm) (month/yr) (months) (n=)
NVR1 Oct. 1996 9 6 2 C. anguina 1:1:0 4 C. macrolepis 1:2:1 NVR2 Jun. 1995 25 31 10 C. anguina 5:3:2 21 C. macrolepis 5:5:11 LVR1 Aug. 1995 23 10 10 C. anguina 3:4:3
drainage line can also halt a fire. Other factors that stop fires include unharmed beneath rocks in burnt firebreaks in June and July. Although a change in the wind direction and precipitation in the form of rain Jacobsen (1989) recorded one Chamaesaura macrolepis in a hollow or mist, and at night, dew. Precipitation in any form is able to render under a rock on soil, the grass dwelling Chamaesaura species do not the vegetation too wet to burn. These factors may, to a certain degree, normally shelter under objects in their habitat and would have to serve to naturally limit the impact of fire on Chamaesaura populations contend with fires at this time. in highveld grassland, provided the incidence of annual fires is In Malolotja Nature Reserve no section of grassland is burned controlled. purposely every year. However, with some of the protective firebreaks Fire management programmes differ depending on the land-use along fence lines, around buildings and tourist accommodation practices in highveld grassland. Within protected areas the main facilities this is not always possible. In these instances the firebreaks objective is to try and maintain plant species diversity by protecting are not placed in the exact same position as the previous year but are certain sections of the habitat from fire each year, while deliberately shifted slightly or the width changed. Firebreaks along roads or tracks burning selected sections on a rotational basis of two, three or four are alternated over two to four years (see Fig.9). Ecologically sensitive years. Outside protected areas, annual burning of the grassland is areas, such as those containing wetlands, are left unburned for four or practiced to provide winter grazing for cattle and sheep. In forestry five years. Lightning strike fires are allowed to burn, unless they occur areas firebreaks are burned annually to protect the plantations from in a compartment burned the previous year, in which case they are fire. In the cultivation of maize, at least in Swaziland, cattle are allowed extinguished. The objective of the fire-burning programme is to try into maize fields after the harvest to feed on the left-over plant and achieve a patchwork of different-aged grassland compartments material and to spread their manure. What is left is often burned to each year. In respect of Chamaesaura and undoubtedly other reptiles, return nutrients to the soil, and these fires are often left unchecked this allows for recruitment to take place from older to younger and escape into the adjacent grassland. grassland. Concern was expressed previously as to the susceptibility of The river, road and track network in the southern part of Chamaesaura to grass fires when mortalities were recorded during Malolotja allows this part of the reserve to be divided into ten fire-burning programmes in Malolotja Nature Reserve (Boycott compartments for controlled block burning. The size of the 1990, 1992b). Perhaps this concern was somewhat premature, as compartments range from approximately 1.5 km² to 15.7 km², with the observations presented in the present paper might indicate that five compartments being less than 5 km² and five being more than 5 Chamaesaura is actually a fire-adapted genus of lizards. km² (Table 4). This allows for a controlled fire-burning programme that reduces the chances of large extensive areas being burned. FIRE MANAGEMENT IN MALOLOTJA NATURE RESERVE EFFECT OF FIRE FREQUENCY ON CHAMAESAURA Both natural and man-made fires occur in Malolotja Nature Reserve POPULATION NUMBERS IN MALOLOTJA NATURE and these vary in extent, intensity and time. Natural fires are those RESERVE started by lightning strikes, while man-made fires consist of controlled firebreak burning and controlled block or compartment burning. Some Introduction lightning-strike fires occur outside the boundaries of the reserve and The annual fire-burning programme at Malolotja Nature Reserve burn into the reserve. Similarly, some fires spread into the reserve as provides the opportunity to assess what effect fire has on a result of the traditional burning of the grassland by Swazis in areas Chamaesaura populations in different-aged grassland habitats. bordering the reserve. Fires are also started deliberately by poachers Consequently, in July 1997, an informal investigation was carried inside the reserve. During controlled firebreak burning, there is an out with the assistance of reserve management staff involved in the element of risk, as a sudden change in wind direction could lead to a firebreak burning. The aim of the investigation was to determine controlled burn getting out of control. whether there were larger numbers of lizards in older grassland. Between 1988 and 1998 there were eight lightning-strike fires recorded in the southern portion of the reserve during the months Method of September (1), October (6) and January (1). Controlled firebreak Three 1 km sections of grassland adjacent to gravel roads, in burning is carried out in June, July and August, while the controlled different-aged grassland, were selected for firebreak burning. The block or compartment burning is carried out in September, October firebreaks were approximately 40 m wide. Two of the sections were or in early November, depending on the amount of rainfall recorded. located along the Nkomati Viewpoint road and one along the Logwaja The controlled block burning is therefore closely aligned to the Viewpoint road (see Fig. 1). The three sections are located at much natural fire regime of the Swaziland highveld. During June, July and the same altitude, between 1 260 m and 1 520 m. The sections were August any hibernating reptiles will be unaffected by the firebreak all burned at roughly the same time, between 12h00 and 15h00, and burning. Boycott (1990) listed several species of reptiles observed during similar weather conditions that were sunny, clear and warm 36 OBSERVATIONS ON AFRICAN GRASS LIZARDS Durban Natural Science Museum Novitates 37
with little or no wind, over a period of approximately three weeks. more than half a century, will ultimately lead to the local extinction During the burning of each firebreak all Chamaesaura that were of Chamaesaura populations. This appears to be the case in Swaziland flushed into the road were collected. It is understood that some with C. aenea and C. anguina having being classified as Near- lizards might have fled into the adjacent unburned grassland next Threatened (Monadjem et al. 2003). to the firebreak and some may have remained in the burnt area. No attempt was made to capture these and only those that were CHAMAESAURA AS A FIRE-ADAPTED GENUS flushed into the road were collected. Later they were identified, OF LIZARDS measured, sexed and their reproductive state (in females) noted. After processing these data (see Appendix 2), all the specimens were The issue of fire and its effect on Chamaesaura populations has released into the unburned areas next to the respective firebreaks. been documented on a number of occasions (Bruton & Haacke 1980; Jacobsen 1989; Boycott 1990, 1992b; Monadjem et al. 2003; History of burning of the three sample sections Spawls et al. 2004; de Villiers & de Villiers 2004; Alexander & Marais The first Nkomati Viewpoint road sample section (NVR1) was last 2007). Jacobsen (1989) believes that fire is probably responsible for burned in October 1996, approximately nine months previously, the paucity of these lizards in most of the areas where they occur. when both sides of the road were burned as part of a block or Observations made in Swaziland clearly support this. It is not easy compartment burn. The second Nkomati Viewpoint road sample to find these lizards while searching for them in their grassland section (NVR2) was last burned in June 1995, approximately 25 and savanna habitats. However, large numbers may be flushed from months before, as part of an internal protective firebreak. The cover during controlled fire-burning operations and during wild fires opposite side of the road had been burned the previous year in (Bruton & Haacke 1980; Boycott 1990). Although mortalities have August 1996, 11 months before, as part of an internal protective been recorded after fires (Jacobsen 1989; Boycott 1990, 1992b; de firebreak. The Logwaja Viewpoint road sample section (LVR1) was last Villiers & de Villiers 2004) it has also been observed that many lizards burned in August 1995, approximately 23 months previously, as part survive fires in highveld grassland (pers. obs and this study). of an internal protective firebreak. The opposite side of the road had Having evolved in the grassland and savanna regions of Africa, it been burned 12 months before in July 1996, as part of an internal would seem that Chamaesaura has had to adapt to fire throughout protective firebreak. This history provided the opportunity to observe its evolutionary history. It is suggested that in the grassland biome of the numbers of lizards in three different grassland sections of various south-eastern Africa and probably in the fynbos biome of southern ages, ranging from nine months to 25 months. Africa, these lizards are fire-adapted species. Female lizards of the three species are capable of producing 5 - 17 live young (Bates Results of investigation 2014). This is important in respect of the recruitment potential of The three sampling sites yielded a total of 47 Chamaesaura, 22 C . these species and could be interpreted as a strategy to counter anguina and 25 C. macrolepis (Table 5). At the first Nkomati Viewpoint the effects of fire. The females of C. anguina and C. macrolepis in the road sample site (NVR1), six specimens of Chamaesaura were flushed highveld grassland of western Swaziland and adjacent South Africa are into the road during the burning of the firebreak and were collected. gravid during the winter and spring, and give birth in November or Four of these were C. macrolepis and the other two were C. anguina. December (FitzSimons 1943; Jacobsen 1989; Boycott 1990). At the second Nkomati Viewpoint road sample site (NVR2), thirty- During the present study five of the eight adult female C . one specimens of Chamaesaura were flushed into the road during anguina, and all seven adult female C. macrolepis were gravid the burning of the firebreak. Twenty-one of these were C. macrolepis (Appendix 2). It seems that parturition in these two species occurs and ten were C. anguina. At the Logwaja Viewpoint road sample site once the incidence of natural lightning-strike fires has passed, (LVR1), ten specimens of Chamaesaura were flushed into the road thereby enhancing the survival of neonates and achieving successful during the burning of the firebreak, all of which were C. anguina. recruitment in grassland habitats. This could also be interpreted as a The sex ratios recorded in the three samples show a fairly even fire-adaptive strategy. It has been established that some gravid females distribution between males and females of both species and the may perish during grass fires but some will survive, and the survivors numbers of immatures indicate that recruitment at the three sites are capable of giving birth to up to 17 young after the natural fire is taking place (Table 5). The numbers of lizards recorded from the season. These two aspects of their biology could be important fire- three sampling sites indicate that more lizards were recorded in older adaptive strategies for Chamaesaura and may be applicable elsewhere grassland habitats. In the two older grassland sites that had not been in southern Africa. In respect of the third species, C. aenea, two burned for 25 and 23 months, 31 and 10 lizards respectively were Malolotja females collected in February and May were gravid (pers. counted over a 1 km section, whereas in the nine month old grassland obs). Branch & Haagner (1992) report on a female from nearby only six lizards were counted over a 1 km section. Lothair, collected in November, containing a single full-term embryo. They also report on another female, collected in April from montane Conclusion grassland in the north-eastern Cape, containing eight yolked ova with Although a far more detailed investigation would have been desirable, little or no embryonic development. This evidence seems to suggest it can be concluded from the above investigation, that the oldest that C. aenea females are also gravid at that time of the year when section of grassland had the greatest number of lizards. This is an natural lightning-strike fires are likely to occur and are able to give important observation and should be considered when deciding birth after the threat of fires has passed. on an appropriate fire-management programme for grassland If Branch’s (1988) statement of parturition for C. anguina in habitats that support Chamaesaura populations. Annual burning of April applies to Cape populations in the fynbos biome, then it could grassland as practiced in the surrounding areas and in forestry areas be considered that the same strategy is at work. After recording a is undoubtedly detrimental to Chamaesaura populations because gravid female C. anguina from Malolotja Nature Reserve in August, it is likely to inhibit recruitment into and re-colonisation of burnt Boycott (1990) suggested that this species might either give birth habitats. Annual burning of grassland habitats outside protected areas, to multiple broods or have different geographical breeding seasons. especially in forestry areas, some of which have been in existence for On the available evidence it appears as if the latter is the case. Such Durban Natural Science Museum Novitates 37 OBSERVATIONS ON AFRICAN GRASS LIZARDS 37
a strategy could have evolved as a result of the natural incidence of A.M., BURGER, M., MARAIS, J., ALEXANDER, G.J. & DE VILLIERS, lightning-strike fires in grassland and fynbos. Although the incidence of M.S. (eds.), Atlas and red list of the reptiles of South Africa, Lesotho natural lightning-strike fires in the fynbos is far less frequent than in and Swaziland. Suricata 1, pp.183-185. South African National the grasslands of south-eastern Africa, they occur at a different time Biodiversity Institute: Pretoria. of the year, usually during the hot dry summer months. De Villiers & BOURQUIN, O. 2003. Reptiles (Reptilia) in KwaZulu-Natal: I – de Villiers (2004) report on a devastating fynbos fire that occurred diversity and distribution. Durban Museum Novitates 29: 57-103. in January in the Gansbaai region of the south-western Cape. One BOYCOTT, R.C. 1990. Chamaesaura anguina – Size, reproduction and of the casualties in the fire was a gravid female. April is believed susceptibility to fire. Journal of the Herpetological Association of to be after the natural fire season in that biome and parturition at Africa 37: 49. this time would enhance the survival of neonates and recruitment BOYCOTT, R.C. 1992a. An annotated checklist of the amphibians in fynbos habitats. It would be difficult to explain in any other way, and reptiles of Swaziland. The Conservation Trust of Swaziland: or to offer an alternative argument for, any selective advantage of Mbabane. different parturition times in two different biomes for C. anguina. BOYCOTT, R.C. 1992b. A herpetofaunal survey of Swaziland. Furthermore, it seems likely that a group of highly specialized grass Unpublished MSc. Thesis. University of Natal: Durban. lizards, which evolved in the grasslands and savannas of Africa, should BOYCOTT, R.C. 1996. Lamprophis fuscus – size, lepidosis and show adaptations to fires, considering that fire is an integral part of distribution. African Herp News. 25: 40-41. the ecology of these regions. BOYCOTT, R. & PARKER, V. 2003. Birds of the Malolotja Nature Reserve, Hopefully as more is learned of the biology and ecology of Chamaesaura, the observations reported on in the present paper Swaziland. Bright Continent Guide 3. Avian Demography Unit and may be substantiated further. It is very important for the continued the Conservation Trust of Swaziland: Cape Town and Mbabane. survival of these highly specialized lizards that appropriate fire BRANCH, B. 1988. Field guide to snakes and other reptiles of southern management of highveld grassland, such as that practiced in Malolotja Africa. First Edition. Struik Publishers: Cape Town. Nature Reserve, be carried out in the areas where they occur. The BRANCH, B. 1998. Field Guide to snakes and other reptiles of southern primary objective would be to ensure that large extensive areas are Africa. Third Edition. Struik Publishers: Cape Town. not burned and that smaller compartments be managed appropriately BRANCH, W.R. & HAAGNER, G.V. 1992. Chamaesaura aenea – with fire. The presence of healthy Chamaesaura populations is likely to Reproduction. Journal of the Herpetological Association of Africa 41: be indicative of sound and appropriate fire management in highveld 42-43. grassland. BROADLEY, D.G. 1988. A check list of the reptiles of Zimbabwe, with synoptic keys. Arnoldia Zimbabwe 9: 369-430. ACKNOWLEDGEMENTS BROADLEY, D.G. & HOWELL, K.M. 1991. A check list of the reptiles of Tanzania, with synoptic keys. Syntarsus 1: 1-70. This paper is dedicated to the Malolotja Nature Reserve contract BRUTON, M.N. & HAACKE, W.D. 1980. The reptiles of Maputaland. firefighters and rangers in recognition of a difficult and dangerous In: BRUTON, M.N. & Cooper, K.H. (eds), Studies on the Ecology of job well done. In particular the following are thanked for collecting Maputaland, pp. 251-287. Rhodes University & Natal Branch of the the lizards: J. Mabuza, M. Makhanya, M. Masimula, L. Mkhabela, D. Wildlife Society of Southern Africa: Grahamstown & Durban. Msimango and P. Ngwenya. Ara Monadjem and Kim Roques provided DE VILLIERS, A.L. & DE VILLIERS, M.E. 2004. Chamaesaura anguina – assistance during other field surveys. The Animal Demography Unit Fire, population size and density. African Herp News 37: 22-23. of the University of Cape Town and the Conservation Trust of DOBSON, L. & LOTTER, M. 2004. Vegetation map of Swaziland. In: Swaziland are thanked for permission to reproduce the map from an MUCINA, L. & RUTHERFORD, M.C. (eds.), Vegetation Map of South earlier publication on the birds of Malolotja Nature Reserve (Bright Africa, Lesotho and Swaziland. National Botanical Institute: Cape Continent Guide 3). Johan Marais and Jaco van Wyk are thanked for Town. providing photographs. Timothy Boycott is thanked for scanning the FITZSIMONS, V.F. 1943. The Lizards of South Africa. Memoirs of the photographs and digitizing the figures. For fruitful discussion and encouragement the writer would like to thank Orty Bourquin, Michael Transvaal Museum 1: 1-528. Cunningham, Niels Jacobsen and Angelo Lambiris. JACOBSEN, N.H.G. 1989. A herpetological survey of the Transvaal. Unpublished PhD. Thesis. University of Natal: Durban. REFERENCES MONADJEM, A., BOYCOTT, R.C., PARKER, V. & CULVERWELL, J. 2003. Threatened Vertebrates of Swaziland - Swaziland Red Data Book: ALEXANDER, G. & MARAIS, J. 2007. A guide to the reptiles of southern Fishes, Amphibians, Reptiles, Birds and Mammals. Ministry of Tourism, Africa. Struik Publishers: Cape Town. Environment and Communications: Mbabane, Swaziland. BATES, M.F. 2014. Chamaesaura aenea, Chamaesaura anguina anguina, SPAWLS, S., HOWELL, K., DREWES, R. & ASHE, J. 2004. A Field Guide to Chamaesaura macrolepis, In: BATES, M.F., BRANCH, W.R., BAUER, the Reptiles of East Africa. Reprinted Edition. A & C Black: London. 38 OBSERVATIONS ON AFRICAN GRASS LIZARDS Durban Natural Science Museum Novitates 37
APPENDIX 1.
Voucher specimens of Chamaesaura from Swaziland.
Chamaesaura aenea (n = 7) Log Cabins, MNR (DM1699) (2631AA); Main Gate, MNR (DM1682) (2631AA); Majolomba picnic site, MNR (DM1690, 1695, 1698 TM71104) (2631AA); btwn. Majolomba picnic site and Nkomati Viewpoint turnoff, MNR (TM70967) (2631AA).
Chamaesaura anguina (n = 17) Logwaja Viewpoint road, MNR (DM1691) (2631AA); Majolomba River crossing, MNR (DM1687) (2631AA); Malanti area, MNR (DM1697, RCBS1598) (2631AA); Manzini (as Bremersdorp) (TM24360) (2631AD); Mbabane (KNM782, Fitz. 1943) (2631AC); Main Campsite, MNR (TM64732, 64745, 64746) (2631AA); btwn. Main Campsite and workshop, MNR (TM71817) (2631AA); Ngwenya Mountain, MNR (DM1696) (2631AA); Nkomati Viewpoint road, MNR (DM1693, TM67797) (2631AA); Nkomati Viewpoint turnoff, MNR (DM1684) (2631AA); Sibebe Rock, 7km NE of Mbabane (DM1672) (2631AC); Warden’s house, MNR (DM1683, TM67939) (2631AA).
Chamaesaura macrolepis (n = 13) Mafutseni, 1 mile E of (TM47459) (2631BC); Malolotja jeep track, MNR (DM1685, 1688, 1689) (2631AA); Mlilwane Wildlife Sanctuary (TM53278, 54100, 54101, 54102) (2631AC); Mnyokane, 5km E of (RCBS1947) (2631AA); Nhlangano (as Goedgegun) (TM19417, Fitz. 1943) (2731AA); Nkomati South trail, MNR (DM1694) (2631AA); Nkomati Viewpoint road, MNR (DM1686, 1692) (2631AA).
Abbreviations: Ditsong National Museum of Natural History (formerly Transvaal Museum) (TM); Durban Natural Science Museum (DM); FitzSimons 1943 (Fitz.); Malolotja Nature Reserve (MNR); KwaZulu-Natal Museum (KNM); R.C. Boycott Swaziland Collection (RCBS). Durban Natural Science Museum Novitates 37 OBSERVATIONS ON AFRICAN GRASS LIZARDS 39
APPENDIX 2.
Details of Chamaesaura specimens examined from three sample sites in Malolotja Nature Reserve (* - denotes loss of original tail with truncations or re-growth)
Sample site Species Total length Sex NVR1 C. anguina 94 + 308 = 402 mm adult male " not recorded adult female (gravid) C. macrolepis 149 + 543 = 692 mm adult female (gravid) " 110 + 270* = 380 mm adult male " 147 + 500 = 647 mm adult female (gravid) " 82 + 291 = 373 mm immature
NVR2 C. anguina 74 + 235 = 309 mm immature " 98 + 122* = 220 mm adult male " 96 + 315 = 411 mm adult male " 84 + 286 = 370 mm adult male " 86 + 277 = 363 mm adult female " 142 + 97* = 239 mm adult female (gravid) " 145 + 337 = 482 mm adult female (gravid) " 84 + 288 = 372 mm adult male " 77 + 259 = 336 mm immature " 97 + 370 = 467 mm adult male
C. macrolepis 88 + 311 = 399 mm immature " 121 + 476 = 597 mm adult male " 121 + 474 = 595 mm adult male " 80 + 316 = 396 mm immature " 92 + 338 = 430 mm immature " 87 + 335 = 422 mm immature " 95 + 355 = 450 mm immature " 136 + 315* = 451 mm adult female (gravid) " 141 + 497 = 638 mm adult female (gravid) " 83 + 309 = 392 mm immature " 80 + 307 = 387 mm immature " 81 + 300 = 381 mm immature " 78 + 263 = 341 mm immature " 75 + 209* = 284 mm immature " 94 + 328 = 422 mm immature " 115 + 261* = 376 mm adult male " 108 + 437 = 545 mm adult male " 130 + 443 = 573 mm adult female (gravid) " 105 + 238* = 343 mm adult male " 124 + 434 = 558 mm adult female (gravid) " 136 + 493 = 629 mm adult female (gravid)
LVR1 C. anguina 73 + 263 = 336 mm immature " 80 + 285 = 365 mm adult male " 93 + 215* = 308 mm adult male " 93 + 331 = 424 mm adult male " 63 + 105* = 168 mm immature " 72 + 258 = 330 mm immature " 86 + 282 = 368 mm adult female " 94 + 105* = 199 mm adult female " 132 + 184* = 316 mm adult female (gravid) " 123 + 201* = 324 mm adult female (gravid) 40 MOULT OF THE GREY-HEADED GULL Durban Natural Science Museum Novitates 37
MOULT OF THE GREY-HEADED GULL LARUS CIRROCEPHALUS IN SOUTH AFRICA
A.M. MCINNES1, D.G. ALLAN2 & L.G. UNDERHILL3 1DST-NRF Centre of Excellence at the Percy FitzPatrick Institute of African Ornithology, University of Cape Town, Rondebosch 7701, South Africa email: [email protected]. 2Durban Natural Science Museum, P.O. Box 4085, Durban 4000, South Africa 3Animal Demography Unit, Department of Biological Sciences, University of Cape Town, Rondebosch 7701, South Africa
cInnes, A.M., Allan, D.G. & Underhill, L.G. 2014. Moult of the Grey-headed Gull Larus cirrocephalus in South Africa. Durban MNatural Science Museum Novitates 37: 40-46. A total of 284 adults and 49 immature Grey-headed Gulls, trapped in Gauteng and KwaZulu-Natal, South Africa, and 14 museum specimens, were inspected for moult. Twenty-eight adults and 17 immatures were in active primary, secondary or tail moult. Primary moult duration was estimated using the percentage feather mass grown (PFMG) technique. Mean primary moult duration for adults was calculated as 136 days between 12 October (mean starting date) and 24 January (mean completion date), primarily during the non-breeding season, although six gulls were in active primary moult in late August. The timing of primary moult in immatures was more variable than adults. In both adults and immatures the timing of moult in the outermost secondaries (S1 – S10) coincided approximately with the moulting of the central primaries. This was followed by a second wave of moult in the innermost secondaries. Tail moult was mostly associated with the early stages of primary moult for both age groups. Observational data from monthly waterbird counts in Durban Bay showed head moult in adults occurred in November and December (into non- breeding head) and again between March and May (into breeding hood). Two first-year birds had active wing and tail moult; this is the second record of this phenomenon for Grey-headed Gulls. This is unusual for ‘masked’ gulls, Hartlaub’s Gull, the closest relative of Grey-headed Gull, being the only other known species that undergoes this premature moult. This may have been influenced by similar selection pressures for increased post-natal dispersion for these African sister species.
KEYWORDS: Grey-headed Gull, Larus cirrocephalus, moult.
INTRODUCTION migration having been recorded for birds between St Lucia and Durban (McInnes & Allan 2011). The Grey-headed Gull Larus cirrocephalus is one of ten species in The literature on moult in masked gulls is extensive, see reviews the ‘masked’ gull species group. These are relatively small gulls that by Dwight (1925) for all gull species, Cramp & Simmons (1983) invariably possess a contrasting hood during the breeding season and Olsen & Larsson (2003) for Palaearctic and North American (Crochet et al. 2000; Given et al. 2005). The two largest breeding species (including Grey-headed Gull), and by Higgins & Davies (1996) populations in South Africa are in Gauteng (estimated breeding for Australian and New Zealand species. Information on moult in population for 2005: 2185 breeding pairs) and at Lake St Lucia where Hartlaub’s Gull Larus hartlaubii has been synthesised by Crawford & they frequently breed on Bird and Lane islands (estimated breeding Underhill (2003). Grey-headed Gulls typically undergo descendent population for 2000: 678 breeding pairs), with smaller colonies primary moult and, in KwaZulu-Natal, this is initiated between July dispersed throughout all the other provinces except Limpopo and October, and is completed between November and March, while (Brooke et al. 1999; McInnes & Allan 2011). Breeding phenology in Zambia primary moult has been noted in all months sampled, varies between sites but is predominantly during the austral winter including during the breeding season (Crawford & Hockey 2005). To (reviewed by Crawford & Hockey 2005). Ring recoveries of chicks date, most moult studies have been concerned with the primaries and reared in South Africa have shown widespread juvenile dispersal there is little information published on secondary moult, especially as far as Zambia (Underhill et al. 1999) and adult birds are known for the northern hemisphere species. to have dispersive/nomadic movements during the non-breeding The following account describes the estimated phenology of season (reviewed by Crawford & Hockey 2005), with regular partial primary and head moult, as well as secondary and tail moult Durban Natural Science Museum Novitates 37 MOULT OF THE GREY-HEADED GULL 41
processes, for the Grey-headed Gull from populations sampled in months and three years. For the purposes of individual accounts of Gauteng during the breeding season and from non-breeding birds immature moult we further classified these birds into two year classes trapped or collected in KwaZulu-Natal. based on Olsen & Larsson (2003). Each gull trapped was examined for active moult in the primaries and tail. If active moult in the primaries was evident, then secondary moult was also examined. A standard moult scoring system (Ginn & Melville 1983) was used to record the stage of moult for each primary, secondary and tail feather. Scores: (0) old feather, (1) missing or new feather completely in pin, (2) new feather just emerging from sheath, (3) new feather 1/3 – 2/3 grown, (4) new feather 2/3 to fully grown but with some sheath remaining, (5) new feather but no trace of sheath, (8) fully grown feather, age uncertain. Information on the timing of head moult in adults was taken from monthly waterbird counts conducted in Durban Bay during 2004 and 2005 (Allan & McInnes unpublished data). During these counts, adults were partitioned into two categories (breeding/non-breeding) by assessing the state of breeding hood development with the aid of Fig. 1. Total monthly numbers of adult Grey-headed Gulls sampled in Gauteng binoculars from distances of approximately 10 m - 200 m. and Durban between April 2004 and July 2005 (black bars) and total number of adult Grey-headed Gulls recorded in active primary moult (white bars). Primary moult duration Estimates of the duration of primary moult and start and finish dates METHODS were obtained using the model of Underhill & Zucchini (1988). This method uses the percentage feather mass grown (PFMG) as the Capture techniques and procedure governing protocol for estimating the stage of progression through All birds were trapped using a spear-gun-driven net (McInnes et primary moult. We used this method’s “data type 2” moult index al. 2005). This trap was used on landfill sites in Gauteng and on which includes all birds sampled, i.e. not just birds actively moulting. Blue Lagoon Beach in Durban. Grey-headed Gulls were lured within In order to calculate PFMG, all primary feathers of four Grey- range of the net by baiting them with restaurant discards in Gauteng headed Gull wings, coming from four specimens collected as freshly and with fishing bait (pilchards) in Durban. All birds caught were dead in Gauteng, were weighed to the nearest 0.01 g using an Ohaus temporarily stored in ventilated cardboard boxes until processed. Each electronic balance. Additional moult data from 14 adult specimens bird was weighed, ringed with both an individually engraved colour from Durban Natural Science Museum collected from KwaZulu- ring and a metal ring. Each bird was then measured and the moult Natal between 1965 and 1977 were used in this analysis (Accession was assessed. Age classification was based on plumage and bare-part numbers/dates (yyyymmdd): characteristics as described by Grant (1978) and Olsen & Larsson (2003) and the month in which they were trapped: breeding adult - pale eye, grey hood, white mirrors on outer primaries, observed DNSM832/19611117 DNSM30712/19751023 between May and August; non-breeding adult - as with breeding adult DNSM30153/19730125 DNSM29694/19731216 but hood usually pale and observed between November and March; DNSM18396/19650304 DNSM30759/19751229 juvenile – dark eye, extensive brown and dusky markings in plumage DNSM1045/19611227 DNSM1016/19611127 (notably wing feathers and coverts), black terminal bar to the tail. DNSM30982/19770102 DNSM834/19611117 Using this classification, we combined all birds that were not juveniles DNSM408/19611012 DNSM1017/19611127 or adults into a generic ‘immature’ class due to a limited sample of these birds – these include birds aged between approximately six DNSM15028/19630405 DNSM835/19611117
Table 1. Sample effort details of trapped Grey-headed Gulls inspected for moult between 2004 and 2005. The number and proportion of birds that showed evidence of primary, secondary or tail moult is shown.
Adults Immatures Site Coordinates Province No. No. % No. No. % sampled moulting sampled moulting
Linbro Park 2605S 2807E Gauteng 1 0 0 0 - - Landfill Rooikraal 2618S 2815E Gauteng 80 7 8.8 20 7 35 Landfill Simmer & 2612S 2808E Gauteng 48 0 0 8 2 25 Jack Landfill Weltevreden 2612S 2821E Gauteng 139 6 4.3 14 1 7.1 Landfill Blue Lagoon 2948S 3202E KwaZulu-Natal 16 15 93.8 7 7 100 42 MOULT OF THE GREY-HEADED GULL Durban Natural Science Museum Novitates 37
Table 2. Mass (in grams) and proportionate mass (%) of primary feathers taken During January and February, four of the six gulls sampled were from one wing each of four Grey-headed Gulls collected in Gauteng during June moulting their proximal secondaries (S11-S14) inwardly (ascending), 2004. during the final stages of primary moult. Active moult at S17 was evident in all months between November and February, except Primary %Total December. In all months sampled, between 20% and 86% of all no. Mass1 Mass2 Mass3 Mass4 Mean mass moulting birds had some degree of moult in the innermost three 10 0.39 0.41 0.47 0.48 0.44 16.83 secondaries (S18 – S20). 9 0.36 0.38 0.45 0.45 0.41 15.77 Of all Grey-headed Gulls recorded in active primary moult, birds 8 0.32 0.34 0.4 0.43 0.37 14.33 were only recorded in tail moult during November (six of seven) and 7 0.27 0.29 0.33 0.37 0.32 12.12 December (one of two). The pattern of moult here was generally 6 0.24 0.26 0.29 0.32 0.28 10.67 synchronous starting with the innermost retrices and ending with the 5 0.21 0.22 0.25 0.26 0.24 9.04 4 0.16 0.17 0.2 0.2 0.18 7.02 3 0.13 0.14 0.16 0.17 0.15 5.77 2 0.11 0.12 0.13 0.13 0.12 4.71 1 0.08 0.1 0.11 0.11 0.10 3.85 Total 2.27 2.43 2.79 2.92 2.60 100.00
RESULTS
Sample effort A total of 284 adult Grey-headed Gulls and 49 immature Grey- headed Gulls, trapped between April 2004 and July 2005, were inspected for moult (Table 1.). The majority (n= 310, 93%) of birds Fig. 2. Proportion (%) of adult Grey-headed Gulls with breeding hoods were trapped in Gauteng during the breeding season (May - August between January and December 2004, 2005. Data from aged counts 2004, June/July 2005) while the remainder (n=23, 7%) were caught conducted monthly in Durban Bay. on Durban’s northern beachfront during the non-breeding season (November 2004 - February 2005). Birds caught in Durban had a outermost retrices except for one gull (SAFRING ID: 37776) that had higher proportion (adults – 93.8 %, immature - 100%) of birds in all of its retrices moulting simultaneously. The limited sample suggests active moult when compared to those caught in Gauteng (adults - 4.9 that the timing of tail moult fell within the first half of the primary %, immature - 23.8%) (Table 1). moult period, i.e. during the austral spring and early summer. Moult scores for all birds (adults and immatures) recorded with The timing of head moult for adult Grey-headed Gulls counted active moult are shown in Appendix 1. in Durban Bay is illustrated in Fig. 2. Head moult of adults into non-breeding hoods occurred during the months November and Adults December. Grey-headed Gulls acquired their breeding hoods between Twenty-eight of 284 adults (9.8%) captured were recorded in active March and May, with the majority (80%) of birds having moulted into moult. Twenty-one adults (7.4%) were in active primary moult (Fig. 1). their breeding hoods by April. Of the birds sampled during the breeding season, only six were found moulting the inner three primaries in late August. The majority (94%) Primary moult duration using PFMG of Grey-headed Gulls trapped between November 2004 and February The average weight of each primary feather and the proportionate 2005 during the non-breeding season were recorded with active mass of each feather to the total mass of all primaries are given in primary moult. Six adult Grey-headed Gulls, trapped between May and Table 2. The mean mass of all 10 primaries was 2.6 g. The outermost August 2004, which had not initiated primary moult yet, had active primary (P10) was on average 4.4 times heavier than the innermost moult in their inner tail feathers. primary (P1). The estimated mean starting date for Grey-headed Gull Two birds (SAFRING IDs: 37806, 37811) trapped in December primary moult was 12 October and the estimated mean completion and January that were in the final stages of primary moult, i.e. date was 24 January with a mean estimated duration of 136 days moulting P8 – P10, were simultaneously moulting one inner (Table 3). The standard deviation of the mean starting date was 25 primary each (P2 and P3, respectively). This could be as a result of days with a standard error of four days. It is therefore estimated that a simultaneous, second wave of moult or, alternatively, may indicate 95% of Grey-headed Gulls, in the sample population, initiated primary an initial partial moult starting at P4. All other adults recorded with moult between 24 August and 2 December. primary moult were moulting their feathers sequentially from the innermost primaries distally. Immatures Although no adult Grey-headed Gulls were observed actively Fourteen immature Grey-headed Gulls (29%) were in active primary moulting their outermost ten secondaries (S1-S10), all birds sampled moult (Fig. 3). Immatures in active primary moult were recorded in all between December and February had these feathers newly replaced. months sampled, both in the breeding and non-breeding season. Two
Table 3. Timing and duration of primary moult in adult Grey-headed Gulls calculated from percentage feather mass grown.
Mean Duration Mean completion 95% CI for mean 95% CI for mean starting date date starting date completion date
12 October 136 days 24 January 4 October - 20 October 6 January - 11 February Durban Natural Science Museum Novitates 37 MOULT OF THE GREY-HEADED GULL 43
Fig 4. Example of immature Grey-headed Gull in active primary moult with new feathers sequentially moulting from P1 to P6 and old outer primaries P7 - P10. Fig 3. Monthly numbers of immature Grey-headed Gulls sampled in Gauteng Note the similar age of the outer secondaries S1 - S10, scored as indeterminate and Durban between April 2004 and July 2005 (black bars) and total number of age (moult score 8). Bird (SAFRING ID: 37880) trapped in Gauteng on 14 July immature Grey-headed Gulls recorded in active primary moult (white bars). 2005. birds (SAFRING IDs: 35868, 37685) trapped in July and August, and may relate to the differences in seasonal movements between respectively, and which had not initiated primary moult yet, were these species. Hartlaub’s Gull is endemic to coastal areas adjacent moulting their retrices. to the Benguela upwelling in south-west Africa where they are There was greater variation in the timing of primary moult in mostly sedentary (Williams et al. 1990). Grey-headed Gulls are immatures, compared with adults. Exceptions included: the only partly migratory/dispersive, with juveniles having been regularly two first year birds sampled in November, which were both actively recorded substantial distances from their natal colonies (Underhill moulting their inner primaries, and the two second-year birds sampled et al. 1999) and adult birds are known to move between Gauteng in January and February, respectively, that were both in the latter and Kimberley, and St Lucia and Durban (McInnes & Allan 2011). stages of primary moult. The primary moult stage of immature birds The disproportionately larger outer primary in Grey-headed Gulls trapped during the breeding season in Gauteng was more advanced could, therefore, be an adaptation for migratory/dispersal movements than that of moulting adult birds during August, e.g. for immature bird (Berthold 1993). The mean total mass of all primaries feathers (SAFRING ID: 37880) moulting its central primaries during July (Fig. 4). for Grey-headed Gulls (2.6 g) was higher than that recorded for Moulting of the outer secondaries was recorded in four immature Hartlaub’s Gull (2 g; Crawford & Underhill 2003). The Grey-headed Grey-headed Gulls: one in June (SAFRING ID: 35846), two in August Gull is on average larger than Hartlaub’s Gull (Hockey & Crawford (SAFRING IDs: 37673, 37687), and one first-year bird (SAFRING ID: 2005) and this difference is therefore expected. 37777) in November. The bird trapped in June was actively moulting The estimated mean primary moult duration for Grey-headed S5, S6, S7 (scores 2 and 3, i.e. emerging feathers to 2/3 grown) and Gulls of 136 days is intermediate between Hartlaub’s Gull (115 days, S13 (score 1, feather completely in pin), and was in the middle Crawford & Underhill 2003) and Kelp Gull Larus dominicanus vetula stages of primary moult, i.e. P5 and P6. The two birds in August were (170 days, Ward et al. 2007). These are the only gull species for which actively moulting their outermost secondaries (S1). In both birds, all primary moult duration has been calculated using the PFMG (Ward secondaries from S2 – S14 were old feathers while the innermost et al. 2007). This is probably related to the greater feather mass that secondaries (S15 – S20) were fully grown but of indeterminate age. needs to be generated by Grey-headed Gulls compared to Hartlaub’s These birds were in the middle stages of primary moult, i.e. P5 and Gulls. However, this inter-specific comparison suggests a non- P6.The first-year bird, trapped in November, was moulting most of its linear relationship between primary moult duration and body mass, secondaries (S1 – S14) simultaneously, except for S11 that was fully- suggesting that factors unrelated to feather mass may be influencing grown. Of these feathers, S1 and S14 were 1/3 to 2/3 grown (score 3) the seemingly protracted duration of primary moult in the Grey- while the remaining 11 were still contained within the feather sheath headed Gulls sampled. or just emerging (scores 1 and 2). The innermost secondaries (S15 The discrepancy between the primary moult starting dates for - S20) in this bird were all new fully-grown feathers and the stage of the six adult Grey-headed Gulls trapped in August (in Gauteng) and primary moult was centrally situated, i.e. P4 – P6. In the remaining the expected starting dates as calculated from birds mostly trapped in birds (n=6), secondary moult was limited to the innermost feathers Durban, may be related to the occurrence of two separate breeding (S15 – S20) and these birds were trapped in all sampled months other populations. Re-sightings of colour-ringed adult Grey-headed Gulls than June, August and November. in South Africa indicate a tendency for Durban’s Grey-headed Gulls Eight of the 14 immature Grey-headed Gulls in active primary to constitute, at least in part, birds that return to St Lucia during the moult were recorded with tail moult. The three birds sampled in breeding season (McInnes & Allan 2011). The breeding season for January, February and May, respectively, had no active moult in the tail. Grey-headed Gulls at St Lucia has historically been more restricted The limited sample indicates that tail moult is initiated within the early (June-September) than the range of months when they have been stages of primary moult. recorded breeding in Gauteng (January-November) (Brooke et al. 1999). The onset of primary moult towards the end of the breeding DISCUSSION season is a common occurrence in gulls and the great variation in the timing of moult, as is related to different breeding populations, The comparative proportionate masses of all primary feathers for has been documented in five other masked gull species: Slender-billed Hartlaub’s Gull, taken from Crawford & Underhill (2003), and Grey- L.genei and Black-headed L. ridibundus gulls (Cramp & Simmons 1983; headed Gull are illustrated in Fig. 5. The proportions are almost Olsen & Larsson 2003), Silver L. novaehollandiae and Black-billed L . identical, except the innermost and outermost primaries, viz. P1 and bulleri gulls (Higgins & Davies 1996), and Hartlaub’s Gull (Crawford P10. These are smaller and larger for Grey-headed Gull, respectively 44 MOULT OF THE GREY-HEADED GULL Durban Natural Science Museum Novitates 37
compared to species that employ the sequential moult strategy. Higher aspect ratios are beneficial in gliding and lower wing loading values are beneficial in take-off (Pennycuick 1986). If Grey-headed Gulls have higher aspect ratios and/or lower wing loading values compared to these other species, then they could afford to sacrifice this part of the integument temporarily. Alternatively, this strategy may have evolved in conditions that favour soaring and take-off flight movements where the temporary absence of large tracts of secondaries may be compensated for by consistent and amplified wind velocities, e.g. windy conditions such as those that persist as on-shore north-easterly winds during Durban’s summer. Of particular interest in this study was the occurrence of two first-year birds in active wing and tail moult. Our data are in support of Vangeluwe (cited in Olsen & Larsson 2003), who also recorded 1st year Grey- headed Gulls moulting flight and tail feathers in Senegal. Grey-headed Fig. 5. Comparison of proportionate masses (%) of each primary feather to Gull and probably Hartlaub’s Gull (see Crawford & Underhill 2003), total mass of all primaries for Hartlaub’s and Grey-headed gulls. are the only gull species that are known to have this moult strategy; & Underhill 2003). The predicted time of primary moult initiation, most gulls undergo their first complete moult during their second which was based mostly on a sample of birds trapped during their post-breeding season. The recent molecular phylogeny of masked non-breeding season in Durban, would coincide with the termination gulls (Given et al. 2005) recognises a strongly supported “southern of breeding activities for Grey-headed Gulls at St Lucia. The 95% clade” including Hartlaub’s, Grey-headed and Black-headed gulls, confidence limits for completion dates span a longer period than the with the former two being each others’ closest ‘relatives’ and having initiation time span, and more sampling towards the final stages of diverged as little as 70 000 years ago. The apparent limitation of moult are needed to gain more accuracy here. this moult strategy to these two species is probably indicative of The variation in timing of primary moult in immature Grey- certain selective processes happening within Africa sometime after headed Gulls is probably related to the non-reproductive state these species split from their sister species, Black-headed Gull, ca of these birds during the breeding season, the variety of different 380 000 years ago (Given et al. 2005). These selective processes may immature stages present in the sample (i.e. 1st- and 2nd-year birds), have included more demanding conditions related to flight activity evolutionary constraints associated with intraspecific competition (e.g. increased post-natal dispersion), with a subsequent accelerated for food and/or breeding sites, and the potential for these birds to wearing of these remiges and therefore the need to replace them have originated from disparate breeding localities (Harris 1971). earlier. There is a tendency for the immatures of other ‘masked’ gull species to initiate primary moult earlier than adults, e.g. Slender-billed and CONCLUSION Black-headed gulls (Higgins & Davies 1996; Olsen & Larsson 2003). This phenomenon has also been documented for larger gulls: Herring The information on moult coming from this study is the first L argentatus, Great Black-backed L. marinus and Lesser Black-backed attempt to describe this part of the life cycle of the Grey-headed L. fuscus gulls (Harris 1971), and could explain the advanced stage of Gull. The existence of geographically discrete breeding populations moult for Grey-headed Gull immatures sampled between May and on the Gauteng Highveld and northern KwaZulu-Natal lowlands August, in Gauteng. warrants a more thorough investigation into the timing of moult Despite the small sample size there is evidence that, in both between different regions. This may be complicated by a lack of adults and immatures, the onset of secondary moult coincides with knowledge of movements between these areas and a project of this the moulting of the central primary feathers. The same situation has nature would benefit by simultaneously incorporating this aspect. been recorded for Black-headed, Silver and Black-billed gulls (Cramp Finally, the validation of wing and tail moult in first-year birds, an & Simmons 1983; Olsen & Larsson 2003; Higgins & Davies 1996), unusual phenomenon for masked gull species, provides an exciting and Dwight (1925) and Harris (1971) describe this phenomenon opportunity for further research into this area. as the norm for gulls in general. There appears to be two waves of secondary moult in the Grey-headed Gull, starting with the outer ACKNOWLEDGEMENTS secondaries (S1-S10) and then followed by a second wave between S11 and S14. The absence of any observable secondary moult in This work would not have been possible were it not for the help of adult Grey-headed Gulls, despite birds having been sampled in the staff of Gauteng Nature Conservation, who contributed many successive months during the study period, is probably indicative hours to this project. In this regard we would like to convey a special of rapid moult in this region, with the possibility of simultaneous thanks to the following: Craig Whittington-Jones, Sean West, Richard moult as shown by first-year bird (SAFRING ID: 37777). Both the Koko, Job Motsamai, Jacob Makola, Andries Mputhi, Wilson Molaba and Silver and Black-billed gulls moult their outer secondaries (S1-S14) Aaron Matabane. We would like to thank the following bird-ringers sequentially inward, followed by a second wave between S15 and S19, and volunteers who contributed to the ringing effort in Gauteng and which also moult inwardly (Higgins & Davies 1996). In contrast to Durban: Pat Cochran, Joel Avni, Lynne Bingham, Terry Walls, Murray these species, results coming from this study indicate that Grey- Christian, Fran de Jager, Richard White, Greg and Fiona Brown, Koos headed Gulls are more similar to the large gulls studied by Harris Stovold and Herman Bos. Finally a special thanks to Tony Tree and an (1971) who noted these birds having “many feathers” missing in each anonymous reviewer who provided valuable comments on the original wing during secondary moult. This may be related to the relative manuscript. wing-aspect ratios and wing-loading values of these species when Durban Natural Science Museum Novitates 37 MOULT OF THE GREY-HEADED GULL 45
REFERENCES HIGGINS, P.J. & DAVIES, S.J.J.F. 1996. Handbook of Australian, New Zealand & Antarctic birds. Oxford University Press: Melbourne. BERTHOLD, P. 1993. Bird migration. Oxford University Press: Oxford. HOCKEY, P.A.R. & CRAWFORD, R.J.M. 2005. Hartlaub’s Gull Larus BROOKE, R.K., ALLAN, D.G., COOPER, J., CYRUS, D.P, DEAN, W.R.J., hartlaubii. In: HOCKEY, P.A.R., DEAN, W.R.J. & RYAN, P.G. (eds), DYER, B.M., MARTIN, A.P. & TAYLOR, R.H. 1999. Breeding Roberts birds of southern Africa, VIIth ed., pp. 444 – 446. The distribution, population size and conservation of the Grey- Trustees of the John Voelcker Bird Book Fund: Cape Town. headed Gull Larus cirrocephalus in southern Africa. Ostrich 70: MCINNES, A.M., ALLAN, D.G., BRYAN, M.C. & MERSON-DAVIES, M. 157-163. 2005. A speargun-driven net for catching gulls. AFRING News 34: CRAMP, S. & SIMMONS, K.E.L. 1983. Handbook of the Birds of Europe, 46-50. the Middle East and North Africa: the Birds of the Western Palearctic. MCINNES, A.M. & ALLAN, D.G. 2011. Distribution, abundance and Volume 3. Waders to Gulls. Oxford University Press: London. movements of the Grey-headed Gull Larus cirrocephalus in South CRAWFORD, R.J.M., & UNDERHILL, L.G. 2003. Aspects of breeding, Africa. Durban Museum Novitates 34: 1-20. moult, measurements and population trends of Hartlaub’s Gull in OLSEN, K.M., & LARSSON, H. 2003. Gulls of Europe, Asia and North Western Cape, South Africa. Waterbirds 26: 139-149. America. Christopher Helm: London. CRAWFORD, R.J.M. & HOCKEY, P.A.R. 2005. Grey-headed Gull Larus PENNYCUIK, C.J. 1986. Flight of seabirds. In: CROXALL, J.P. (ed), cirrocephalus. In: HOCKEY, P.A.R., DEAN, W.R.J. & RYAN, P.G. Seabirds: feeding ecology and role in marine ecosystems, pp. 43-62. (eds), Roberts birds of southern Africa, VIIth ed., pp. 443 – 443. The Cambridge University Press: Cambridge. Trustees of the John Voelcker Bird Book Fund: Cape Town. UNDERHILL, L.G., TREE, A.J., OSCHADLEUS, H.D. & PARKER, V. 1999. CROCHET, P., BONHOMME, F. & LEBRETON, J. 2000. Molecular Review of ring recoveries of waterbirds in southern Africa. Avian phylogeny and plumage evolution in gulls (Larini). Journal of Demography Unit, University of Cape Town: Cape Town. Evolutionary Biology 13: 47-57. UNDERHILL, L.G. & ZUCCHINI, W. 1988. A model for avian primary DWIGHT, J. 1925. The gulls (Laridae) of the world; their plumages, moult. Ibis 130: 358-372. moults, variations, relationships and distribution. Bulletin of the WARD, V.L., OSCHADLEUS, H.D. & UNDERHILL, L.G. 2007. Primary American Museum of Natural History 52: 63-411. moult of the Kelp Gull Larus dominicanus vetula in the Western GINN, H.B. & MELVILLE, D.S. 1983. Moult in birds. BTO Guide 19. Cape, South Africa. In: KIRKMAN, S. (ed.), Final Report of the British Trust for Ornithology: Tring. BCLME (Benguela Current Large Marine Ecosystem) Project on Top GIVEN, A.D., MILLS, J.A. & BAKER, A. J. 2005. Molecular evidence for Predators as Biological Indicators of Ecosystem Change in the BCLME, recent radiation in southern hemisphere masked gulls. Auk 122: pp. 201-204. Avian Demography Unit, University of Cape Town: 268-279. Cape Town. GRANT, P.J. 1978. Field identification of west Palearctic gulls. British WILLIAMS, A.J., STEELE, W.K., COOPER, J. & CRAWFORD, R.J.M. 1990. Birds 71: 145-176. Distribution, population size and conservation of Hartlaub’s Gull HARRIS, M.P. 1971. Ecological adaptations of moult in some British Larus hartlaubii. Ostrich 61: 66-76. gulls. Bird Study 18: 113-118. 46 MOULT OF THE GREY-HEADED GULL Durban Natural Science Museum Novitates 37 ;
ad (adult), imm (immature). imm (adult), ad ; Age2 – Age2 ; - year) Y (second nd , 2 Y (first Y - year) st
RL (RooikraalRL Landfill), WL (Weltevreden Landfill), BL (Blue Lagoon, Durban), SJ (Simmer & Jack Landfill) 1.
APPENDIX breeding adults), imm (immature), 1 - BA (breeding adult), NBA (non adult), (breeding BA
, refer to text for score definitions . Abbreviations: Location (Loc.) – Natal); Age1 – - Natal); Age1 headed Gulls headed - Gtg (Gauteng), KZN (KwaZulu (Gauteng), Gtg
– Province (Prov.) Moult scores for adult and immature Grey and immature adult for scores Moult Durban Natural Science Museum Novitates 37 DECREASE OF COQUI FRANCOLIN 47
DECREASE OF COQUI FRANCOLIN PELIPERDIX COQUI (AVES: PHASIANIDAE) IN SOUTHERN KWAZULU-NATAL, SOUTH AFRICA
G.B.P. DAVIES Ditsong National Museum of Natural History, P.O. Box 413, Pretoria, South Africa 0001 email: [email protected]; [email protected]
avies, G.B.P. 2014. Decrease of Coqui Francolin Peliperdix coqui (Aves: Phasianidae) in southern KwaZulu-Natal, South Africa. DDurban Natural Science Museum Novitates 37: 47-53. The change in range occupation and relative abundance of the Coqui Francolin Peliperdix coqui in southern KwaZulu-Natal (south of 29° S) was evaluated by analysing historical records, Southern African Bird Atlas Project (SABAP) data and records from the Birds in Reserves Project (BIRP). Since 1980, the Coqui Francolin has apparently disappeared from 12 quarter-degree-grid cells (15’ x 15’ grid-cells) and has been currently recorded from only six quarter-degree-grid cells (= eight pentads, 5’ x 5’ grid-cells) south of the Tugela River, with all of these grid-cells having very low indices of abundance (reporting rates < 5%). Apparently remnant sub-populations in southern KwaZulu-Natal are also isolated by large distances from each other. The decrease of the Coqui Francolin has been steeper than other broadly sympatric gamebirds, which in some cases have even increased (e.g. Natal Spurfowl Pternistis natalensis), suggesting it is a more sensitive species. Reasons for the decrease of the Coqui Francolin in southern KwaZulu-Natal remain to be elucidated but are possibly a concatenation of various factors including the transformation of its grassland and light savanna habitat for sugarcane farming, subsistence agriculture and urbanisation, intensive burning and grazing of remnant grasslands, hunting, and the indirect consequences of pesticide and herbicide usage on its food base.
KEYWORDS: atlas data, Coqui Francolin, decrease, grassland, KwaZulu-Natal, museum specimens, Peliperdix coqui.
INTRODUCTION METHODS
In this paper, the change in range occupancy and status of a small To determine the historical range in KwaZulu-Natal, details of The (approximately 250 g) gamebird, the Coqui Francolin Peliperdix coqui in Coqui Francolin study skins and eggs held in eight major South African, southern KwaZulu-Natal (south of 29° S), South Africa is investigated. European and American natural history museums were assembled Although the francolin has never been listed in any avifaunal (unpubl. data, except for Aubrecht et al. 2004). Historical primary and conservation assessment (e.g. Brooke 1984; Barnes 2000), and Little secondary data pertaining to The Coqui Francolin in KwaZulu-Natal (1997) commented that “there is no evidence that its distribution in were collated from old papers and the newsletters of bird clubs in southern Africa has changed significantly in recent times”, the author KwaZulu-Natal (between 1951 and 2014). recently became concerned regarding the status of the francolin in More recent distributional information was extracted from the southern KwaZulu-Natal when it was noticed that specimens were faunal atlases by Mentis (1974), Cyrus & Robson (1980) and Harrison lodged in natural history museums that would be impossible to et al. (1997). These references plotted distribution of species at the replicate today. Furthermore, Underhill & Brooks (2014) reported the scale of quarter-degree-grid cells (i.e. 15’ x 15’). Data in Mentis (1974) species as one of the 50 worst performing species (i.e. with lowered and Cyrus & Robson (1980) was collected between 1969 and 1980, indices of abundance) in the current Southern African Bird Atlas while data in Harrison et al. (1997; SABAP 1) was assembled between Project (SABAP 2). This prompted an investigation into the status of 1986 and 1992. the francolin by asking several questions: (a) where did the francolin Unpublished data for the Coqui Francolin was also extracted occur historically south of the Tugela River, (b) how common was it from the Birds in Reserves Project (BIRP), managed by the Animal where it occurred, (c) how has its distribution changed since then, and Demography Unit, University of Cape Town (www.birp.adu.org. (d) what factors might be operating to cause this range change? za). BIRP entailed the collection of daily checklists in protected 48 DECREASE OF COQUI FRANCOLIN Durban Natural Science Museum Novitates 37
areas throughout South Africa between 1992 and 2009. Lastly, the national vegetation map of Mucina & Rutherford (2006) to detect any distribution and reporting rates for The Coqui Francolin were broad-scale vegetation type (habitat) preferences. Information on the assessed from current unpubl. data in the South African Bird Atlas distribution and status of the francolin south of the Tugela River was Project (SABAP 2; www.sabap2.adu.org.za), which has collected data also solicited by email from experienced bird-watchers and land- on a finer, pentad (5’ x 5’ grid-cell), scale between 2007 and 2014. owners known to have properties within the historic range of the The Southern African Bird Atlas Projects calculated a simple francolin (see Acknowledgements). index of abundance (called the ‘reporting rate’) by dividing the Acronyms and abbreviations: AMNH = American Museum number of checklists with the species for that quarter-degree-grid of Natural History, New York, USA; BIRP = Birds in Reserves cell by the total number of checklists for that grid-cell expressed as a Project (1992-2009); c/ = number of eggs per clutch; DNSM = percentage (Harrison et al. 1997). A key assumption is that changes in Durban Natural Science Museum, Durban, South Africa; FMNH = reporting rate correlate with changes in bird numbers. As the Coqui Field Museum, Chicago, USA; ha = hectares; MCZ = Museum of Francolin regularly vocalises at dawn, around midday and again at dusk Comparative Zoology, Harvard University, Cambridge, USA; NHBZ = (Vincent 1934), and has vocalisations dissimilar to other gamebirds Natural History Museum, Bulawayo, Zimbabwe; NHM = The Natural (Little 2005), it is assumed that the atlas data adequately represent History Museum, Tring, United Kingdom; NHRM = Royal Natural its distribution and approximate status. This assumption may not be History Museum, Stockholm, Sweden; OOLM = Museum of Upper valid, however, given the different levels of expertise by participants Austria, Linz, Austria; QDGC = quarter-degree-grid-cell; SABAP 1 in SABAP and the secretive nature of this francolin when not calling = First Southern African Bird Atlas Project (1986-1992); SABAP 2 and raising chicks (J. van Niekerk in litt.). In the absence of other = Second Southern African Bird Atlas Project (2007-present); TM = quantitative data, the SABAP information has had to be treated at Ditsong National Museum of Natural History, Pretoria, South Africa. face-value. For analysis the data was divided into four time periods, pre-1965, RESULTS 1969-1980, 1980-1992 and 1992-recent. The reasons for the divisions are that active collecting of museum skins in KwaZulu-Natal largely Distribution over time ended around the mid-1960s; the second period covers the period Prior to 1965 of the first systematic attempts to plot avian distribution ranges The Coqui Francolin was recorded from 13 QDGC from 1839 to in KwaZulu-Natal on a grid-cell basis; the third period covers the 1965 inclusive, based on 27 specimen records (Table 1; Fig. 1) and SABAP 1 atlas, and the last period covers BIRP and SABAP 2. Where literature references as to its occurrence at Durban North (da point-locality data were available this was manually plotted on the Fonseca 1960), Shongweni Dam (Oatley 1964) and Greytown (McLean
Table 1. Specimens (study skins, taxidermic mounts and eggs) of Coqui Francolin Peliperdix coqui from southern KwaZulu-Natal, South Africa in natural history museum collections arranged chronologically. Vegetation Types are as per Mucina & Rutherford (2006).
Museum reg. no. Locality Vegetation Specimen Date Collector type type
NHRM 554638 Congella, Durban CB 3 skin 19 Aug 1840 J.A. Wahlberg NHRM 6854 Congella, Durban CB 3 skin 24 June 1839 J.A. Wahlberg AMNH 541097 Ifafa River CB 3 skin Undated (late 1800s) Woodward brothers? NHM 1891.5.1.20 Pinetown, Durban CB 3 skin 19 Mar 1874 G.E. Shelley NHM 1891.5.1.20 Durban CB 3 skin 25 Mar 1874 G.E. Shelley NHM 1891.5.1.19 Pinetown, Durban CB 3 skin 10 June 1875 T. L. Ayres DNSM 34376 Fields Hill, Pinetown, Durban CB 3 C/5 10 Mar 1902 A.D. Millar OOLM 1941/1531 Kronsberg, Noodsberg SVs 4 skin 11 Jan 1905 A.G.H. Rudatis OOLM 1941/1532 Hlutankungu, Jolivet SVs 4/5 skin 5 June 1905 A.G.H. Rudatis OOLM 1941/1533 Fairfield, Jolivet SVs 4/5 skin 7 June 1905 A.G.H. Rudatis OOLM1941/1534 Fairfield, Jolivet SVs 4/5 skin 8 June 1905 A.G.H. Rudatis OOLM 1941/1536 Jolivet SVs 4/5 skin 11 June 1905 A.G.H. Rudatis OOLM 1941/1535 Fairfield, Jolivet SVs 4/5 skin 15 Aug 1905 A.G.H. Rudatis NHBZ Not known Durban CB 3 C/2 October 1917 James (1970) SAM-ZE060294 Mount Edgecombe, Durban CB 3 C/3 20 Mar 1918 A.D. Millar FMNH 406372 Umzumbe CB 3 skin 3 Oct 1925 R. Cowles FMNH 406373 Umzumbe CB 3 skin 1 Mar 1926 R. Cowles FMNH 406374 Umzumbe CB 3 skin Undated (1920s) R. Cowles NHM 1933.7.14.106 Coombe Barton Farm, Richmond SVs 4 skin 23 Aug 1926 J. Vincent TM 14571 Mt Edgecombe, Durban CB 3 mount Jun 1927 W.A. Campbell TM 14572 Mt Edgecombe, Durban CB 3 mount Jun 1927 W.A. Campbell NHM 1933.7.14.105 Whitson Farm, Umhlongo Nek SVs 4 skin 22 Sep 1929 J. Vincent TM 187597 Near Kloof SVs 5 skin 14 Feb 1932 R.B. Wilson MCZ 98169 Karkloof Gs 9/10 skin Undated (1950s) H. Friedmann TM 187601 Currie’s Post Gs 9 skin 17 Aug 1951 P.A. Clancey DNSM 137 Drummond SVs 5 skin 11 July 1954 Unknown DNSM 27634 Ntuzuma, Inanda, Durban CB 3 skin 29 Jul 1971 J. White Durban Natural Science Museum Novitates 37 DECREASE OF COQUI FRANCOLIN 49
reporting rates (Table 2) for Albert Falls Dam Nature Reserve (45%) and Lake Eland Reserve (33%). The BIRP data suggest (Table 2) that the francolin is absent or extremely rare in most southern KwaZulu-Natal nature reserves and protected areas, such as Vernon Crookes, Oribi Gorge and Cumberland, where the francolin might be anticipated to occur. No museum specimen records are available for this period. The existing sub-populations are apparently centred on Albert Falls Dam Nature Reserve and Oribi Flats, west of Port Shepstone (SABAP 2). These sub-populations are isolated by a distance of approximately 140 km from each other. The nearest populations north of the Tugela River are in the Vryheid-Dannhauser- Babanango and Mtubatuba districts, approximately 160 km and 220 km respectively away from the Albert Falls population (SABAP 2).
Overall trends in historical distribution The number of occupied QDGC in the respective periods is summarised in Fig. 1, and since more thorough coverage began in the second time period, the spatial pattern of occurrence is consistently negative, with the sub-population west of Port Shepstone now particularly isolated (Fig. 1). Since 1980, the Coqui Francolin has Fig. 1. Distribution of Coqui Francolin in southern KwaZulu-Natal, South Africa by apparently disappeared from 12 QDGC (Fig. 1) with especially severe quarter-degree-grid cells over four temporal periods: (A) pre-1965, (B) 1969-1980, (C) 1986-1992 (SABAP 1), (D) 2007-2014 (SABAP 2). Note the steady attrition in contraction in the greater Pietermaritzburg and Durban areas (Fig. 1). range occupancy since 1980. Association with vegetation types & Jackson 1966). The number of QDGC occupied during this early Table 1 provides 27 point localities and seven additional point- period must be taken as a conservative figure given the fragmentary localities are available for Albert Falls Dam Nature Reserve (BIRP), and incomplete coverage of the study area. Contemporary accounts Beaumont-Faraway Farm (Eston; J. Davies pers. comm.), Cato Ridge from this period reveal that the species was “generally dispersed all (Tait 1989), Durban North (da Fonseca 1960), Minnehaha Farm (Oribi over the colony of Natal” (Ayres in Gurney 1860: 215), that “these Flats; M. Neethling pers. comm.), Lake Eland Reserve (Oribi Flats; birds are plentiful throughout Natal, although more partial to the BIRP) and Shongweni Dam (Oatley 1964). The point-locality records coast lands” (Millar in Stark & Sclater 1900: 199), and Vincent (1934: for Coqui Francolin cover seven vegetation types (Table 3), with 332) also implied it was common in Natal. According to Lawson the majority of point-localities falling within KwaZulu-Natal Coastal (1966), in Durban it was “[u]ncommon, though may be found wherever Belt (37.2%), Ngongoni Veld (23.3%) and KwaZulu-Natal Sandstone suitable areas still exist throughout”. At Beaumont (Faraway) Farm, Sourveld (16.3%). Eston district (7 km south-west of Shongweni Dam), the francolin was “fairly common in light Acacia veld and the edges of wattle Comparison with other gamebirds in southern KwaZulu-Natal plantations” from the late 1940s to the 1950s (J.A. Davies pers. Over the three atlassing periods, and relative to other broadly- comm.). sympatric gamebirds such as Natal Spurfowl Pternistis natalensis and Shelley’s Francolin Scleroptila shelleyi, the Coqui Francolin has 1969-1980 performed worst, with a nett decrease of 12 grid-cells versus five in The Coqui Francolin was recorded from 18 QDGC between 1969 Shelley’s Francolin, and an increase of 12 grid-cells in Natal Spurfowl and 1980 inclusive, based on one museum record (Table 1; Fig. 1) and (Fig. 2). The Coqui Francolin also has by far the lowest grid-cel atlas data in Mentis (1974) and Cyrus & Robson (1980). No indices of occupancy currently (Fig. 2). Although not shown in Fig. 2, two other abundance can be extracted from the atlas data in Mentis (1974) and gamebirds (Helmeted Guineafowl Numida meleagris and Common Cyrus & Robson (1980). Contemporary accounts indicate the species Quail Coturnix coturnix) have occupancies of > 50 grid-cells in southern was “[c]ommon in the low-lying and intermediate regions” (Mentis KwaZulu-Natal in the SABAP 2 period. 1974: 21; with the caveat that this comment may have applied to the entire province). No other published statements on its abundance in DISCUSSION southern KwaZulu-Natal during this time frame have been traced. To answer the questions posed in the Introduction, it is clear that the SABAP 1 (1986-1992) Coqui Francolin occupied large parts of southern KwaZulu-Natal east The Coqui Francolin was recorded from 13 QDGC between 1986 of about 30° E and as far south as the Port Shepstone area (31° 45’ S). and 1992 inclusive (Fig. 1). No museum specimen records are available All published pre-1965 references reflect that the francolin was locally for this period. No contemporary accounts of its status for the study common. Woodward & Woodward (1899) even cited an informant area have been traced in the published literature. Bird-watchers active who mentioned a strong increase in numbers in the Howick district. at that period recall it was “common” at Albert Falls Dam Nature One caveat is that there are no historical quantitative population Reserve (R.G.C. Boon in litt., late H.A. Campbell pers. comm.). estimates or indices for any sub-populations in KwaZulu-Natal prior to SABAP 1. Since at least 1980 there has been a consistent negative BIRP (1992-2009) and SABAP 2 (2007-2014) trend in range occupation and today, aside from Albert Falls Nature The Coqui Francolin was recorded from six QDGC (covering eight Reserve and Lake Eland Reserve (Oribi Flats), it is scarce in southern pentads) in SABAP 2 between 2007 and present (Fig. 1). Reporting KwaZulu-Natal with extremely low reporting rates in both BIRP rates in southern KwaZulu-Natal in both the BIRP and SABAP 2 and SABAP 2. Relative to other broadly-sympatric gamebirds such data were very low, generally < 5%. The only exceptions were BIRP as Shelley’s Francolin, Natal Spurfowl, Helmeted Guineafowl and 50 DECREASE OF COQUI FRANCOLIN Durban Natural Science Museum Novitates 37
Table 2. Birds in Reserves Project (BIRP) data for significant protected areas north and south of the Tugela River.
Protected Area Total # of # of checklists Reporting Rate checklists with Coqui Francolin
South of Tugela River Albert Falls Nature Reserve 126 57 45.2 Bisley Valley Nature Reserve 64 0 0.0 Cumberland Nature Reserve 128 0 0.0 Hazelmere Dam 20 0 0.0 Hillandale Conservancy (Bartlett Estate) 89 0 0.0 Lake Eland Nature Reserve 3 1 33.3 Midmar Dam 73 1 1.4 Oribi Gorge Nature Reserve 255 1 0.4 Nagle Dam 19 0 0.0 Umgeni Valley Nature Reserve 174 1 0.6 Umsuluzi Lodge, Colenso 7 1 14.2 Vernon Crookes Nature Reserve 252 0 0.0
North of Tugela River Amatikulu Nature Reserve 69 0 0.0 Bonamanzi Private Game Reserve 93 2 2.2 Goudhoek Farm, Babanango 145 43 29.7 Hluhluwe Game Reserve 117 0 0.0 Imfolozi Game Reserve 103 2 1.9 Ithala Game Reserve 148 3 2.0 Mkhuze Game Reserve 552 0 0.0 Ndumo Game Reserve 250 2 0.8 Phinda Private Game Reserve 14 1 7.1
Common Quail, it is evident that the Coqui Francolin has suffered cells, I am satisfied that the data correctly reflect that a significant greater attrition in grid-cell occurrence (Fig. 2), suggesting it has been change in range occupation by the species has occurred. more sensitive to changes in the environment. The Coqui Francolin has been identified as an inhabitant of dense, A key assumption in this paper has been that the absence from fairy tall grasslands, savanna and well-grassed woodland (Little 1997, particular grid-cells in later atlassing periods is genuinely reflective of 2005), especially areas with paler, sandier soils (Van Niekerk & Von non-occurrence by the francolin. Without a systematic programme of Ginkel 2003). The vegetation analysis for southern KwaZulu-Natal ground-truthing, this assumption cannot be directly tested. However, broadly supported these published conclusions and suggested a several lines of circumstantial evidence suggest that the pattern positive relationship between the Coqui Francolin and KwaZulu- revealed by the atlas data is reliable. Firstly, I encountered Shelley’s Natal Coastal Belt (CB 3), Ngongoni Veld (SVs 4) and KwaZulu-Natal Francolin on the outskirts of Durban during the 1990s and 2000s (e.g. Sandstone Sourveld (SVs 5) vegetation types of Mucina & Rutherford at Mount Moreland and the upper Umbogintwini River), but never (2006). The CB 3 is a mosaic of habitats including various types the Coqui Francolin. Like the Coqui Francolin, Shelley’s Francolin can of subtropical coastal forest, and grassland (Mucina & Rutherford be highly unobtrusive when not calling, and thus the difference does 2006: 578-579), but the Coqui Francolin would only be found in the not appear related to furtiveness of the former species. Secondly, experienced bird-watchers who have regularly visited remnant areas Table 3. Number of point-localities for Coqui Francolin Peliperdix coqui of grassland and light savanna in the greater Durban area during the organised by vegetation type (Mucina & Rutherford 2006). Note some localities 1990s and 2000s have never encountered the Coqui Francolin (R.G.C. are not refined enough to unambiguously place to vegetation type in certain instances; in these cases the locality is scored to all vegetation types in the Boon in litt., D. Styles in litt.). Thirdly, fairly recent, localised extinction general point-locality. events have occurred in southern KwaZulu-Natal, although the evidence is anecdotal and not amenable to formalised analysis. For Vegetation Type No. of point localities example, the Coqui Francolin occurred regularly on the Beaumont CB 3 – KZN Coastal Belt 16 (Faraway) Farm, Eston from the late 1940s to the 1950s (J.A. Davies CB 4 – Pondoland/Ugu Sandstone Coastal Sourveld 2 pers. comm.), but began to decrease when the grassland and light SVs 3 – KZN Hinterland Thornveld 3 thornveld on the farm was transformed to sugarcane fields such that, SVs 4 –Ngongoni Veld 10 by the 1980s, it was much scarcer (late E.B. Hampson pers. comm.) SVs 5 – KwaZulu-Natal Sandstone Sourveld 7 and had entirely disappeared by the 1990s (late M.A. Hampson pers. Gs 9 – Midlands Mistbelt Grassland 3 comm., M.B. Hampson pers. comm., pers. obs). Consequently, although GS 10 – Drakensberg Foothill Moist Grassland 2 isolated coveys may have been overlooked in the some of the grid- Durban Natural Science Museum Novitates 37 DECREASE OF COQUI FRANCOLIN 51
grassland components. Most museum collecting for birds was done at or close to Durban (a CB 3 area), and thus this artefact may have skewed the specimen records towards that vegetation type. Ngongoni Veld (SVs 4), dominated by the grass Aristida junciformis (Poaceae), is an ‘artificial’ vegetation type, the likely consequence of many decades of heavy livestock grazing (Scott-Shaw & Morris 2014; R.G.C. Boon pers. comm.), a form of pastoralism that encourages the dominance of that grass species (Tainton 1999). The SVs 4 is probably the transformed remnants of areas that were formerly SVs 3 (KwaZulu-Natal Hinterland Thornveld), SVs 5 (KwaZulu-Natal Sandstone Sourveld) and CB 3 grasslands. The factors that might have caused the decrease of the Coqui Francolin in southern KwaZulu-Natal are formally unstudied, but chief among them is likely to be the anthropogenic transformation of grassland and light savanna. Extremely little of the preferred habitat of
the Coqui Francolin remains intact south of the Tugela River, although Fig. 2. Comparison of number of grid cells occupied by three, broadly published figures of the extent, proportions and rates of habitat sympatric gamebirds (Natal Spurfowl, Shelley’s Francolin and Coqui transformation in KwaZulu-Natal are scarce. Mucina & Rutherford Francolin) in southern KwaZulu-Natal, South Africa, during the three (2006) stated that 68% of KwaZulu-Natal Sandstone Sourveld, 50% atlassing periods. Coqui Francolin shows a nett decrease of -12 (18 to 6 of KwaZulu-Natal Coastal Belt, 39% of Ngongoni Veld and 22% of grid-cells), Shelley’s Francolin a nett decrease of -5 (31 to 26 grid-cells) and Natal Spurfowl a nett gain of +12 (37 to 49 grid-cells). KZN Hinterland Thornveld has been “transformed” for cultivation, plantations, urban development and road building. By the early 1990’s, Von Ginkel 2003). It is probable that there is little or no exchange only approximately 1% of Ngongoni Sourveld (the Acocks Veld Type of individuals currently between the remaining sub-populations of approximately equating to KZN Sandstone Sourveld, Ngongoni Veld the Coqui Francolin in southern KwaZulu-Natal, given the inimical and Coastal Belt) was protected in formal conservation areas in intervening matrix (e.g. sugarcane fields, timber plantations, high- KwaZulu-Natal (Scott-Shaw et al. 1996). The area under sugarcane in density rural populations). The sub-population on the Oribi Flats, in KwaZulu-Natal has also grown substantially from 6680 ha in 1904, to particular, is apparently now isolated by > 100 km from the nearest 145,389 ha by 1936, approximately 323,000 ha by the late 1960s and other birds. Negative demographic consequences of fragmentation around 420,000 ha by the early 2000’s (G.B.P. Davies unpubl. data), may also be operative as Milstein (1989) noted that “decreasing much of it having been planted on grassland and light savanna formerly populations [of Coqui Francolin] seem to develop a preponderance amenable to the Coqui Francolin. The anecdotal observation from of males” and in a brood reared from a wild nest at Cato Ridge, Tait Beaumont Farm, Eston mentioned above serves as an example. (1989) only had a single female in four chicks. These observations are Other predominant inimical factors may include annual burning consistent with Donald’s (2007) conclusion that the adult sex ratio of grasslands and light thornveld, over-grazing, fragmentation per se, in threatened/decreasing bird species invariably becomes skewed commercial and subsistence gamebird hunting, and pesticide/herbicide towards the male sex. usage in agriculture. Annual burning has widely been identified as Potential negative impact from organised, commercial gamebird especially harmful to the Coqui Francolin (e.g. Ayres in Gurney 1860; hunting is unknown, as published hunting statistics for KwaZulu-Natal Meyer 1971; Irwin 1981; Milstein 1989). Due to the nutrient-poor do not appear to be available. There is also no published, quantitative soils in much of KwaZulu-Natal (especially those soils derived from data on subsistence and recreational hunting in communal land but sandstone and quartzite), livestock farmers burn their grasslands it is likely to be severe as snaring and hunting with packs of dogs annually to provide grazing with sufficient protein content and, on the appears common (e.g. Pooley 1967; Anon. 1991; pers. obs in the Upper Oribi Flats, west of Port Shepstone, regular burning has “I believe … Umbogintwini and Umzinto districts). Much earlier, Woodward & caused a drop in Coqui numbers” (M. Neethling in litt.). Although fire Woodward (1899: 160) indicated that rural peoples “kill and snare a would have been an intrinsic feature of the habitats occupied by the number”. Coqui Francolin, the inferred increase in burning frequency (above the The use of agrochemicals such as dieldrin and aldicarb (‘Temik’), ‘natural’ baseline situation of lightning-induced fires and less intensive which are directly harmful to wildlife, was evidently widespread in burning by pre-colonial human communities) may have impacted the the KwaZulu-Natal sugarcane industry at least until the 1970’s (e.g. francolin negatively by changing the sward physiognomy and botanical Carnegie & Wood 1972). Current data on the usage of such chemicals composition of the grasslands and open savanna. in KwaZulu-Natal agriculture is sparse, but Ratcliffe & Crowe (2001) Over-grazing has likewise been implicated in decreases of the found the use of 31 different types of herbicide and 28 different types Coqui Francolin through much of its southern African range (e.g. of insecticide by farmers in the KwaZulu-Natal Midlands during the Irwin 1981; Milstein 1989; Herremans in Little 1997). Dramatic 1990’s. The impact of agrochemicals may be especially pertinent in the transformation of grass and forb species composition due to intensive case of the Coqui Francolin because soft-leaved forbs (which are often grazing by livestock has been demonstrated for KwaZulu-Natal coastal the ‘weeds’ targeted by agricultural herbicides) and invertebrates grasslands (Scott-Shaw & Morris 2014). Many remnant grassland areas (which are often negatively impacted by generic insecticides) appear in regions inhabited by high density rural human populations are important in the diet of the species (e.g. Meyer 1971; Tait 1989), heavily grazed, with a very short grass sward (pers. obs), presumably possibly far more so than in Scleroptila francolins and Pternistis conditions unsuitable for the Coqui Francolin. Furthermore, sensitivity spurfowl, which tend to feed on geophytic corms (Scleroptila) and to both annual burning and heavy grazing has been found in the Red- fallen seeds and grain (Pternistis). winged Francolin Scleroptila levaillantii (Jansen et al. 1999) and Orange Other specific deleterious impacts, both natural and River Francolin S. levaillantoides (Van Niekerk 2012). anthropogenic, mentioned by others, include fatalities from road traffic Fragmentation of suitable habitat is liable to have serious effects (Wolff & Milstein 1987: 52), collisions with game fences when the for the Coqui Francolin given their sedentary nature (Van Niekerk & 52 DECREASE OF COQUI FRANCOLIN Durban Natural Science Museum Novitates 37
francolin are put to flight (I. Weiersbye in litt.), disease (Van Niekerk Murray Hampson, Geoff Nichols, Mike Neethling, David Styles and & Verwoerd 2002), nest predators (Tait 1989) and predation by small Ingrid Weiersbye. carnivores such as mongoose species (M. Neethling in litt.). This paper has focused on southern KwaZulu-Natal, but there REFERENCES are indications that a diminution in numbers and range occupation of the Coqui Francolin has also taken place in parts of northern ANONYMOUS. 1991. Hunting in KwaZulu. African Wildlife 45(2): 96. KwaZulu-Natal. At Mtunzini, Garland (1967: 5) observed that it was AUBRECHT, G., SCHIFTER, H., PLASS, J., WEIGL, S. & ALLAN, D.G. “at one time freely seen – resident and breeding – [but] no records 2004. The A.G.H. Rudatis bird (Aves) collection (1904-1906) from for about ten years” i.e. since late 1950’s. The extent of coastal Natal (KwaZulu-Natal) at the Museum of Upper Austria. Durban grassland at Mtunzini (demarcated by the area east of the N2 highway, Museum Novitates 29: 31-49. south of the Umlalazi River, west of the Indian Ocean and from the BARNES, K. (ed.) 2000. The ESKOM Red Data Book of Birds of South southern municipal boundary) has shrunk from approximately 500 Africa, Lesotho and Swaziland. BirdLife South Africa & Avian ha in 1937 to < 5 ha in 2014 (G.B.P. Davies unpubl. analyses of aerial Demography Unit: Johannesburg & Cape Town. photographs). Macdonald & Birkenstock (1980) likewise recorded a BROOKE, R.K. 1984. The rare and vulnerable birds of South Africa. sharp reduction in the Coqui Francolin at both Hluhluwe and Imfolozi South African National Sciences Programmes Report 97: 1-203. game reserves since the 1960s. Substantial changes in the vegetation CARNEGIE, A.J.M. & WOOD, R.A. 1972. Sugarcane pesticides and and birdlife of Hluhluwe-Imfolozi have been in progress over several their residue analyses. Proceedings of the 46th South African Sugar decades and the deterioration in general gamebird numbers has been Technologists’ Association no volume: 220-223. tentatively attributed to heavy nest predation by chacma baboons CYRUS, D. & ROBSON, N. 1980. Bird atlas of Natal. University of Natal Papio ursinus and Pied Crows Corvus albus (Macdonald 1984: 620). Press, Pietermaritzburg. These extinctions and decreases appear permanent because there DA FONSECA, A.H. 1960. The birds of a Durban North garden. Natal were no records for this species at Mtunzini during SABAP 1 and Bird Club News Sheet 64: 2. SABAP 2, and records from just one pentad each (reporting rates < DONALD, P.F. 2007. Adult sex ratios in wild birds. Ibis 149: 671-692. 5%) for Hluhluwe and Imfolozi during SABAP 2. The Coqui Francolin GARLAND, I. 1967. List of birds on the farm Twinstreams, Mtunzini is also curiously absent or exceedingly rare in Ithala, Mkhuze and district, Zululand. South African Avifaunal Series 46. Ndumo game reserves (Table 2), although the light savanna in at least GURNEY, J.H. 1860. On birds collected in the colony of Natal, in parts of these reserves might have been deemed suitable. On the south-eastern Africa. Ibis 2(7): 203-221. other hand, the Coqui Francolin appears to be fairly common in the HARRISON, J.A, ALLAN, D.G., UNDERHILL, L.G., HERREMANS, M., Vryheid-Dannhauser-Babanango triangle of KwaZulu-Natal (SABAP TREE, A.J., PARKER, V. & BROWN, C.J., 1997. The atlas of southern 2 data), an area that is dominated by Income Sandy Grassland (Gs 7; African birds, vol. 1. BirdLife South Africa: Johannesburg. Mucina & Rutherford 2006). IRWIN, M.P.S. 1981. The Birds of Zimbabwe. Quest: Salisbury. In conclusion, this study has documented the apparently JAMES, H.W. 1970. Catalogue of the bird eggs in the collection of the precipitous regional decrease of a gamebird species not listed in any National Museums of Rhodesia. Trustees of the National Museums Red Data assessment (e.g. Brooke 1984; Barnes 2000). Causes for of Rhodesia: Salisbury. the decrease point towards habitat modification and fragmentation, JANSEN, R., LITTLE, R.M. & CROWE, T.M. 1999. Implications of but insufficient evidence exists to rule out the involvement of grazing and burning of grasslands on the sustainable use of other causes. The available data also suggest the Coqui Francolin francolins (Francolinus spp.) and on overall bird conservation in is also more sensitive to landscape modification in KwaZulu-Natal the highlands of Mpumalanga province, South Africa. Biodiversity than other co-occurring gamebirds (e.g. Natal Spurfowl, Helmeted and Conservation 8: 587-602. Guineafowl, Shelley’s Francolin). Future work should attempt to LAWSON, W.J. 1966. Check list of the birds of Durban. Privately ground-truth the results of this mainly desk-top study, empirically printed, Durban. investigate the postulated causes of decrease, and attempt to LITTLE, R.M. 1997. Coqui Francolin. In: HARRISON, J.A., ALLAN, elucidate why the Coqui Francolin appears more sensitive to D.G., UNDERHILL, L.G., HERREMANS, M., TREE, A.J., PARKER, V. landscape changes in southern KwaZulu-Natal than other gamebirds. & BROWN, C.J. (eds), The Atlas of Southern African Birds, Vol.1, pp. Close monitoring of the remnant sub-populations of the Coqui 276-277. BirdLife South Africa: Johannesburg. Francolin in southern KwaZulu-Natal, especially at Albert Falls Nature LITTLE, R.M. 2005. Coqui Francolin. In: HOCKEY P.A.R, DEAN, W.R.J., Reserve, is also recommended. & RYAN, P.G. (eds) Roberts – Birds of Southern Africa, 7 th edition, pp. 62-63. John Voelcker Bird Book Fund: Cape Town. ACKNOWLEDGEMENTS MACDONALD, I.A.W. 1984. An analysis of the role of the Hluhluwe- Umfolozi Game Reserves Complex in the conservation of the The unpublished data contained in the Birds in Reserves Project and avifauna of Natal. In: LEDGER, J. (ed.), Proceedings of the Fifth Second Southern African Bird Atlas (SABAP 2), managed by the Animal Pan-African Ornithological Congress, pp. 601-637. Southern African Demography Unit, has proven critical for this paper. Information of Ornithological Society: Johannesburg. specimens in natural history museums in the United States of America MACDONALD, I.A.W. & BIRKENSTOCK, P.J., 1980. Birds of the was sourced from the ORNIS website (www.ornisnet.org). Data in Hluhluwe-Umfolozi Game Reserve complex. Lammergeyer 29: the Natural History Museum, Tring, was extracted from an electronic 1-56. database of South African specimens compiled by W.R.J. Dean and McLEAN, T. & JACKSON, G. 1966. The birds of Greytown and district. supplied by M. Adams. David Allan provided data held at the Durban Natal Bird Club News Sheet 142: 2-4. Natural Science Museum. Richard Peek visited the Bulawayo Museum MEYER, H.F. 1971. The Coqui Francolin. Honeyguide 65: 29-30. to check that collection on my behalf. For answering queries and MENTIS, M.T. 1974. Distribution of some wild animals in Natal. providing information I thank Richard Boon, Hugh Chittenden, Jenifer Lammergeyer 20: 1-68. Davies, the late Evelyn Brian Hampson, the late Michael Hampson, Durban Natural Science Museum Novitates 37 DECREASE OF COQUI FRANCOLIN 53
MILSTEIN, P. LE S. 1989. Coqui Francolin. In: GINN, P.J., MCILLERON, TAIT, C.C. 1989. Observations on a brood of Coqui Francolins. Birding W.G. & MILSTEIN, P. LE S. (eds), The Complete Book of Southern in Southern Africa 41(3): 70-72. African Birds. Struik: Cape Town. UNDERHILL, L.G. & BROOKS, M. 2014. Preliminary summary of MUCINA, L. & RUTHERFORD, M.C. (eds) 2006. The Vegetation of South changes in bird distributions between the first and second Africa, Lesotho and Swaziland. South African National Biodiversity Southern African Bird Atlas Projects (SABAP 1 and SABAP 2). Institute: Pretoria. Ornithological Observations 5: 258-293. OATLEY, T. 1964. List of birds of Shongweni area. Natal Bird Club News VAN NIEKERK, J.H. 2012. Habitat use by Orange River Francolin Sheet 109: 4-6. Scleroptila levaillantoides on farmland in the Heidelberg district, POOLEY, A.C. 1967. Illegal trapping. Bokmakierie 19(3): 72-73. Gauteng province, South Africa. Ostrich 83(1): 43-49. RATCLIFFE, C.S. & CROWE, T.M. 2001. Declining populations of VAN NIEKERK, J.H. & VERWOERD, D.J. 2002. Avian pox in Swainson’s Helmeted Guineafowl in the midlands of KwaZulu-Natal. South francolin in South Africa. South African Journal of Wildlife Research African Journal of Wildlife Research 31(3&4): 161-171. 32(1): 43-48. SCOTT-SHAW, C.R., BOURQUIN, O. & PORTER, R.N. 1996. The VAN NIEKERK, J.H. & VAN GINKEL, C.M. 2003. Notes on the conservation status of Acocks’ veld types in KwaZulu-Natal. behavioural ecology of Coqui Francolin in the Rustenburg district, Lammergeyer 44: 50-63. South Africa. South African Journal of Wildlife Research 33(1): 59-62. SCOTT-SHAW, C.R. & MORRIS, C.D. 2014. Grazing depletes forb VINCENT, J. 1934. The birds of northern Portuguese East Africa, part species diversity in the mesic grasslands of KwaZulu-Natal, South 3. Ibis 4(2), 13th series: 305-340. Africa. African Journal of Range & Forage Science DOI:10.2989/1022 WOLFF, S.W. & MILSTEIN, P. LE S. 1987. Limiting factors for gamebirds 0119.2014.901418. in southern Africa. South African Journal of Wildlife Research TAINTON, N.M. 1999. Veld management in South Africa. University of Supplement 1: 51-53. Natal Press: Pietermaritzburg. WOODWARD, R.B. & WOODWARD, J.D.S. 1899. Natal birds. Davis & Sons: Pietermaritzburg. 54 GORONGOSA WATERBIRDS Durban Natural Science Museum Novitates 37
A MAJOR WATERBIRD BREEDING COLONY AT LAKE UREMA, GORONGOSA NATIONAL PARK, MOÇAMBIQUE
STALMANS M.1, DAVIES G.B.P.* 2, TROLLIP J. 1 & POOLE G. 1 1 Gorongosa Restoration Project, Chitengo, Sofala Province, Moçambique 2 Ditsong National Museum of Natural History, P.O. Box 413, Pretoria, South Africa *Author for correspondence: [email protected]; [email protected]
talmans, M., Davies, G.B.P., Trollip, J. & Poole, G. 2014. A major waterbird breeding colony at Lake Urema, Gorongosa National SPark, Moçambique. Durban Natural Science Museum Novitates 37: 54-57. A very large breeding colony of eight waterbird species, comprising 5003 nests and active over March and April 2014 in flooded (0.5-1 m deep), open Faidherbia-Acacia woodland, is described from Lake Urema, Gorongosa National Park, Moçambique. The colony contained breeding Yellow-billed Stork Mycteria ibis (983 nests), African Openbill Anastomus lamelligerus (531 nests), Reed Cormorant Phalacrocorax africanus (2276 nests), African Darter Anhinga rufa (547 nests), Great Egret Egretta alba (330 nests), White-breasted Cormorant (230 nests), Grey Heron Ardea cinerea (82 nests) and African Sacred Ibis Threskiornis aethiopicus (24 nests). The number of Yellow-billed Storks and African Darters present during the 2014 count exceeds the 1% criterion for listing as a Ramsar site of international importance.
KEYWORDS: Gorongosa, heronry, Moçambique, nests, waterbirds.
INTRODUCTION trees were counted from a boat within ca 20 - 100 m of the colony. All nests were individually counted per nesting tree and per species. This note provides quantitative data on a large waterbird breeding colony (‘heronry’) on Lake Urema, Gorongosa National Park, RESULTS Sofala Province, Moçambique. Although there have been tangential references to significant numbers of waterbirds breeding and feeding The colony was located at the south-east corner of Lake Urema (Fig. on Lake Urema in the past (e.g. Tinley 1971), there have been no 1) and extended across a distance of 1.6 km from 18° 54’ 2.4” S; 34° published counts of the waterbirds, as far as we are aware, and Parker 30’ 1.7” E to 18° 54’ 38.1” S; 34° 30’ 44.3” E (covering approximately (2001: 635) observed that “the wetlands in the National Park have not 15 ha). Nesting activity in the colony was initially detected in early been comprehensively surveyed [for birds]”. The size of the breeding March 2014 when the area was first accessed. The birds nested in colony documented below indicates that Lake Urema is a significant open, flooded woodland dominated by ana trees Faidherbia albida, nesting site for certain waterbirds. fever trees Acacia xanthophloea and paper-bark acacias A. sieberana Lake Urema is a shallow, permanent lake (ca 1.6 m deep) located (Figs 2 and 3). Only F. albida trees seemed to be used for nesting. within the Urema Graben, a continuation of the East African Rift During the April survey, these trees were standing in water that was System. The extent of the lake fluctuates widely between ca 10-25 0.5 to 1.0 m deep (Figs 2 and 3). Ground observations of water and km2, with extreme flooding events of up to ca 105 km2, dependent on mud marks on the nesting trees during the 2014 dry season indicated seasonal rainfall, annual flooding by the Vunduzi and Nhandue rivers, that maximum flood levels (probably attained in February and March) and groundwater recharging (Tinley 1977: 31-33, plate 22; Böhme et had been approximately 2 m deep. al. 2006). The lake has a large, seasonally-inundated floodplain (tando) Eight species of medium- to large-bodied waterbirds were found on its perimeter. Maximal flooding of the lake and adjoining floodplain nesting at the colony during the count of 7 April (e.g. Figs 4 and 5) occurs between January-March (Tinley 1977: 145). with a total of 5003 active nests in 463 trees. Reed Cormorants On 7 April 2014 a detailed count was undertaken by the authors Phalacrocorax africanus exceeded 2000 active nests and Yellow- of a large waterbird colony discovered on the lake edge. The nesting billed Storks Mycteria ibis approached 1000 active nests (Table 1). Durban Natural Science Museum Novitates 37 GORONGOSA WATERBIRDS 55
Fig. 2. Representative view of the Lake Urema waterbird breeding colony situated in leafless Faidherbia trees in flooded water. Waterbird species breeding in the area pictured include Yellow-billed Stork, African Sacred Ibis and Reed Cormorant (photo Piotr Naskrecki).
In Moçambique, aside from Gorongosa, significant numbers of waterbirds also nest in the 11,000 km2 Marromeu Complex, Zambezi Delta (Dutton 1987; Beilfuss & Bento 1998; Bento 2000). Indications are that nesting waterbird numbers at Marromeu exceed those of Lake Urema (R. Beilfuss in litt.) as “three immense breeding colonies Fig. 1. Map of Gorongosa National Park, Moçambique, showing the location of of storks and herons” have been observed in the central part of Lake Urema and the waterbird breeding colony described in this paper. that massive wetland (Beilfuss & Bento 1998). When flooded, the ephemeral Banhine wetland (6588 ha) in Gaza Province (Stalmans & Assuming that each nest represented a pair of birds, the Urema colony Wishart 2005) also attracts very large numbers of nesting waterbirds contained approximately 10,000 breeding waterbirds. (Pietersen & Pietersen 2010; D.W. Pietersen pers. comm.) but detailed More than 30% of the nesting trees held three or more species counts are not yet available from that wetland. It should also be of birds (Table 2). A maximum of five nesting species were recorded borne in mind that many significant wetlands in Moçambique remain per breeding tree. The maximum number of nests recorded for a poorly surveyed or completely unknown for nesting waterbirds, e.g. single breeding tree was 58, mostly Reed Cormorants. Some vertical large coastal wetlands in northern Moçambique. Nonetheless, even stratification of the birds was observed, with Reed Cormorants if comprehensive data were to become available from these other occupying the higher, thinner branches and African Openbill Anastomus wetlands, it is still likely that Lake Urema would rank highly. lamelligerus being found on the lowest branches. The breeding colony at Lake Urema is also large when compared Although the colony was active since at least early March, against ‘heronries’ in Botswana (e.g. Tyler & Hancock 2006; Tyler 2011; breeding was staggered for the respective species. African Sacred Ibis Brown 2012; Muller & Flatt 2013), Zambia (e.g. Leonard 2001, 2005; Threskiornis aethiopicus were only observed starting to nest a week Dowsett et al. 2008) and Tanzania (Baker & Baker 2001), with the before the survey, i.e. late March. On the other hand, by the time of caveat that many major wetlands in Zambia (e.g. Lukanga Swamps) and the survey, the chicks of African Darter Anhinga rufa and Great Egret Tanzania (e.g. Wembere Steppe) are largely inaccessible when flooded Egretta alba were well grown, whereas Yellow-billed Storks were only and the full extent of regular waterbird breeding in such wetlands is starting to hatch. The nesting birds were relaxed and allowed approach inadequately known. within 30 m or less of the nesting trees by the boat, except for some diving by African Darters at the northern end of the colony.
DISCUSSION
The above figures indicate that Lake Urema is a highly significant waterbird breeding locality. The attributes that make Urema attractive as a breeding locality are probably the large amounts of food (fish and aquatic invertebrates) provided by the lake and the receding flood waters of the adjacent floodplain, as well as the protection from disturbance provided by its location deep within the national park. There is very little past information on this Urema colony, but informal observations by the authors between 2009 and 2013 indicate that the site is an annually active colony and not an episodic occurrence. Currently, there are no direct threats to the breeding colony as the site is at least 20-25 km from any human settlements Fig. 3. General aspect of the Lake Urema waterbird breeding colony showing (e.g. Muanza town), except for the park headquarters (Chitengo), extensive flooding of open, largely leafless, woodland and including a view of the which lie approximately 17 km to the south-west. The general area is boat used to visit the colony (photo Bob Poole). also patrolled by the park’s game guards. 56 GORONGOSA WATERBIRDS Durban Natural Science Museum Novitates 37
Table 1. Results of count of colonial waterbirds breeding at Lake Urema, Gorongosa National Park, on 7 April 2014.
Species No. of No. trees Average Max. nests nests with nests nests/tree per tree African Darter Anhinga rufa 547 118 4.6 25 Reed Cormorant Phalacrocorax africanus 2276 219 10.4 53 White-breasted Cormorant P.lucidus 230 54 4.3 22 Great Egret Egretta alba 330 131 2.5 12 Grey Heron Ardea cinerea 82 66 1.2 3 African Sacred Ibis Threskiornis aethiopicus 24 4 6.0 12 African Openbill Anastomus lamelligerus 531 112 4.7 13 Yellow-billed Stork Mycteria ibis 983 259 3.8 23
Total (all species) 5003 463 10.8 58
Table 2. Number of bird species per nesting tree.
Number of bird Frequency of trees species per tree (%) 1 35.2 2 31.7 3 23.8 4 8.4 5 0.9
With regard to individual species, the count of nesting Yellow- billed Storks is especially noteworthy. An indication of the relative size of this stork aggregation may be gauged by comparing the number of nests (893) against published data from Botswana, Moçambique, Zambia and Tanzania. The main nesting sites for Yellow- billed Stork in Botswana (e.g. in Moremi Game Reserve and Gadikwe Heronry) have varied between six and 108 nests or breeding pairs (Gaosafelwe et al. 1997; Tyler et al. 2002; Tyler & Hancock 2006: table 1; Brown 2012), although colonies of “a few hundred” have also been reported from unspecified localities in the Okavango Delta (Anderson Fig. 5. Yellow-billed and African Openbill stork nests and adults in the 1997). A huge colony of “approximately 500-1000 pairs” of Yellow- Lake Urema breeding colony; African Sacred Ibis also present (photo Piotr billed Storks was observed at one colony in the Marromeu Complex, Naskrecki). Zambezi Delta by Beilfuss & Bento (1998). In Zambia, Dowsett et al. southern and eastern Africa (1000 birds; Wetlands International 2012). (2008: 95) reported Yellow-billed Storks nesting in only three areas, None of the other waterbird nest numbers meet the 1% Ramsar with colonies in the Luangwa Valley of “up to 150 pairs”. In Tanzania, criterion. approximately 800 breeding pairs were recorded at Lake Manyara To date, the importance of Lake Urema for nesting waterbirds in May 2002 (N. Baker per T. Dodman in litt.). The large numbers has gone unreported in the ornithological and conservation of nesting Yellow-billed Storks at Urema exceeds the 1% Ramsar literature, e.g. neither Clancey (1996) nor Parker (2005) in their threshold for sub-Saharan Africa of 870 birds (Wetlands International synoptic Mozambican works mentioned ‘heronries’ at Lake Urema. 2012). The count of 547 African Darter nests (equating to 1094 The presence of this major waterbird breeding colony also heightens birds) also just exceeds the 1% Ramsar threshold for this species in the importance of Gorongosa National Park as a conservation area. Potentially, this wildlife spectacle could also be an important ecotourism drawcard in future and augment the ongoing efforts to rehabilitate the National Park as a leading ecotourism destination and biodiversity sanctuary.
ACKNOWLEDGEMENTS
We thank Jen Guyton (Princeton University) and Piotr Naskrecki (Harvard University) for helpful comments, and the latter also for the use of his photos. Constructive criticism of the paper was provided by Richard Beilfuss (International Crane Foundation), Tim Dodman (Wetlands International), and Stephanie Tyler (BirdLife Botswana). Fig. 4. African Openbill nests and adults in the Lake Urema breeding colony (photo Piotr Naskrecki). Durban Natural Science Museum Novitates 37 GORONGOSA WATERBIRDS 57
REFERENCES LEONARD, P.M. 2005. Important Bird Areas in Zambia: priority sites for conservation. Zambian Ornithological Society: Lusaka. ANDERSON, M.A. 1997. Yellowbilled Stork. In: HARRISON, J.A., MULLER, M. & FLATT, A. 2013. A survey of the Kanana heronry, 20-22 ALLAN, D.G., UNDERHILL, L.G., HERREMANS, M., TREE, A.J., October 2012. Babbler 58: 2-8. PARKER, V., & BROWN, C.J. (eds), The atlas of southern African PARKER, V. 2001. Moçambique. In: FISHPOOL, L.D.C. & EVANS, birds, Vol. 1, pp. 100-101. BirdLife South Africa: Johannesburg. M.I. (eds), Important Bird Areas in Africa and associated islands: BAKER, N.E. & BAKER, L.M. 2001. Tanzania. In: FISHPOOL, L.D.C. & priority sites for conservation, pp. 627-638. BirdLife International: EVANS, M.I. (eds), Important Bird Areas in Africa and associated Cambridge, UK. islands: priority sites for conservation, pp. 897-945. BirdLife PARKER, V. 2005. The atlas of the birds of central Moçambique. International: Cambridge, UK. Endangered Wildlife Trust & Avian Demography Unit: Johannesburg BEILFUSS, R.D. & BENTO, C. 1998. Impacts of hydrological changes & Cape Town. on the Marromeu Complex of the Zambezi Delta, with special PIETERSEN, D.W. & PIETERSEN, E.W. 2010. Annotated checklist attention to the avifauna. In: DAVIES, B.R. (ed.), Proceedings of of the birds of Banhine National Park, southern Moçambique. the Workshop on the Sustainable Use of Cahora Bassa Dam and Ornithological Observations 1: 7-37. the Zambezi Valley, 29 September to 2 October 1997. Arquivo STALMANS, M. & WISHART, M. 2005. Plant communities, wetlands Patrimonio do Cultura (ARPAC): Maputo, Moçambique. and landscapes of the Parque Nacional de Banhine, Moçambique. BENTO, C. 2000. Wetland bird survey of the Zambezi Delta. In: Koedoe 48: 43-58. TIMBERLAKE, J. (ed.), Biodiversity of the Zambezi Basin Wetlands, Vol. TINLEY, K.L. 1971. Sketch of Gorongosa National Park. In: 2, pp. 259-278. Zambezi Society: Harare. ANONYMOUS (ed.), Nature conservation as a form of land use - BROWN, C.J. 2012. Heronry at Kasane Rapids. Babbler 57: 14-15. Proceedings of the SARCCUS Symposium, pp. 163-172. Government CLANCEY, P.A. 1996. The birds of southern Moçambique. African Bird Printer: Pretoria. Book Publishing: Westville, Durban. TINLEY, K.L. 1977. Framework of the Gorongosa Ecosystem. DSc. DOWSETT, R.J., ASPINWALL, D.R. & DOWSETT-LEMAIRE, F. 2008. The thesis, University of Pretoria, South Africa. birds of Zambia: an atlas and handbook. Tauraco Press & Aves: Liège, TYLER, S.J. & HANCOCK, P. 2006. Heronries in Botswana. Babbler 48: Belgium. 18-39. DUTTON, P. 1987. A qualitative survey of the wetlands of TYLER, S.J. 2011. A review of waterbird counts in Botswana 1991- Moçambique. South African Journal of Wildlife Research Supplement 2010. Babbler Special Supplement 4. BirdLife Botswana: Maun. 1: 9-11. TYLER, S.J., TYLER, L., ORCHARD, C. & ORCHARD, D. 2002. A visit to GAOSAFELWE, E.S., HANCOCK, P. & MADONGO, M.H. 1997. A count Xakanaxa and Gadikwe ‘heronries’ in September 2001. Babbler 40: of Marabou and Yellowbilled storks breeding at Gadikwe, Xhobega 18-22. and Xakanaxa in Moremi Game Reserve during January 1997. WETLANDS INTERNATIONAL. 2012. Waterbird population estimates Babbler 32: 26-28. 5: Population details, estimates and trends, www.wpe.wetlands. LEONARD, P.M. 2001. Zambia. In: FISHPOOL, L.D.C. & EVANS, M.I. org/Imaps (eds), Important Bird Areas in Africa and associated islands: Priority Sites for Conservation, pp. 1005-1024. BirdLife International: Cambridge, UK. 58 WHITE-BROWED SPARROW-WEAVER IN KWAZULU-NATAL Durban Natural Science Museum Novitates 37
WHITE-BROWED SPARROW-WEAVER PLOCEPASSER MAHALI (AVES: PLOCEIDAE) IN KWAZULU-NATAL, SOUTH AFRICA
H. DIETER OSCHADLEUS Animal Demography Unit, Department of Zoology, University of Cape Town, Rondebosch 7701, South Africa email: [email protected]
schadleus, H.D. 2014. White-browed Sparrow-Weaver Plocepasser mahali (Aves: Ploceidae) in KwaZulu-Natal, South Africa. Durban ONatural Science Museum Novitates 37: 58-61. Records of the White-browed Sparrow-Weaver Plocepasser mahali in KwaZulu-Natal have increased in frequency over recent decades. A total of 24 records are documented from a variety of sources and these serve as a baseline from which to monitor future records of this species in the region. Two early records dated from 1910 and 1958, and the remaining records are from 1977 onwards. The only confirmed breeding records in KwaZulu-Natal are from Canigou Farm near Estcourt. The White-browed Sparrow-Weaver in KwaZulu-Natal most likely originated from the west (Free State), although birds in Zululand may have visited from the north (Mpumalanga). The apparently increasing frequency of the White-browed Sparrow-Weaver in KwaZulu-Natal may be related to increasing bush encroachment. Factors limiting its range expansion in KwaZulu-Natal may include rainfall and diet. Continuing to monitor this species at the edge of its range may provide clues to its ecological requirements.
KEYWORDS: KwaZulu-Natal, Plocepasser mahali, range records, White-browed Sparrow-Weaver.
INTRODUCTION METHODS
White-browed Sparrow-Weavers Plocepasser mahali are cooperatively Digital biodiversity data sets held at the Animal Demography Unit, breeding passerine birds of eastern and southern Africa. They live in collected largely by citizen scientists, were searched for records of pairs or small groups year-round consisting of a dominant breeding White-browed Sparrow-Weavers in KwaZulu-Natal. These projects pair and subordinate, non-breeding males and females (Collias & are the South African Bird Ringing Unit (SAFRING, 1948-to date), Collias 1978; Lewis 1982). Breeding is usually restricted to the Birds in Reserves Project (BIRP, 1992-to date), the Southern African rainy season, particularly November to February, but breeding of Bird Atlas Project (SABAP1, 1987-1991), the Southern African Bird neighbouring groups is not necessarily synchronized (Earlé 1983). In Atlas Project (SABAP2, July 2007-to date), and the Nest Record southern Africa, the White-browed Sparrow-Weaver occurs mainly Cards (NRC, 1952-to date). Published references were based on an in dry woodland and savanna (Colahan 1997). Its main stronghold unpublished bibliography of the Ploceidae family. Some specimen data is in north-central southern Africa, with isolated populations in the are included from visits to museums and databases received from Eastern Cape and eastern Free State (Colahan 1997). museum curators. Unpublished sightings emailed to the author are Clancey (1964) mentioned one record of this species in included. The ADU has a digital database of bird club outings, but KwaZulu-Natal, South Africa, but in recent decades the number of the original cards have not been traced – these records are thus not records has increased. Biotic changes in distribution, and the rates at included in the table of results but are mentioned in the Results. which these are occurring, are important to document, especially in The raw data covering White-browed Sparrow-Weaver records the light of global climate change. This paper details all records of the in KwaZulu-Natal were extracted from bird-atlas projects. Other White-browed Sparrow-Weaver in KwaZulu-Natal up to December records were included if there was a date (at least to the resolution 2014, based on a variety of sources. of a year) and a locality. Coordinates to the nearest minute were assigned to localities obtained from the literature. Records appearing Durban Natural Science Museum Novitates 37 WHITE-BROWED SPARROW-WEAVER IN KWAZULU-NATAL 59
Table 1. Confirmed, geo-referenced records of the White-browed Sparrow-Weaver in KwaZulu-Natal, listed chronologically. * Vegetation types: 1 - Valley Thicket, 2 - Natal Lowveld Bushveld, 3 - NE Mountain Grassland, 4 - Natal Central Bushveld, and 5 - Wet Cold Highveld Grassland.
Locality Dates Veg. type Observer Sources
Pongola River 13 Jul. 1910 2 F.O. Noomé Ditsong Nat. Mus. Nat. Hist.-6413 Cathkin Park 2-10 Oct. 1958 3 D.R. Calder Calder 1959 Canigou farm, Estcourt 1977 4 Les Nutting Nutting 1979, Cyrus & Robson 1980 Canigou farm, Estcourt 1978 4 Les Nutting Nutting 1979, Cyrus & Robson 1980 Canigou farm, Estcourt 20 Oct. 1979 4 Les Nutting Nutting 1979, Cyrus & Robson 1980 Canigou farm, Estcourt 11 Jan. & 1 Feb. 1980 4 Les Nutting NRC 780.001 Spioenkop Dam Mar. 1982 4 Ian Trench Johnson 1982 2829DA [Spioenkop] 18 Jul. 1984 4 Unknown SABAP1-049156 Spioenkop Dam 8 Sept. 1984 4 I. Trench Cyrus 1985 2729DC [S of Memel] Oct. 1984 [5 or 3] Unknown SABAP1-049190 2730DC [W of Vryheid] Feb. 1988 [4 or 3] Unknown SABAP1-058636 2730DC [W of Vryheid] March 1988 [4 or 3] Unknown SABAP1-058637 Weenen N.R. 30 Nov. 1996 1 W.L. Branford BIRP-2600965 Spioenkop N.R. 16 Jun. 2000 4 C.N. Burne BIRP-2602679 Spioenkop N.R. 30 Apr. 2001 4 Chris Mathews BIRP-2506798 Cotswold farm Aug. 2001 4 Tony Roberts pers. comm. Glencoe dump site Aug. 2005 4 Peter Dearlove pers. comm. Hluhluwe GR 4 Nov. 2007 2 Andrew Kruger Davis 2008a, Davis 2008b Pongola NR 22 Nov. 2007 2 Tim Wood BIRP-2509655 Spioenkop Dam NR 16 Dec. 2007 4 E. Russell, S. du Preez & G. Leisegang pers. comm. Ndumo GR, main camp 25-26 Oct. 2008 2 Francois Furstenburg Davis 2008c, Davis 2009a Pentad 2850_2955 [Weenen] 6 Jul. 2012 4 Johan de Villiers Davis 2012, SABAP2 Pentad 2850_3000 [Weenen] 24 Sept. 2013 4 Michael Cunningham SABAP2 Pongola NR 29 Sept. 2013 2 Unknown Davis 2013
in more than one source were cross-referenced. Habitat categories birds and on 20 October 1979 there were 13 nests with two or three were assigned to White-browed Sparrow-Weaver records, according having one entrance (Nutting 1979), indicating breeding. There are to the South African vegetation map (Low & Rebelo 1996). monthly bird club records from June to December 1979. Mr L. Nutting submitted a Nest Record Card, noting an eggshell below the nests RESULTS on 11 January 1980 and the adults at the nest on 1 February. Cyrus (1982) gives one confirmed record for the species in KwaZulu-Natal There are 24 dated records of White-browed Sparrow-Weavers in for the period 1980-1982, presumably referring to the latter breeding KwaZulu-Natal (Table 1, Fig. 1). No records were found for KwaZulu- record. Natal in the SAFRING database. In March 1982 there was a record from Spioenkop Dam by Ian The earliest dated record of a White-browed Sparrow-Weaver Trench (Johnson 1982). A bird club field card indicates that this in KwaZulu-Natal is from 13 July 1910. Specimen TM_6413 from the sighting was on 14 March. This was followed by an atlas card on 18 Ditsong National Museum of Natural History (formerly Transvaal July 1984 for the grid covering the eastern part of the dam (the Museum), Pretoria, South Africa, was collected by F.O. Noomé, a entrance side), and presumably the record was from the dam area. Museum taxidermist, on the Pongola River. This record was not known On 8 September 1984 Ian Trench again recorded this weaver at to Clancey (1964), presumably since the collection of the Ditsong Spioenkop Dam; this being the third published record for the species Museum had not then been digitized. in KwaZulu-Natal (Cyrus & McCosh 1985), and a bird club field card Calder (1959) reported a group of up to six birds seen at El was submitted. Further records from this locality were on 16 June Mirador hotel, Cathkin Park (28°59’S; 29°28’E); several were seen daily 2000 and 30 April 2001 (BIRP records) and 16 December 2007 (Elena from 2-10 October 1958 (Calder 1958, 1959). Calder saw no nests Russell, Sandi du Preez and Gill Leisegang, as reported on the internet and attributed the presence of the birds as being related to below- listserver: SABirdNet). No nests have been reported from this locality. average rainfall in the eastern Free State at that time. In October 1984 the White-browed Sparrow-Weaver was recorded The next record is from the farm of Mr G. Reynolds: Canigou from the quarter-degree-grid cell 2729DC on the border of the Free Farm (29°03’S; 29°59’E), 12 km ESE of Estcourt (Nutting 1979). A State and KwaZulu-Natal (SABAP1); this atlas card was labelled as pair built a nest in a gum tree in the cattle kraal near the farm house KwaZulu-Natal. In February and March 1988 there were SABAP1 atlas (NRC, Nutting 1979). Up to six birds were present from 1977 to 1980 records of White-browed Sparrow-Weavers from 2730DC, west of (Nutting 1979, Cyrus & Robson 1980). Mr Reynolds saw five or six Vryheid. The exact localities and observers for these SABAP1 records 60 WHITE-BROWED SPARROW-WEAVER IN KWAZULU-NATAL Durban Natural Science Museum Novitates 37
The closest region to central KwaZulu-Natal where most of the White-browed Sparrow-Weaver records come from is the Free State, where the species is widespread (Skead 1966; Colahan 1997). This suggests that the records in central KwaZulu-Natal relate to birds originating from the west. Clancey (1964) noted the distribution as the western Free State, east as far as between Winburg and Senekal. The earliest records east of 28°45’E are from 1984, soon after the Free State atlas project (1983-1986) began (Earlé & Grobler 1987). It is not clear if the White-browed Sparrow-Weavers in Zululand originated from the west (Free State) or from the north (Mpumalanga). Although particular in its micro-habitat requirements (Vernon 1983; Ferguson 1985), the White-browed Sparrow-Weaver finds suitable conditions in a variety of vegetation types, with high reporting rates in particular from the dry woodland and savanna areas, and the much-altered Sweet Grasslands of central southern Africa. Nest trees are usually at the ecotone between two habitats: one with reduced ground cover, where the birds forage, and the Fig. 1. Confirmed, geo-referenced records of the White-browed Sparrow- Weaver for KwaZulu-Natal and the eastern Free State; white circles 1836-1970, other having good grass cover, providing nest material (Vernon 1983; solid triangles 1970-1989, solid squares 1989-2009. Also shown are SABAP1 Ferguson 1985). Such ecotones have been extensively generated in records (1987-1991, quarter-degree-grid cells, Harrison et al. 1997), with darker the Sweet Grasslands by crop farming and livestock grazing. In these shades of grey indicating higher reporting rates. areas alien trees planted as windbreaks, or along roads, are commonly used for nesting (Colahan 1997). KwaZulu-Natal probably offers are not captured in the database. On 30 November 1996 there was a a variety of habitats near human dwellings that the White-browed BIRP record from Weenen Nature Reserve (28°56’S; 30°04’E), some Sparrow-Weaver can utilize. 25 km from Canigou Farm. The apparently increasing records of the White-browed More recent sightings are from Cotswold Farm (28°04’S; 30°19’E) Sparrow-Weaver in KwaZulu-Natal may be related to increasing where three birds and roost nests were seen in August 2001 by Tony bush encroachment. Woody cover has increased in the Weenen area Roberts. Birds and nests were seen at the Glencoe dump (28°10’S; (Hoffman & O’Connor 1999) and in the Hluhluwe area (Wigley et al. 30°09’E) in August 2005 by Peter Dearlove; a week later only two 2009, Wigley et al. 2010) over the last century, both areas in which the nests were seen at the site by Tony Roberts. David Hoddinott, Tony White-browed Sparrow-Weaver has been recorded. Too much bush Roberts and Garth Aiston did not find them here in March 2006. The encroachment, however, would be a limiting factor for this species species was seen in Hluhluwe Game Reserve (~28°04’S; 32°04’E) on 4 (Colahan 1997). November 2007 by Andrew Kruger and this record was accepted by There have been slight range expansions of this species in other the KZN Regional Rarities Committee (Davis 2008a, 2008b). On 25- parts of southern Africa. The White-browed Sparrow-Weaver has 26 October 2008 a pair was seen in the main camp of Ndumo Game increased in numbers and range in the Eastern Cape, spreading Reserve (26°53’S; 32°16’E) (Davis 2008c). southwards through the Cradock district since 1950 (Skead 1966, On 6 July 2012 Johan de Villiers recorded the species in a 1967; Collett 1982). The distribution map in Colahan (1997) also SABAP2 pentad near Weenen (Davis 2012) and a year later Michael shows records from the northern and eastern Transvaal beyond Cunningham recorded it in an adjacent pentad on 24 September the eastern edge of the range shown by Tarboton et al. (1987). It is 2013. These records may have been in the Weenen Nature Reserve spreading towards and into the Kruger National Park (Arkell 1997; and are not far from Canigou Farm. A few days later this species Lawson 2010; SABAP 2) and bush encroachment is suspected to be a was recorded in the Pongola Nature Reserve, reported initially on contributing factor (Arkell 1997). SA Rare Bird Forum and subsequently published (Davis 2013). No The White-browed Sparrow-Weaver has not become permanently records from 2014 were found. established in KwaZulu-Natal, in spite of the apparent suitable The habitat categories for White-browed Sparrow-Weaver habitat and increasing records of birds. Possible reasons may be records from KwaZulu-Natal were mostly Natal Central Bushveld related to rainfall (Ferguson 1989). Another possibility is the lack of (Table 1). At least 14 records came from this vegetation type; Hodotermes harvester termites in KwaZulu-Natal (Ian A.W. Macdonald two were in in Natal Lowveld Bushveld, and one in NE Mountain pers. comm.). The importance of termites in the diet of the White- Grassland. Habitat was not determined for the SABAP1 records, due browed Sparrow-Weaver varies, however. For instance, Ferguson to the large grid size spanned by these records. (1985) notes that beetles were the most important arthropod taken. While the optimal ecological factors required by the White-browed DISCUSSION Sparrow-Weaver are not clearly understood, continuing to monitor this species at the edge of its range (including KwaZulu-Natal) may There are two early records from KwaZulu-Natal, in 1910 and 1958 provide clues to its ecological requirements. (Table 1). From the 1970’s there are some records of this species in KwaZulu-Natal every decade. The only confirmed breeding records in ACKNOWLEDGEMENTS KwaZulu-Natal are on Canigou Farm near Escourt. The birds appear to have arrived on this farm in 1977 and remained until about 1980. SAFRING is sponsored by the South African National Biodiversity Nests were built at some other sites but no breeding records were Institute and the Namibian Ministry of Wildlife and Tourism. The obtained. University of Cape Town provides accommodation and services. Durban Natural Science Museum Novitates 37 WHITE-BROWED SPARROW-WEAVER IN KWAZULU-NATAL 61
SAFRING data, Nest Record Cards and Cape Bird Club field cards EARLÉ, R.A. 1983. Aspects of the breeding biology of the whitebrowed were computerized at the ADU with funds from SABIF via the NRF sparrowweaver Plocepasser mahali (Aves: Ploceidae). Navorsinge van and SANBI. The records from the Ditsong National Museum of die Nasionale Museum, Bloemfontein 4: 177-191. Natural History were supplied by Tamar Cassidy. Thanks to David EARLÉ, R. & GROBLER, N. 1987. First atlas of bird distribution in the Hoddinott, Tony Roberts, Garth Aiston, Peter Dearlove, Gill Leisegang, Orange Free State. National Museum: Bloemfontein. Steve Davis, Richard Dean and Carl Vernon for providing information. FERGUSON, J.W.H. 1985. The influence of crop-farming on Richard Dean, Les Underhill, David Johnson and Ian A.W. Macdonald Whitebrowed Sparrow-weavers Plocepasser mahali. Proceedings of a provided comments on earlier drafts. Symposium on Birds and Man, Johannesburg 1983: 45-54. FERGUSON, J.W.H. 1989. Preferred habitat of White-browed Sparrow REFERENCES Weavers Plocepasser mahali. South African Journal of Zoology 24: 1-10. ARKELL, G. 1997. Whitebrowed Sparrow-weavers. Hornbill 47: 21. HOFFMAN, M.T. & O’CONNOR, T.G. 1999. Vegetation change over CALDER, D.R. 1958. [No title]. Natal Bird Club News Sheet 58: 1-2. 40 years in the Weenen/Muden area, KwaZulu-Natal: evidence CALDER, D.R. 1959. White-browed Sparrow-weaver Plocepasser mahali. from photo-panoramas. African Journal of Range & Forage Science Ostrich 30: 163. 16(2&3): 71-88. CLANCEY, P.A. 1964. The Birds of Natal and Zululand. Oliver & Boyd: JOHNSON, D. 1982. New distributional data. Albatross 269: 9-10. London LAWSON, P. 2010. Interesting local sightings. Hornbill 98: 26-27. COLAHAN, B.D. 1997. Whitebrowed Sparrowweaver Plocepasser LEWIS, D.M. 1982. Cooperative breeding in a population of White- mahali. In: HARRISON, J.A., ALLAN, D.G., UNDERHILL, L.G., browed Sparrow Weavers Plocepasser mahali. Ibis 124: 511-522. HERREMANS, M., TREE, A.J., PARKER, V. & BROWN, C.J. (eds), The LOW, A.B. & REBELO, A.G. (eds). 1996. Vegetation of South Africa, atlas of southern African birds. Vol. 2: Passerines, pp. 532-533. BirdLife Lesotho and Swaziland. Dept Environmental Affairs & Tourism: South Africa: Johannesburg. Pretoria. COLLIAS, N.E. & COLLIAS, E.C. 1978. Cooperative breeding behavior NUTTING, L. 1979. White-browed Sparrow Weavers in Natal. Albatross in the White-browed Sparrow Weaver. Auk 95: 472-484. 257: 4. COLLETT, J. 1982. Birds of the Cradock District. Southern Birds 9: 1-76. SKEAD, C.J. 1966. White-browed Sparrow-weaver, Plocepasser mahali A . CYRUS, D. & ROBSON, N.F. 1980. Bird atlas of Natal. University of Smith, in the eastern and north-eastern Cape. Ostrich 37: 128-129. Natal Press: Pietermaritzburg. SKEAD, C.J. 1967. Ecology of birds in the Eastern Cape Province. CYRUS, D. 1982. Observations of rare and out-of-range birds in Natal Ostrich Supplement 7: 1-103. (1980-82). Albatross 269: 6-9. TARBOTON, W.R., KEMP, M.I. & KEMP, A.C. 1987. Birds of the Transvaal. CYRUS, D. & MCCOSH, B. 1985. Recent field notes. N.B.C. recorder & Transvaal Museum: Pretoria. rare birds panel. Albatross 281: 6-8. VERNON, C.J. 1983. Aspects of the habitat preference and social DAVIS, S. 2008a. KwaZulu-Natal Honorary Recorder’s report. 7 May behaviour of the White-browed Sparrow Weaver. Honeyguide 113: 2008. KZN Birds 23: 21-22. 11-14. DAVIS, S. 2008b. KwaZulu-Natal Honorary Recorder’s report. August WIGLEY, B.J., BOND, W.J. & HOFFMAN, M.T. 2009. Bush 2008. KZN Birds 24: 21-24. encroachment under three contrasting land-use practices in a DAVIS, S. 2008c. KwaZulu-Natal Honorary Recorder’s report. mesic South African savanna. African Journal of Ecology 47: 62-70. November 2008. KZN Birds 25: 20-23. WIGLEY, B.J., BOND, W.J. & HOFFMAN, M.T. 2010. Thicket expansion DAVIS, S. 2012. Honorary Recorder’s report. KZN Birds 37: 26-29. in a South African savanna under divergent land use: local vs. DAVIS, S. 2013. Honorary Recorder’s report. KZN Birds 40: 28-31. global drivers? Global Change Biology 16: 964–976. 62 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37
RECENT RECORDS OF THE WHITE- WINGED FLUFFTAIL SAROTHRURA AYRESI (AVES, SAROTHRURIDAE) IN SOUTH AFRICA, INCLUDING DETAILS OF A SURVEY OF HIGH-ALTITUDE WETLANDS IN 2013-14
G.B.P. DAVIES1, H.A. SMIT-ROBINSON2, I.M.M. DRUMMOND3, B. GARDNER4, S. RAUTEN- BACH5, D. VAN STUYVENBERG6, C. NATTRASS7, M. PRETORIUS8, D.W. PIETERSEN9 & C.T. SYMES10
1Ornithology Section, Ditsong National Museum of Natural History, P.O. Box 413, Pretoria 0001, South Africa Author for correspondence: [email protected]; [email protected] 2Terrestrial Bird Conservation Programme, BirdLife South Africa, P.O. Box 515, Randburg 2125, South Africa 3Middelpunt Wetlands Trust, P.O. Box 47410, Parklands 2121, South Africa 4Johannesburg Zoological Gardens, Private Bag X13, Parkview 2122, South Africa 5c/o Middelpunt Wetlands Trust, P.O. Box 47410, Parklands 2121, South Africa 6Stellaland Raptor Project Coordinator, P.O. Box 928, Montana Park 0159, South Africa 7Witwatersrand Bird Club, P.O. Box 641, Cresta 2118, South Africa 8Reserve Manager, Seekoeivlei Nature Reserve, 9 Voortrekker Street, Memel 2970, South Africa 9Department of Zoology and Entomology, University of Pretoria, Private X20, Hatfield, Pretoria 0028, South Africa 10School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Private Bag 3, Wits 2050, South Africa
avies, G.B.P., Smit-Robinson, H.A., Drummond, I.M.M., Gardner, B., Rautenbach, S., van Stuyvenberg, D., Nattrass, C., Pretorius, DM., Pietersen, D.W. & Symes, C.T. 2014. Recent records of the White-winged Flufftail Sarothrura ayresi (Aves, Sarothruridae) in South Africa, including details of a survey of high-altitude wetlands in 2013-14. Durban Natural Science Museum Novitates 37: 62-75. The White-winged Flufftail Sarothrura ayresi is a poorly known, globally threatened species that only occurs with any regularity in South Africa and Ethiopia. A survey of 13 high-altitude South African wetlands, including nine important sites for the flufftail, was undertaken in the summer of 2013-14 to attempt to assess the flufftail’s current status. Additionally, all miscellaneous sightings of White-winged Flufftail in South Africa since the last (2005) review of its general status and distribution in South Africa were collated. The wetland survey was largely unsuccessful in detecting White-winged Flufftail, the species only being recorded at one of the 13 wetlands. Twenty- four miscellaneous sighting episodes comprising 32-34 flufftails between 2005 and the summer of 2014-2015 were available from eleven localities in eastern South Africa, most (46%) from Middelpunt Wetland, Mpumalanga Province. The records span September to March inclusive. Sighting episodes usually comprised one flufftail and never more than four birds at any particular wetland. Ageing, morphometric and moult details of three caught White-winged Flufftails are provided. Twenty-two sighting episodes (92%) were in high-altitude wetlands with 50-100 cm high sedges and hygrophilous grasses (cover 80-100%), on shallowly flooded ground (water depth 5-25 cm). Two of the post-2005 records were at low altitude in rather atypical habitat (edge of savanna air-strip and close to an airport run-way). No records were made on the basis of vocalisations, perhaps because of difficulties associated with differentiating the flufftail calls from the roosting (unison) calls of Grey Crowned Crane Balearica regulorum.
KEYWORDS: conservation, Sarothrura ayresi, South Africa, wetland, White-winged Flufftail, waterbirds. Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 63
INTRODUCTION
The White-winged Flufftail Sarothrura ayresi is one of Africa’s rarest birds with most records emanating from Ethiopia and South Africa (Collar et al. 1985; Taylor 1994, 1999, 2000 b, 2005 d, Taylor et al. 2004; Allan et al. 2006). The South African population has been estimated at approximately 200-250 birds in 2005 (Taylor 2005 d) and it is listed as ‘Critically Endangered’ in the South African Red Data Book (M. Taylor in prep.) and by BirdLife International (2015). The species was the subject of dedicated study in South Africa and Ethiopia by Taylor (1994, 1997 a, b, 1999) and Taylor et al. (2004), research which greatly improved our knowledge of the species. In South Africa, the flufftail was detected in small to moderate numbers, usually on the basis of auditory cues, at nine sites during the 1990s, typically in permanent, high-altitude Carex-dominated peat wetlands (Taylor 1994, 1997 a, b, 2000 b, 2005 d). Given the paucity of information on the flufftail, its precarious conservation position and the lack of clarity on its general status in Fig. 1. Immediate area where White-winged Flufftail flushed on 14 December 2013, Middelpunt Wetland, Mpumalanga, showing knee-high Pycreus sedges in South Africa, especially whether it is a regular migrant from Ethiopia fairly dense formation in very shallow (< 10 cm deep) water and cover 90-100%. or not, it is important for details of all sightings to be published. Our Photo: G. Davies. objective in this paper is to document all records of White-winged Flufftail made in South Africa since Taylor (2005 d) last reviewed the Between October 2013 and February 2014, thirteen wetlands overall status and distribution of this flufftail in southern Africa, i.e. in were surveyed for White-winged Flufftail and other waterbirds by the time frame 2005 to the summer of 2014-2015. the first author (Appendix 1). Nine of the surveyed wetlands were sites where the flufftail had been previously recorded (Taylor 1994, METHODS 2000 b, 2005 d). Two Carex-dominated wetlands (Vrederus and Rush Valley) and one mid-altitude floodplain (Minnehaha) were selected for Sightings of White-winged Flufftails over the last decade were solicited investigation in the Eastern Cape, because of the historical occurrence and collated from bird-watchers and other interested parties. The of White-winged Flufftail in the Eastern Cape (Keith et al. 1970; Taylor authenticity of the sightings reported herein is accepted because all 1994, 2000 b, 2005 d). The remaining site (Verloren Vallei Nature observers referred to a smallish crake flushed from wetland vegetation Reserve) was chosen because of the presence of lightly modified that displayed prominent white patches in the wings (secondaries). wetlands and the hitherto unpublished reports of the flufftail from the Possible confusion with other small wetland birds with white or pale reserve (see under Results and Discussion). For logistical reasons, the wing patches, such as Little Grebe Tachybaptus ruficollis, were ruled out following South African wetlands, where White-winged Flufftails have after discussing the observations with each observer. The possibility been reported since the 1980s (Taylor 1994, 2000 b, 2005 d), were not that some of the records mentioned below might be misidentifications, surveyed: (1) Coleford Nature Reserve, KwaZulu-Natal, (2) Mfabeni, however, remains, as photographic evidence was typically not available. iSimangaliso Wetland Park, KwaZulu-Natal and (3) Murphy’s Rust, Free State.
Table 1. Summary of results for waterbird community richness and diversity.
WETLAND NUMBER OF NUMBER OF WATERBIRDS/ WATERBIRD WATERBIRDS MINUTE SPECIES
Vrederus (Nov) 29 749 1.4 Vrederus (Jan) 21 519 2.7 Rush Valley (Nov) 9 20 0.1 Rush Valley (Jan) 11 25 0.1 Minnehaha (Nov) 9 134 1.0 Minnehaha (Jan) 12 94 0.7 Middelpunt (Oct) 9 50 - Middelpunt (Dec) 12 49 0.2 Vogelvlei-Tramorne 29 400 1.1 Penny Park 25 209 0.8 Hebron 12 46 0.2 Wakkerstroom 22 165 0.5 Vanger 7 32 0.2 Seekoeivlei 39 775 5.0 Bedford-Chatsworth 15 43 0.2 Suikerbosrand 8 58 1.3 Verloren Vallei 5 5 0.1 64 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37
2014, and 11 and 27 February 2015. Detailed notes were not kept of these flushing efforts, but they generally consisted of approximately five beaters walking abreast, beating the surface of the vegetation with light sticks and calling out. Rope-dragging was not usually employed in these walks except for the flushing efforts in February 2014 and 2015. The area traversed in these ‘flushes’ fell between Trout Dam #3 and the northern fence line. These flushing efforts were performed in the morning (ca 08h00-11h00). Search effort was not standardised as the object was to confirm the presence of the flufftails in the wetland and, on certain occasions, to allow groups of bird-watchers, congregated on the sides of the wetland, to see the flufftails. From 13 to 14 February 2014 and 26 to 27 February 2015, approximately 250-300 m of mist-nets were deployed in the eastern part of the wetland and a beating party of approximately 10 observers dragged a very long (> 50 m) rope, with soft-drink cans and plastic bottles containing small stones attached, through this area in an attempt to flush and catch White-winged Flufftails for blood and feather samples (for genetic and isotopic studies). These capture exercises were performed under permit (MPB 5384 (2014) and MPB 5511 (2015)) from Mpumalanga
Fig. 2. Dominant botanical species flowering in sedge meadows at Middelpunt Wetland where White-winged Flufftails were flushed in December 2013 and February 2014: (1) Pycreus macranthus, (2) Fuirena ciliaris, (3) Pennisetum thunbergii and (4) Leersia hexandra. Photos: G. Davies.
The survey technique consisted either of the first author walking steadily through the wetland from dawn to around 11 am and again in the afternoon from around 15h00 to dusk, recording all waterbirds seen or heard, or rope-dragging by two to three people with a thin, 10-20 m long nylon rope (no cans or bottles attached to the rope). Particular attention was paid to listening for the low-pitched, short series of oop notes, which has been reported as the main call of the flufftail (Taylor 1994, 2005 d, Taylor & Van Perlo 1998). Additionally, the first author camped in a tent adjacent to six of the wetlands (Bedford-Chatsworth, Hebron, Middelpunt, Penny Park, Rush Valley, Seekoeivlei and Vogelvlei) to listen for any possible nocturnal Fig. 4. View of sedge meadow habitat on east side of Middelpunt Wetland, Mpumalanga, photographed December 2013, showing flowering Pycreus and vocalisations or other potential activity by the flufftails. At Middelpunt Fuirena sedges and Pennisetum grass. During December 2013 and February 2014, Wetland, during December 2013, the first author also undertook White-winged Flufftails were flushed from microhabitat of this type. Photo: G. gross plotting of the broad vegetation communities between Trout Davies. Dam #3 (25°33’07.88”S; 30°06’42.39”E) and the northern fence line (25°32’03.75”S; 30°07’24.52”E) by walking in the wetland and Tourism and Parks Agency, the regulating conservation authority. demarcating the boundaries of subjectively-assessed vegetation Ageing of caught White-winged Flufftails followed the terminology communities (e.g. Phragmites reed-beds, Typha rush-beds, monospecific and criteria of Taylor & Van Perlo (1998) and Taylor (2005 d) . Carex sedge areas) with a hand-held GPS. Excluding the down-covered chicks, three age classes have been The Middelpunt Wetland Trust organised ‘flushes’ at Middelpunt recognised in White-winged Flufftail: juvenile, immature and adult Wetland on 26 February and 19 March 2005, 4 and 7 January 2006, (Taylor 2005 d). The timing of the moults between these different age February 2011 (specific date not recorded), 13-14 and 22 February classes does not appear to be known in the White-winged Flufftail. In other Sarothrura species partial post-juvenile (first pre-basic) moult (excluding remiges and rectrices) to assume immature plumage occurs between 3-10 weeks in Buff-spotted Flufftail S. elegans, 5/6 to 10/11 weeks in Red-chested Flufftail S. rufa and from 5 weeks in Streaky-breasted Flufftail S. boehmi (Taylor 2005 a-c). Very little appears to be known about the timing of first pre-breeding (pre- alternate) moult to assume adult plumage, but in Streaky-breasted Flufftail it occurs six to seven months after the completion of the post-juvenile moult (Taylor 2005 c), therefore ca 8-10 months after hatching; full adult plumage being attained at about 11-12 months old. The post-breeding moult in adult flufftails is complete with erratic, synchronous or descendant moult of the primaries (Taylor Fig. 3. Cattle grazing in sedge meadow on east side of Middelpunt Wetland, 2005 a-c). Juvenile White-winged Flufftails are identified by their Mpumalanga on 14 December 2013, approximately 50 m from area where “predominantly grey-brown” (♂) or “blackish brown” (♀) heads with White-winged Flufftails were flushed. Photo: G. Davies. no rufous or chestnut wash to the breast (Taylor 2005 d). Immatures Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 65
Table 2. Details of oop vocalisations corresponding to description of White-winged Flufftail call (Taylor 1994, 2005 d) heard by the first author during the 2013-14 wetland survey.
WETLAND DATE AND CIRCUMSTANCES CONCLUSION TIME Vogelvlei-Tramorne, 22 January 2014, Oop calls were heard from three different locations in tall Grey Crowned Crane Franklin 04h30-05h07 Phragmites/Typha vegetation. Mahem calls were heard from the unison calls same direction as two of the calling oop areas and from the vicinity of the third area a Grey Crowned Crane voluntarily flew out of the reedbed at 05h10. Vogelvlei-Tramorne, 23 January 2014, Oop calls were heard again from three different areas in tall Grey Crowned Crane Franklin 04h30-05h35 Phragmites/Typha, probably referring to the same birds as above. unison calls Mahem calls were heard from the general direction of these oop noises, although there was no overlapping in the two different sounds. Uncertain Penny Park 23 January 2014, Oop calls were heard far out in dense, tall Typha bulrushes. The 19h50 sounds were difficult to pinpoint and were only briefly uttered. Uncertain Penny Park 24 January 2014, Single bouts of oop calls were heard ca 50-200 m north-west of 04h50-05h00 the bird hide in thick Typha bulrushes and also to the north of the small open water area but difficult to pinpoint. Grey Crowned Crane Penny Park 25 January 2014, Single bout of oop calls heard from tall Typha bulrushes on north- unison calls 08h07 west side of Typha area. Mahem calls were heard immediately afterwards and 2-3 Grey Crowned Cranes were also flying over the Typha area at this point, possibly stimulating the bird in the Typha to call. Uncertain Hebron 27 January 2014, Brief oop calls were heard from a Typha area, but the spot was 04h42 difficult to pinpoint in the heavy dawn mist. Grey Crowned Cranes were later seen in this Typha area. Grey Crowned Crane Seekoeivlei 24 February 2014, Two Grey Crowned Cranes were seen standing next to a small pan unison calls ca 07h00 immediately opposite the Waterval Section entrance. One crane was seen to point its bill vertically, while holding its neck in a ‘S’ shape, and give brief, deep oop calls. The crane was watched doing this with 10x40 Swarovski binoculars at a distance of ca 150 m. Grey Crowned Crane Bedford-Chatsworth 26 February 2014, Oop calls were heard emanating from thick reeds and sedges unison calls 04h50-05h05 just beyond the westernmost (lower) fence-line. A pair of Grey Crowned Cranes were shortly thereafter seen voluntarily to fly out from the same spot.
are intermediate with some development of chestnut or rufous on the billed Duck Anas undulata, South African Shelduck Tadorna cana and head and breast (Taylor 2005 d). Unlike most Sarothrura species, sexual African Snipe Gallinago nigripennis (Appendix 2). The most-regularly dimorphism in adult White-winged Flufftails is not pronounced, with encountered rallids, aside from Red-knobbed Coot, were African Rail both sexes having chestnut or rufous toned heads, necks and sides to Rallus carulescens and Common Moorhen Gallinula chloropus the breast (Taylor & Van Perlo 1998, Taylor 2005 d). (Appendix 2). Although traditionally the genus Sarothrura has been placed in the Of the Sarothruridae, Red-chested Flufftails were found at Rallidae (e.g. Keith et al. 1970; Taylor & Van Perlo 1998), recent genetic six wetlands (Appendix 2). At Vogelvlei-Tramorne, eight calling research (Hackett et al. 2008) suggests that the flufftails should be birds (either low, evenly-spaced hoots or excited dueh-dueh-dueh reclassified as a separate family (Sarothruridae). This latter view has vocalisations) were located during the transects in 1.5 to 2 m tall been followed here. Typha with dense 1 m high Carex and Cyperus sedges and 2 to 3 m tall Phragmites with dense, mixed sedges about 1 m high; water depth RESULTS was < 15 cm in both areas. At Penny Park, four were heard during transects and two outside of transect time, mostly in 2 m tall Typha Wetland survey of 2013-14 areas with knee- to waist-high, thick sedges and hygrophilous grasses Fifty-seven species of waterbirds were recorded at the 13 wetlands on lightly-flooded ground (< 20 cm deep). At Middelpunt, only one surveyed, including ten species of Rallidae s.l. (Appendix 2). The Red-chested Flufftail was recorded during transects, but several (ca range of waterbird species recorded per wetland varied between 5-10) were heard calling (low, evenly-spaced hoots and excited dueh- five and 39 species (average = 16.1, median = 12 species) and counts dueh-dueh calls) outside of transect time in the Typha and Phragmites varied between five and 775 waterbirds (average = 198.4, median = beds in the central part of the wetland. Red-chested Flufftails did not 58 birds; Table 1). The Waterval Section of Seekoeivlei was the most occur in the low sedge meadows of the wetland in December 2013, diverse wetland surveyed, both in terms of species richness and total although one female was flushed in such habitat in February 2014 (F. waterbirds counted (Table 1). Peacock pers. comm.). The five most regularly-encountered waterbirds were Western White-winged Flufftail was only detected at Middelpunt Wetland Cattle Egret Bulbulcus ibis, Red-knobbed Coot Fulica cristata, Yellow- during these field surveys. On 14 December 2013, a White-winged 66 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37
Fig. 5. Deployment of subjectively assessed palustrine vegetation communities at Middelpunt Wetland, Mpumalanga in December 2013. Light grey areas = sedge meadows dominated by flowering Pycreus macranthus, Fuirena ciliaris and Kyllinga erecta sedges (Cyperaceae) and Pennisetum thunbergii and Leersia hexandra hygrophilous grasses (Poaceae), 1 = mixed area of Pycreus macranthus, Carex sp., Schoenoplectus corymbosus and Cyperus cf. fastigiatus (Cyperaceae), 2 = Pycreus macranthus (Cyperaceae) sedges but strongly intermixed with Dierama pauciflorum (Iridaceae: Ixioideae) and Disa chrysostachya (Orchidaceae: Orchidoideae), 3 = Pycreus macranthus (Cyperaceae) sedges intermixed with Kyllinga erecta (Cyperaceae) sedges and Albuca sp. (Asparagaceae: Scilloideae) forbs, 4 = mixed area of Pycreus, Kyllinga and Carex sedges (Cyperaceae) with Dierama pauciflorum (Iridaceae) and Pennisetum thunbergii (Poaceae), 5 = monospecific patch of Carex cf. cognata (Cyperaceae), 6 = patch dominated by Gunnera perpensa (Gunneraceae) forb, X = camp spot of first author, black filled circles = areas where White-winged Flufftails were flushed in December 2013 and February 2014.
Flufftail flushed from 55 cm high vegetation at ca 05h10, and flew m away from the flufftail flushing site on 14 December (Fig. 3). The about 5-10 m before dropping down into the sedges; it was not immediate vicinities of the flush sites, however, were ungrazed and flushed again on walking to the landing spot. The immediate area untrampled (Figs. 1 and 4). where the bird flew out was dominated by two sedge (Cyperaceae) There were very few other waterbirds in the immediate area species, Pycreus macranthus and Fuirena ciliaris, intermixed with two where the White-winged Flufftails occurred at Middelpunt in hygrophilous grass (Poaceae) species, Pennisetum thunbergii and Leersia December 2013. Two Baillon’s Crakes Porzana pusilla were heard hexandra (Figs 1-2). Average water depth in the immediate vicinity vocalising (low, ratcheting rattles) very close (ca 15-20 m) to where of the flush site was 6.8 cm (range 5-8 cm, n=4). Vegetation cover White-winged Flufftails flushed and one crake was caught in a mist- was 90-100%, i.e. dense without exposed areas of mud or water. The net deployed about 30-50 m away. following day, 15 December 2013, a single White-winged Flufftail was Deep oop vocalisations, broadly corresponding with the flushed close to the sighting of the previous day at ca 05h50 from 60 description of the flufftails’ call, were heard at five wetlands, but in cm high P. macranthus sedges with water depth approximately 5-10 cm most instances the circumstances indicated that the calls originated and cover 90-100 %. The flufftail flew ca 20 m and, on walking to the from Grey Crowned Cranes Balearica regulorum and not the flufftails landing spot, it flushed again, this time flying much further (ca 50-60 (Table 2). Although the first author camped for three nights on a m). Approximately 75 cows were seen grazing in the sedge meadows grassy knoll adjacent to Middelpunt and within 100-400 m of the in one of the eastern side arms to the wetland on 13 December sites where the flufftails were flushed in December 2013 and later and 23 cows were grazing in the sedge meadows approximately 50 in February 2014, no oop vocalisations were heard nor any other Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 67
vocalisations that could be associated with the flufftails. first visited Middelpunt that season, regrowth was minimal, the The vegetation community plotting at Middelpunt indicated vegetation comprising short, regenerating reed, sedge, and grass tillers that the wetland was dominated by knee-high mixed sedges and (most < 20 cm tall) amidst blackened stubble and open patches of hygrophilous grasses (Fig. 5), viz. P. micranthus, F. ciliaris, Kyllinga erecta oozing red and black mud. There were a few, small (< 0.2 ha), narrow, (Cyperaceae) and P. thunbergii and L. hexandra (Poaceae). There were dense, unburnt clumps of 60-90 high sedges and grasses in shallowly- localised patches of flowering forbs such as red torch orchid Disa flooded (1-3 cm deep) patches, often with tiny pools (1-10 cm deep). chrysostachya (Orchidaceae) and marsh hairbell Dierama pauciflorum The substrate varied from damp to sodden, but water depth did (Iridaceae). The mixed sedge meadows covered approximately 125- not exceed 1 cm in most of the wetland. There was extremely little 150 ha. There was very little Carex sedge evident in December 2013, suitable habitat for flufftails (estimated < 1-2 ha) and the only rallid and only one very small (ca 0.2 ha) monospecific patch of Carex cf. recorded in the wetland in mid-October were two African Rails. cognata (Cyperaceae). The middle and lower sections of the wetland were dominated by tall Phragmites reedbeds and Typha rush-beds (Fig. White-winged Flufftail sightings between 2005 and 2015 5), covering approximately 30-40 ha. The majority of the wetland (ca Twenty-four sighting episodes of White-winged Flufftail comprising 95%) had been burnt in September 2013 (K. Combrink, Elandskloof 32-34 individuals were made between 2005 and 2015 at eleven Farm, pers. comm.). By mid-October 2013, when the senior author localities in South Africa, including the above-mentioned Middelpunt
Fig. 6. Map showing recent (post-2005) records of White-winged Flufftail in South Africa (dots) as well as selected localities of historical significance (asterisks). Note the concentration of recent records along the eastern escarpment. To avoid clutter, not all historical confirmed localities are displayed, e.g. Hebron and Penny Park, East Griqualand. 1 = Makalali Game Lodge, Gravelotte, 2 = Verloren Vallei Nature Reserve, 3 = Middelpunt Wetland, 4 = Wakkerstroom Wetland, 5 = Seekoeivlei Nature Reserve, Memel, 6 = Bedford-Chatsworth, Ingula, 7 = Klerksvley, QwaQwa Nature Reserve, 8 = King Shaka International Airport, Durban, 9 = Ntsikeni Nature Reserve, Swartberg, 10 = 40 km SW of Underberg, and 11 = 15 km N of Kokstad. 68 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37
throat, hind-neck and breast, blackish crown, lores and cheeks and the absence of white spotting on the median and lesser upperwing coverts. It weighed 26.6 g, with wing 74 mm, tail 36 mm, culmen 13.6 mm, tarsus 22 mm. The primaries were fresh and the flufftail showed no signs of active moult. The mist-net had been deployed in an area of low sedges with water level about 15 cm deep and approximately 30 m from 3 m tall Phragmites reedbeds. The mist-nets had been erected the previous night and unfurled very early (before dawn). The male was released unharmed after ringing. On 13 February 2014, a single White-winged Flufftail was flushed twice at Middelpunt Wetland from ca 80 cm high Leersia grass and non-flowering sedges with water depth approximately 15-20 cm and cover about 80-100%. When first flushed, the flufftail flew for ca 50 m, but on the second flushing it flew a shorter distance, ca 15-20 m, landing close to the line of mist-nets. After diving down into the sedges and hygrophilous grasses on the second occasion, the landing spot was surrounded by the beaters, who walked forward disturbing the flufftail, which ran out into the narrow pathway in front of the mist-nets before fluttering into the lowest shelf of mist- nets where it became entangled (Figs 8-9). The captured flufftail’s head was dark brown with chestnut speckling around the eye. The nape was dark brown with chestnut barring while the sides of the neck were dark brown with a slight chestnut wash. The sides of the Fig. 7. White-winged Flufftail caught in mist-net at Wakkerstroom Wetland, Mpumalanga, 18 January 2014, showing bright rufous face, hindneck and breast, breast were largely dark brown. The throat and middle of the breast unspotted median and lesser upperwing coverts and fresh (unworn) remiges. were white. The flanks were blackish brown with small white spots. Photo: S. van Stuyvenberg. The upperparts were blackish with white speckling. The primaries were black except the 1st primary and inner web of the 2nd primary sightings (Appendix 3; Fig. 6). These sightings were concentrated along while the secondaries were white. The greater coverts were black the eastern escarpment of South Africa in high-altitude (> 1500 m with minor white barring, while the median and lesser coverts were asl) wetlands, the bulk (46%) originating from Middelpunt Wetland. blackish and largely unmarked, except for a few, scattered tiny white Six of the localities represent sites where the flufftail has never spots. The tail was bright rufous with narrow black bars. The maxilla been recorded before in the literature, viz., 15 km north of Kokstad, was blackish, the mandible pale horn-pink with an infuscated upper Klerksvley, King Shaka International Airport, 40 km south-west of margin. The eye was dark brown. These plumage features indicated Underberg, Makalali Game Lodge and Ntsikeni Nature Reserve. Few that this individual was a juvenile. The bird weighed 33.7 g with wing of these records were accompanied by detailed information relating 78 mm, culmen 12.3 mm, tail 39 mm and tarsus 20. 9 mm. The remiges to vegetation composition and cover, flooding, or other pertinent were fresh and the bird showed no signs of active moult. The flufftail habitat factors (e.g. time since last burning of the wetland). Those was released unharmed after ringing. sightings with such pertinent ecological information are described On 5 September 2013, a White-winged Flufftail was found next to below. a fence in flooded grass at King Shaka International Airport, Durban At Verloren Vallei Nature Reserve (Wanhoop 78 JT section), a single flufftail was flushed in mid-December 2011 from the edge of a head-water wetland dominated by grasses and Cyperus and Carex sedges, approximately 1 to 1.5 m high, near a small patch of Phragmites reeds and flew for about 20 m before landing in dense cover (F. Krige in litt., January 2015). A single White-winged Flufftail flushed at about 06h30 close to the edge of Ntsikeni wetland in late November 2014 from ca 50 cm high Carex sedges and about 2-3 m from a dense patch of 1 to 1.5 m high Typha bulrushes (J. Cilliers in litt., December 2014, R. Edwards in litt., December 2014). The bird did not flush again. On the Waterval Section of Seekoeivlei Nature Reserve, a single bird flushed in late February 2015 from ca 30 cm high Leersia hexandra grass, Cyperus sedges and Persicaria forbs growing next to a clump of 2.2 m high Typha bulrushes with water depth of approximately 3 cm (M. Pretorius). The flufftail flew for about 5 m landing in a 60 cm high clump of Juncus rushes with water depth of approximately 12 cm. The bird did not flush again.
Morphometrics, moult and other details of mist-netted and live-caught White-winged Flufftails On 18 January 2014, a White-winged Flufftail was serendipitously caught in a mistnet during general bird-ringing at the western edge Fig. 8. White-winged Flufftail caught in mist-net at Middelpunt Wetland, Mpumalanga, 13 February 2014 showing largely dark brown head, largely pale of Wakkerstroom Wetland, Mpumalanga (Fig. 7). The captured flufftail mandible, fresh (unworn) remiges with conspicuous white secondary panel and was an adult male as indicated by the bright chestnut wash to the 1st primary. Photo: F. Krige. Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 69
accumulated much moribund growth due to no burning in the winter/ spring of 2013. The high counts of Western Cattle Egret during the transects (Appendix 2) were deceptive in that they represented roosting birds at Vrederus Farm dam and flocks flying overhead to roost at Wakkerstroom and Seekoeivlei. Red-knobbed Coot numbers were also skewed by large, non-breeding flocks congregated on Vrederus Farm dam, while the count of South African Shelduck was biased by a big flock dabbling in a shallowly-flooded depression on the Seekoeivlei floodplain. Those three species, strictly-speaking, are not regular inhabitants of the palustrine vegetation in high-altitude wetlands. Ignoring them, shows that Yellow-billed Duck, African Snipe and Grey Crowned Crane were the most commonly encountered species (Appendix 2). This corresponds closely to the results of Allan et al.
Fig. 9. White-winged Flufftail caught in mist-net at Middelpunt Wetland, (2006) who found African Snipe, Yellow-billed Duck and Marsh Owl Mpumalanga, 13 February 2014 showing dark brown head with chestnut Asio capensis to be the most regular palustrine species at six high- speckling, dark brown nape with chestnut barring, dark brown sides of neck altitude South African wetlands. Relative to data accumulated for the with slight chestnut wash, white spotting to lower mantle, white chin and centre CWAC project (Taylor et al. 1999), counts of 74 and 63 African Snipes of breast. at Penny Park and Vogelvlei during the 2013-14 survey were high. by airport staff. The bird had apparently flown into the fence during There was good snipe habitat along the south-eastern fringe of the the previous night. It was briefly retained in an airport office where a main Typha area at Penny Park, where extensive trampling and grazing few, low-resolution photographs and video-footage (taken by cellular by cattle of the low Leersia-Paspalum-Juncus-Eleocharis fringe had telephone) were recorded after which the bird was released alive near created a mosaic of open mud, shallow pools and flattened/cropped the recovery point (Figs 10-11). The flufftail had a chestnut and black grasses and sedges. At Vogelvlei, the bulk of the snipe were counted mottled head (including the nape), black upperparts with copious in a small area (at about 30°16’25.9”S, 29°26’37.1”E) where intensive white speckling and barring, white chin and throat and the breast was trampling and grazing by cows had created similar conditions to those largely white with dark brown and chestnut mottled sides. The bill was at Penny Park. Grey Crowned Cranes were encountered at 10 of the largely dark with slightly pale bases to both the maxilla and mandible. 13 surveyed wetlands, with good numbers at the Minnehaha Wetland, The eyes were brownish. The tail was rufous with black bars. On two where flocks of hundreds have been seen in the recent past (P. Sefton photos and the video footage the white secondaries were discernible. pers. comm.). The moult status could not be ascertained from the photographs, although there appeared to be minor abrasion to the remiges and Recent sightings of White-winged Flufftail rectrices. The plumage features indicate that the bird was an immature. Between 2005 and 2015 the majority of sightings (92 %) of White- winged Flufftails were from high-altitude wetlands (above 1500 m DISCUSSION asl), although two were from low-altitude sites (below 500 m asl). Middelpunt Wetland contributed most of the records, underscoring Wetland survey results the importance of this site for the flufftail, although the number of The wetland surveys during 2013-14 were largely unsuccessful in sightings is probably skewed by the sustained search effort over the detecting White-winged Flufftails, aside from at Middelpunt Wetland. last decade relative to other wetlands. Despite the large amount of We can only speculate on the failure to find the species at the other searching at Middelpunt, the usual number of White-winged Flufftails wetlands. As flufftails are extremely skulking birds, it is possible that flushed per visit over the last decade has only been 1-2 birds. It (a) the first author consistently overlooked the flufftails, (b) the survey appears that in the early 1990s the flufftail was more common at techniques (walking and rope-dragging) were inadequate at disclosing Middelpunt when up to eight birds were sometimes flushed (Taylor the flufftails’ presence, (c) the number of flufftails have declined since 1994, 1997 b). This apparent change in status is perhaps linked to the surveys of the 1990s (Taylor 1994, 1997 a, b), or (d) that earlier fluctuations in the extent of Carex-dominated sedge vegetation. During population estimates at the nine wetlands were inflated. The poor the gross vegetation community plotting undertaken in December 2013-14 results are, however, consistent with the last rope-dragging 2013, it was found that Carex sedge patches were highly localised. It is survey conducted during 2001-04 (Allan et al. 2006), which found no possible that the amount of Carex sedges has declined at Middelpunt White-winged Flufftails during 27 hours of transects at six of the nine or that very little was flowering in December 2013 and hence its important wetlands. extent was underestimated. A more detailed, quantitative investigation The largest wetland visited during the survey, Seekoeivlei, was the of the wetland vegetation and successional dynamics at the wetland is most diverse in terms of species richness and waterbirds counted warranted, especially to verify the coverage of Carex-dominated sedge (Table 1; Appendix 2). The recent flushing of a single White-winged vegetation. Flufftail on the Waterval Section confirms an earlier aural report of the species from the wetland (Taylor 1997 b). Although comparatively Evidence provided by live-caught White-winged Flufftails large (ca 300 ha) when contrasted against other wetlands in this Of the three live-caught flufftails, the individual trapped on 13 study, the Waterval Section comprises only approximately 6% of that February 2014 at Middelpunt was possibly the most interesting. The enormous, high-altitude wetland, suggesting that the total Seekoeivlei largely dark brown head and sides to the breast indicated that the bird wetland must routinely support thousands of waterbirds. The numbers was young (Taylor & Van Perlo 1998; Taylor 2005 d). The slight chestnut encountered on the Waterval Section are also liable to be higher in speckling behind, above and below the eyes and faint chestnut years of better rainfall as during 2013-14 the floodplain was largely wash and speckling to the nape and lower neck suggest it may have dry (except for water in oxbows and a few depressions) and had been juvenile male (“head … dark grey-brown with dull chestnut mottling … hind neck, sides of neck and mantle grey-brown, washed 70 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37
dull chestnut”, Taylor 2005 d), although the eyes were dark brown cheeks. The male had fresh remiges suggesting that it had possibly whereas the juvenile male is reported to have grey eyes (Taylor 2005 completed its wing moult within the previous 1-2 months. There d). The Middelpunt bird did not appear to be a full immature because is virtually no data on the post-breeding moult of adult White- the chestnut mottling to the head was rather restricted in extent, winged Flufftails, although a female in north-eastern South Africa the breast did not show any significant development of a chestnut was reported in primary moult in February (Taylor 2005 d) and wash and the (lower) mandible was predominantly pale. The well- two individuals “appearing to have stunted wings” in early January grown primaries (falling within adult length range; Taylor 2005 d) and at Franklin Marsh (Mendelsohn et al. 1983) may have been adults its good flight ability (as observed on the first occasion it flushed) undergoing synchronous or partially synchronous remige moult. suggested the bird was older than about five to six weeks, seeing Buff-spotted and Red-chested Flufftails are able to first fly at 19 and Problems associated with vocalisations 42 days respectively (Taylor 1994: 25). The Middelpunt juvenile was Over the last decade, no records of White-winged Flufftail were made similar in appearance to the specimen collected at Franklin Marsh in on the basis of vocalisations, possibly because of the difficulties in October 1982 (Durban Natural Science Museum, Accession # 33901), distinguishing the oop call attributed to the flufftail (Taylor 1994, 2005 due to the largely dark brown head and pale mandible, although the d) and the roosting (unison) calls of Grey Crowned Cranes (Allan Middelpunt bird had more extensive chestnut speckling to the head 2005). Although oop calls were heard at several wetlands during the and was thus older. The Franklin specimen has been aged as a juvenile 2013-14 survey, all seemed to originate from Grey Crowned Cranes “at least 7 weeks [old]” (Taylor 1994: 25). A young bird hatched in (and one crane was watched giving the call). Despite the presence of Ethiopia would have been about 5.5 to 6 months old by mid-February, White-winged Flufftails at Middelpunt between December 2013 and given a breeding season of July to September in that country (Taylor February 2014, no vocalisations were heard from the sedge meadows et al. 2004). Such an Ethiopian-bred individual by mid-February would that could be linked with the flufftails, suggesting either that they have presumably completed the post-juvenile moult, an age when were silent or that any calls given were missed. The latter possibility a distinct chestnut wash to the breast and head should have been is considered unlikely given that the first author camped next to the becoming apparent, the (lower) mandible should have presumably wetland and that observers listening for any unusual or flufftail-like largely darkened and the remiges may have shown signs of abrasion noises were present in and adjacent to the wetland from dawn to (seeing the remiges are not replaced at the post-juvenile moult). The dusk in both December 2013 and February 2014. absence of these features suggest the Middelpunt bird was older than Remarkably, on the Ethiopian nesting grounds, no oop calls or 7 weeks but younger than about 5.5 to 6 months, i.e. approximately other advertising vocalisations attributable to the flufftails have been 2-5 months old. heard, despite considerable field work (e.g. Taylor et al. 2004; Allan et al. 2006). More research into the vocalisations of the White- winged Flufftail is urgently required, as detection on auditory cues would greatly aid future surveys of the species in South Africa and largely obviate the need to walk and rope-drag wetlands potentially supporting the birds, thereby reducing disturbance to the flufftails and their habitat. The possibility that earlier aural records of White- winged Flufftail are based on misidentifications of unison calls of Grey Crowned Cranes also needs to be ruled out by verifying that the flufftail does indeed make a deep oop vocalisation, an objective perhaps best achieved with captive individuals (seeing crakes and flufftails adjust well to appropriate aviary conditions; C. Wintle in litt. ; Taylor 1994) or by further painstaking observation on the proven nesting grounds in Ethiopia.
Occurrence of White-winged Flufftail at low-altitude sites Most of the existing literature has considered the White-winged Flufftail to be a species of high-altitude wetlands in South Africa (e.g. Fig. 10. White-winged Flufftail recovered alive at King Shaka International Wolff & Milstein 1976; Collar & Stuart 1985; Taylor 1994, 2000 b) . Airport, Durban, 5 September 2013 showing chestnut and black mottled head, The occurrence of White-winged Flufftail at the coastal site of King white speckled upperparts and largely dark bill with slightly pale bases to maxilla and mandible. Photo: M. van Rooyen. Shaka International Airport (early September) and low-altitude area of Makalali, Mpumalanga (early March), however, corresponds to a The individual at King Shaka Airport appeared older than the few enigmatic and poorly-documented historical records from low- Middelpunt bird because of the very extensive chestnut mottling to altitude sites in KwaZulu-Natal and the Eastern Cape, South Africa. the head, the largely dark mandible, slight chestnut tinge to the sides Most of these have been discussed by Taylor (1994, 2000 b, 2005 of the breast (discernible on one of the brief cellular telephone video d) and Taylor & Grundling (2003). Additional low-altitude records sequences) and possibly slightly abraded remiges and rectrices. This include sightings of single White-winged Flufftails flushed near the St individual had evidently completed the post-juvenile moult and was Lucia Crocodile Centre, iSimangaliso Wetland Park, KwaZulu-Natal possibly undergoing or about to undergo the first pre-breeding moult. (ca 30 m asl) on 2 April 2004 (Davis 2004) and at Mataffin Farm, If the evidence from Streaky-breasted Flufftail, which undergoes such Nelspruit, Mpumalanga Province (ca 600 m asl) in April 1999 (Hall a moult 6-7 months after completion of the post-juvenile moult 2006), although neither have been adjudicated by Rarities Committees (Taylor 2005 c), applies to White-winged Flufftail, this would suggest to our knowledge. The dates of these records (early spring and early the Durban Airport individual was approximately eight months old. autumn) are times of the year when we might anticipate the flufftails The Wakkerstroom male was fully adult male, as indicated by to be dispersing from or returning to high-altitude wetlands, which the all dark bill and plumage features, especially the extensive, rich may be inimical to White-winged Flufftails during winter due to the chestnut colouration to the head with blackish crown, lores and withering of the vegetation, desiccation of the substrate, extensive Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 71
that are hitherto unpublished or have tended to be overlooked in the literature. In Zimbabwe, a single bird was seen at Aisleby Farm, 10 km north of Bulawayo, in February 1988, a sighting which was accepted by the Zimbabwe Rarities Committee (Hustler & Irwin 1995). The flufftail was flushed from a slushy path overgrown with sedges on a dam wall (D. Ewbank in litt., June 2013). Also in Zimbabwe, an unconfirmed sighting of a single individual was made in a “swampy area” on Sans Souci Farm, Ruwa district, approximately 20 km east of Harare in February 1989 (J. Jones in litt., June 2013). In the 1980s (exact dates not recorded), single White-winged Flufftails were seen at Draaikraal Wetland, Houtenbek Farm, Mpumalanga, approximately 7 km west of Verloren Vallei Nature Reserve, and in an area of palustrine wetland along the western arm of Jericho Dam, Amsterdam district, Mpumalanga, (W.R. Tarboton in litt., July 2014). On 6 February 2001, a serval Felis serval was flushed from thick sedges and hygrophilous grasses at Palmietfontein Wetland (25°21’41.44”S; 30°08’28.77”E), Nederhorst, 6 km north of Dullstroom, Mpumalanga, the cat in turn then flushed a single White-winged Flufftail while running off; the flufftail flew for about 10 m before dropping back into thick cover (F. Krige in litt., December 2014 and January 2015). Several days later, on 9 February 2001, another White-winged Flufftail flushed from mixed Cyperus and Carex vegetation in a wetland in the Wanhoop section of Fig. 11. Rear view of White-winged Flufftail recovered alive at King Shaka International Airport, Durban, 5 September 2013, showing white barred and Verloren Vallei Nature Reserve (F. Krige in litt., December 2014 and speckled upperparts, rufous tail with black barring, white secondary patch on January 2015). partially folded wing and apparently slightly worn remiges and rectrices. Photo: M. van Rooyen. Future requirements and conclusions Although the 2013-14 and earlier 2001-04 surveys failed to produce burning, extreme coldness and reduced invertebrate biomass. This good data on White-winged Flufftails, future surveys of known and tentatively raises the possibility that White-winged Flufftails may be candidate wetlands should still be undertaken. There are several large ‘wintering’ in small numbers in low-altitude wetlands in eastern South (> 500 ha) wetlands that have not been surveyed for White-winged Africa, although against this scenario are late winter (August) and Flufftail, such as Hlatikhulu, Mvoti and Mgeni wetlands in KwaZulu- early spring (October) specimens from Suikerbosrand Nature Reserve, Natal that might hold the species. Likewise palustrine wetlands in the Heidelberg, Potchefstroom and Bloemfontein on the Highveld (Keith et Potchefstroom, Bloemfontein and King Williams Town areas might al. 1970; Benson & Irwin 1974; Wolff & Milstein 1976; Taylor 1994) and repay careful scrutiny, considering the existence of old specimens summer records from the coastal site of Mfabeni, iSimangaliso Wetland from those places (Keith et al. 1970; Taylor 1994). Wetlands outside Park in November and December 2002 (Taylor & Grundling 2003). of South Africa should also be investigated further given enigmatic King Shaka International Airport does not appear to offer any historic records from Zimbabwe and Zambia (Hopkinson & Masterson suitable habitat and is mostly surrounded by sugarcane fields and 1984; Taylor 1994) and most recently from the Kwando Wetland suburbia, although a Phragmites-dominated marsh (ca 20 ha) occurs system, Caprivi Strip, Namibia, in late September 2012 (C.J. Brown in at Mount Moreland, approximately 1.5 km to the south-west of the litt., December 2014). Additionally, different techniques such as using airport, and areas of mixed palustrine habitat occur along the lower pointer dogs (Hopkinson & Masterson 1984; Taylor 1985, 1987; Muller Mdhloti (ca 15 ha east of N2 highway) and lower Umhlanga rivers (ca 2006) or passive walk-in rallid traps (Schmitt 1973) should probably 30 ha east of N2 highway), approximately 3 and 7 km to the south- also be pursued to see if better results could be achieved with those east, respectively. Noteworthy rallids and other cryptic marshland methods. Unfortunately, surveying and censusing the flufftails in South birds (e.g. Striped Crake Aenigmatolimnas marginalis and African Africa based on aural cues (“the only practical way of locating and Grass-Owl) were detected during intermittent avifaunal study of the identifying grassland and wetland flufftails”, Taylor 1994: 92), which lower Umhlanga River in the early 1990s (Smith 1992; Taylor et al. would also limit disturbance to palustrine habitat, has not been of use 1994, Taylor 1997 b), but no White-winged Flufftails were located. The in the last two surveys (Allan et al. 2006) and this study). Our review wetland avifauna of the Mount Moreland Wetland and lower Mdhloti also underscores the need for all sightings of White-winged Flufftail River is unknown. in South (and southern) Africa, even those from ‘traditional’ sites, to The bush airstrip at Makalali falls within a low-altitude savanna be documented in as much detail as possible and for observers to (‘bushveld’) area, a habitat unsuitable for flufftails and with no large give more attention towards the age and sex of the flufftails seen, palustrine wetland nearby. There are no other White-winged Flufftail composition and physiognomy of the wetland vegetation, flooding records from a savanna area in South Africa, except for a poorly- depth, time since last burning and other pertinent details when documented record from the Rustenburg area in May 1955 (Wolff & sightings are made. With such limited information available on the Milstein 1976; Tarboton et al. 1987). species, every sighting assumes high importance, and it is evident that much valuable information from past sightings has been irretrievably Some noteworthy White-winged Flufftail sightings before lost because such information was not systematically recorded. It is 2005 further recommended that the wetlands where White-winged Flufftail Although this paper has concentrated on the time-frame of 2005 are occasionally seen in South Africa should be afforded a high level to the summer of 2014-2015, it is worthwhile briefly mentioning of protection by government and conservation organisations to limit several other sightings of White-winged Flufftail before this period habitat destruction and degradation. 72 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37
ACKNOWLEDGEMENTS MULLER, M. 2006. Striped Crake and other birds in the Maun area SCHMITT, M.B. 1973. How to catch rails and crakes. Safring News Field work in 2013-14 by G. Davies was financially supported by 2(2): 7-9. ESKOM via BirdLife South Africa. The group of bird ringers and SMITH, E. 1992. Unusual observations – Umhlanga River, October beaters that provided assistance at Middelpunt are warmly thanked 1992. Albatross 312: 9-10. for their help. For assistance in various ways, we also thank Hugh TARBOTON, W.R., KEMP, M.I. & KEMP, A.C. 1987. The birds of the Chittenden, Brian Colahan, Deidre van Straaten, David Ewbank, Jim Transvaal. Transvaal Museum: Pretoria. Feely, John Jones, Frans Krige, Fanie Laubscher, Faansie Peacock, TAYLOR, P.B. 1985. Field studies of the African Crake Crex egregia in Maryna Matthee, Warwick Tarboton, Phyll Sefton and Marius van Zambia and Kenya. Ostrich 56(1-3): 170-185. Rooyen. Renee Reddy (University of Witwatersrand) identified TAYLOR, P.B. 1987. A field study of the Spotted Crake Porzana our plant samples. We acknowledge the various landowners for porzana at Ndola, Zambia. Ostrich 58: 107-117. permission to operate on their properties, especially the owners TAYLOR, P.B. 1994. The biology, ecology and conservation of four of the Middelpunt property. The draft manuscript benefitted from flufftail species, Sarothrura (Aves, Rallidae). Unpublished PhD incisive comments by D.G. Allan and P.B. Taylor. thesis, University of Natal: Pietermaritzburg, South Africa. TAYLOR, P.B. 1997a. The status and conservation of rallids in South REFERENCES Africa: results of a wetland survey in 1995/96. Avian Demography Unit Research Report #23. Avian Demography Unit: Cape Town. ALLAN, D.G. 2005. Grey Crowned Crane. In: HOCKEY, P.A.R., DEAN, TAYLOR, P.B. 1997b. South African palustrine wetlands: the results of W.R.J. & RYAN, P.G. (eds), Roberts – birds of southern Africa, 7 th a survey in summer 1995/96. Avian Demography Unit Research edition, pp. 307-308. John Voelcker Bird Book Fund: Cape Town. Report #24. Avian Demography Unit: Cape Town. ALLAN, D.G., McINNES, A.M., & WONDAFRASH, M. 2006. White- TAYLOR, [P.]B. 1999. White-winged Flufftails in Ethiopia – another winged Flufftail Sarothrura ayresi in Ethiopia: notes on habitat, mystery solved. Africa Birds & Birding 4(5): 23-25. densities, morphometrics, nests and eggs, and associated TAYLOR, P.B. 2000b. Whitewinged Flufftail. In: BARNES, K.N. (ed.), The waterbirds. Bulletin of the African Bird Club 13(1): 28-36. ESKOM red data book of birds of South Africa, Lesotho and Swaziland, BARNES, K. (ed.). 1998. The Important Bird Areas of southern Africa. pp. 28-29. BirdLife South Africa: Johannesburg. BirdLife South Africa & Avian Demography Unit: Johannesburg & TAYLOR, P.B. 2005a. Buff-spotted Flufftail. In: HOCKEY, P.A.R., DEAN, Cape Town. W.R.J. & RYAN, P.G. (eds), Roberts – birds of Southern Africa, 7 th BENSON, C.W. & IRWIN, M.P.S. 1974. On a specimen of Sarothrura edition, pp. 315-316. John Voelcker Bird Book Fund: Cape Town. ayresi from the Transvaal in the Leiden Museum. Ostrich 45(3): 193- TAYLOR, P.B. 2005 b. Red-chested Flufftail. In: HOCKEY, P.A.R., DEAN, 194. W.R.J. & RYAN, P.G. (eds), Roberts – birds of Southern Africa, 7 th BIRDLIFE INTERNATIONAL. 2015. Species fact sheet: Sarothrura edition, pp. 317-318. John Voelcker Bird Book Fund: Cape Town. ayresi. Downloaded from http:// www.birdlife.org. TAYLOR, P.B. 2005 c. Streaky-breasted Flufftail. In: HOCKEY, P.A.R., COLLAR, N.J. & STUART, S.N. 1985. Threatened birds of Africa and DEAN, W.R.J. & RYAN, P.G. (eds), Roberts – birds of Southern Africa, related islands. ICBP: Cambridge, U.K. 7th edition, pp. 318-319. John Voelcker Bird Book Fund: Cape Town. DAVIS, S. 2004. KwaZulu-Natal Honorary Recorder’s report – July TAYLOR, P.B. 2005 d. White-winged Flufftail. In: HOCKEY, P.A.R., 2004. KZN Birds 9: 28-30. DEAN, W.R.J. & RYAN, P.G. (eds), Roberts – birds of Southern Africa, HACKETT, S.J., KIMBALL, R.T., REDDY, S., BOWIE, R.C.K., BRAUN, 7th edition, pp. 321-322. John Voelcker Bird Book Fund: Cape Town. E.L., BRAUN, M.J., CHOJNOWSKI, J.L., COX, W.A., HAN, K-L., TAYLOR, [P.]B. & VAN PERLO, B. 1998. Rails: A guide to the rails, crakes, HARSHMAN, J., HUDDLESTON, C.J., MARKS, B.D., MIGLIA, K.J., gallinules and coots of the world. Pica Press: Robertsbridge, United MOORE, W.S., SHELDON, F.H., STEADMAN, D.W., WITT, C.C., Kingdom. & YURI, T. 2008. A phylogenomic study of birds reveals their TAYLOR, P.B. & GRUNDLING, P-L. 2003. The importance of South evolutionary history. Science 320: 1763-1768. African mires as habitat for the endangered Whitewinged Flufftail HALL, D. 2006. White-winged Flufftail. The Hornbill 83/84: 42. (Sarothrura ayresi). International Mire Conservation Group Newsletter HOPKINSON, G. & MASTERSON, A. 1984. The occurrence and 2003(2): 8-12. ecological preferences of certain Rallidae near Salisbury. In: TAYLOR, [P.]B., SMITH, E., & HERHOLDT, K. 1994. The Striped Crake LEDGER, J. (ed.), Proceedings of the Fifth Pan-African Ornithological in southern Africa. Birding in Southern Africa 46: 18-21. Congress, pp. 425-440. South African Ornithological Society: TAYLOR, P.B., NAVARRO, R.A., WREN-SARGENT, M., HARRISON, Johannesburg. J.A. & KIESWETTER, S.L. 1999. TOTAL CWAC report – coordinated HUSTLER, K. & IRWIN, M.P.S. 1995. Fifth report of the the OAZ waterbird counts in South Africa, 1992-1997. Avian Demography Rarities Committee. Honeyguide 41(2): 103-106. Unit: Cape Town. KEITH, S., BENSON, C.W. & IRWIN, M.P.S. 1970. The genus Sarothrura TAYLOR, [P.]B., WONDAFRASH, M. & DEMEKE, Y. 2004. The nest, eggs (Aves, Rallidae). Bulletin of the American Museum of Natural History and chicks of the White-winged Flufftail Sarothrura ayresi. Bulletin 143: 1-84. of the British Ornithologists’ Club 124(4): 233-239. MENDELSOHN, J.M., SINCLAIR, J.C. & TARBOTON, W.R. 1983. WOLFF, S. & MILSTEIN, P. le S. 1976. Rediscovery of the White-winged Flushing flufftails out of vleis. Bokmakierie 35(1): 9-11. Flufftail in South Africa. Bokmakierie 28(2): 33-36. Babbler 48: 55-58. Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 73 CO- ORDINATES 25°32’34.77”S 30°07’21.71”E 27°21’25.55”S 30°07’33.17”E 25°17’13.85”S 30°09’01.14”E 27°51’54.28”S 29°40’59.87”E 27°37’20.0”S 29°35’02.96”E 28°14’03.60”S 29°32’17.37”E 30°47’20.55”S 28° 16’19.63”E 30° 50’56.57”S 28° 12’53.75”E 30° 59’32.97”S 28° 14’02.96”E 30° 16’37.24”S 29°26’11.74”E 30°30’16.45”S 29°29’07.17”E 30°24’27.26”S 29°33’35.64”E 26°32’42.03”S 28°14’51.25”E TIME ca 9 hr 4 hr 46 min 5 hr 14 min 1 hr 15 min 3 hr 04 min 2 hr 35 min 4 hr 30 min 8 hr 52 min 3 hr 12 min 3 hr 20 min 3 hr 01 min 2 hr 08 min 2 hr 22 min 5 hr 50 min 4 hr 20 min 3 hr 21 min 0 hr 45 min TRANSECT DATES SURVEYED 16-18/10/2013 13-15/12/2013 20-21/2/2014 12/2/2014 22/2/2014 23-24/2/2014 24-26/2/2014 25-27/11/2013 30-31/1/2014 27-28/11/2013 28-30/1/2014 28/11/2013 27-28/1/2014 21-23/1/2014 23-25/1/2014 25-27/1/2014 11/2/2014
and and Typha 20 ha) ca 20 ha) and and Typha 300 ha) mostly ca 300 ha) mostly seen. Apparently Apparently Carex seen. on mud pedestals surrounded pedestals surrounded on mud Carex observed. Wetland with with Wetland observed. or Phragmites Typha APPENDIX 1.
sedges (unburnt in preceding winter with much moribund moribund winter with much sedges (unburnt in preceding Carex sedgebeds. No Typha sedgebeds. monospecific Carex by covered mostly ca 10 ha), 500 m length along upper, western edge of wetland and heavy grazing grazing and heavy edge of wetland western ca 500 m length along upper, DESCRIPTION observed. Wetland modified by channel incision with channel incision with modified by Wetland observed. or Carex Phragmites Typha, Carex Carex and no flowering present little Phragmites Very pools. large oxbow ca seven reeds. Apparently burnt in the preceding winter. winter. burnt in the preceding Apparently reeds. sedges and Phragmites Carex vegetation unburnt in preceding winter with dense moribund material. Wetland Wetland winter with dense moribund material. unburnt in preceding vegetation Carex Carex 400 ha) with diverse sedge and hygrophilous grass communities, including oxbow pools pools including oxbow grass communities, sedge and hygrophilous 400 ha) with diverse ca 0.5-1 m deep) on both eastern and western edges of wetland, partially filled in. Within IBA SA Within IBA SA partially filled in. edges of wetland, 0.5-1 m deep) on both eastern and western 85 ha), with small areas of mixed sedge and hygrophilous grasses. Little grasses. sedge and hygrophilous of mixed with small areas 85 ha), not burnt in preceding winter with much moribund material present. Two arms adjacent ( Two moribund material present. winter with much not burnt in preceding observed. Most of wetland apparently burnt in preceding winter. Wetland minimally disturbed, some grazing some grazing disturbed, minimally Wetland winter. burnt in preceding apparently Most of wetland observed. observed. No burning for several years and much of the wetland thick with moribund vegetation. Wetland fairly fairly Wetland thick with moribund vegetation. of the wetland and much years several No burning for observed. and mixed sedges/hygrophilous grasses. A small pan and several large pools fringed with emergent sedges, reeds and and reeds large pools fringed with emergent sedges, A small pan and several grasses. sedges/hygrophilous and mixed . Burnt in September 2013. Wetland fairly heavily modified by dam construction with heavy grazing by livestock in adjacent in adjacent livestock grazing by dam construction with heavy modified by heavily fairly Wetland Burnt in September 2013. . 180 ha), high-altitude, head-water wetland with shallowly-flooded sedge meadows, and extensive expanses of and extensive sedge meadows, with shallowly-flooded head-water wetland high-altitude, 180 ha), 40 ha) head-water wetland, large areas dominated by dominated by large areas 40 ha) head-water wetland, 50-100 ha with extensive ca 50-100 ha with extensive section covering Vogelvlei-Tramorne ( ca 600 ha), high-altitude wetland -dominated, Phragmites 30 and 50 ha). ( ca 30 and 50 ha). areas sedge meadow extensive but with two ( ca 500 ha), high-altitude wetland -dominated, Phragmites present. Carex present. Phragmites or . Many oxbow lakes with thick fringing reeds/ lakes oxbow Many . and Phragmites Cyperaceae with patches of Typha grass and mixed Leersia hexandra but not The floodplain was damp underfoot observed. No Carex . and submerged Lagarosiphon sedges and floating Potamogeton . IBA SA 022 Suikerbosrand Nature Reserve. Nature IBA SA 022 Suikerbosrand . Phragmites Phragmites Large ( ca flooded (following poor summer rainfall). The floodplain vegetation had not been burnt in the preceding winter and there was much was much winter and there had not been burnt in the preceding The floodplain vegetation poor summer rainfall). flooded (following Within IBA Reserve. SA 020 Grassland Biosphere with minimal disturbance. Wetland moribund material. ( high-altitude floodplain wetland Large, morphology with ‘hummock-ground’ but southern arm displays disturbed, minimally Currently IBA SA 043 Bedford/Chatsworth. past cattle trampling. from possibly deep channels, by sedge mixed , of Phragmites areas ( ca 70 ha) with large farm dam (16 and less extensive high-altitude wetland -dominated, Carex Most grass communities. and hygrophilous in adjacent paddocks. livestock grazing by dam construction with heavy modified by heavily fairly high-altitude pan ( permanent, (< 1.2 m deep), shallow Small, with minimal disturbance. near-pristine, Wetland grasses. sedges and hygrophilous low dominated by oxbow A few Cyperaceae. Leersia grass and mixed by mostly hexandra mid-altitude floodplain ( ca 100 ha) covered large, Fairly No emergent grasses and sedges. by pools covered morphology with grasses and sedges on pedestals ‘hummock-ground’ Parts of the floodplain also display grazing. livestock intensive cattle trampling. of excessive a result possibly of water, deep micro-channels by surrounded of mud Large, of Carex areas Leersia-Juncus-Paspalum floodplain ( ca 60 ha) comprising low by bordered ( ca 40 ha), -dominated wetland mid-altitude Typha Small, with grass-dominated area and drier, communities paddocks. Long (> 700 m) ditches ( ca paddocks. 016 Steenkampsberg. Large, Carex No flowering Small ( ca and easy to walk through). winter/spring, grasses (burnt in preceding sedges and hygrophilous with mixed other areas material), No visited). most of other wetlands common (unlike forbs hygrophilous Flowering Within IBA SA 016 Steenkampsberg. minimal disturbance and no grazing pressure. ( ca -dominated wetland Phragmites Within IBA SA 020 Grassland light grazing in adjacent paddocks. condition, in near-pristine Wetland winter. burnt in preceding Reserve.Biosphere section ( Waterval palustrine vegetation. (>5000 ha) with diverse floodplain wetland high-altitude oxbow Huge, horses in paddock Two but otherwise in excellent condition. canal, small concrete modified by lightly Wetland sedges observed. Park. IBA SA 083 Penny cattle grazing. heavy of recent but signs along edge of wetland during visit, adjacent to wetland No grasses. sedges/hygrophilous and mixed of Typha areas ( ca 50 ha) with additional -dominated head-water wetland Carex reedbeds Phragmites Vlei. Within IBA SA 081 Franklin and cultivation of adjoining paddocks. grasses and patches of sedges and hygrophilous (ca 40 ha) with mixed mid-altitude floodplain wetland small, Fairly grasses also present. Wetland fairly heavily modified by small impoundments with adjacent intensive livestock grazing and maize grazing and maize livestock small impoundments with adjacent intensive modified by heavily fairly Wetland grasses also present. Vlei. Within IBA SA 081 Franklin cultivation. on sedge meadow adjacent to town and localised channel incision in south-eastern sedge meadow (gabions emplaced to arrest (gabions emplaced to arrest and localised channel incision in south-eastern sedge meadow adjacent to town on sedge meadow Within IBAReserve. SA 020 Grassland Biosphere erosion). heavily modified by large dam above wetland, deep ditch of wetland, large dam above modified by heavily with fringing reeds and sedges, large expanses of and sedges, with fringing reeds WETLAND *Middelpunt, Belfast/Dullstroom, Mpumalanga *Wakkerstroom, Mpumalanga Vallei Verloren Dullstroom, NR, Mpumalanga State Free *Vanger, Seekoeivlei *Waterval, State Free NR, *Bedford-Chatsworth, State Free Ingula, Pitseng Pass, Vrederus, E Cape Eland Rush Valley, E Cape Heights, Maclear, Minnehaha, E Cape *Vogelvlei-Tramorne, KwaZulu- Franklin, Natal Kokstad, Park, *Penny KwaZulu-Natal KwaZulu- *Hebron, Natal *Elandslaagte, NR, Suikerbosrand Gauteng Summary of wetlands covered during the 2013-14 survey by the first author. An asterisk next to a wetland indicates that White-winged Flufftail had been recorded at the site before the 2013-14 survey. the 2013-14 survey. at the site before White-winged Flufftail had been recorded indicates that An asterisk next to a wetland the first author. by during the 2013-14 survey Summary covered of wetlands Reserve. NR = Nature (Barnes 1998); Area African Important Bird IBA SA = South 74 WHITE-WINGED FLUFFTAIL Durban Natural Science Museum Novitates 37 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 VER 02/14 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 6 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 21 19 SUI 02/14 0 0 0 0 0 0 0 0 0 0 4 0 0 0 6 0 0 0 2 0 2 0 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 1 3 3 2 0 2 0 0 1 0 0 10 BED 02/14 2 3 4 1 1 9 2 4 6 2 4 4 0 2 0 0 0 2 9 4 0 0 0 0 1 0 1 0 0 0 0 0 2 0 9 4 0 2 1 1 1 7 2 44 46 67 18 28 15 22 26 205 206 SEE 02/14 0 0 0 0 0 0 3 0 4 0 0 0 0 0 0 3 0 3 6 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 2 0 0 0 0 0 1 0 0 0 0 3 0 0 0 0 17 VAN 02/14 0 0 0 0 0 0 9 0 0 0 0 6 0 2 1 3 2 0 4 2 0 0 0 0 0 0 0 0 3 3 0 0 0 0 0 2 0 7 1 1 0 4 0 1 0 0 0 0 11 18 73 02/14 WAK 0 0 2 1 0 0 3 2 0 8 10 HEB 0 5 8 4 2 0 2 14 16 74 0 15 PEN 0 40 21 14 18 13 58 13 11 VOG VOG 0 0 1 1 0 6 1 5 0 0 16 86 30 12 13 MID 0 0 0 0 0 0 0 3 2 0 0 0 1 4 0 0 1 0 2 1 0 3 4 0 1 2 0 1 2 1 2 0 3 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 5 2 0 0 3 1 0 3 0 2 0 0 0 2 3 0 0 6 2 3 0 9 0 0 1 8 4 2 0 0 1 0 5 2 0 1 5 0 0 0 3 1 0 5 0 0 5 0 2 18 15 2 63 MID 10/13 12/13 01/14 0 0 0 0 0 0 0 0 4 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 3 1 0 0 0 0 2 0 0 0 0 8 0 0 0 1 0 1 0 0 0 0 0 0 0 0 36 19 13 MIN 11/13 APPENDIX 2. 0 0 0 0 9 3 4 0 1 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 7 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 48 45 19 MIN 11/13 0 0 0 3 1 0 2 0 0 6 0 0 RUS RUS 0 0 0 0 0 0 0 0 0 7 0 0 0 0 4 2 3 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 2 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 RUS 11/13 02/14 2 0 0 0 8 0 0 0 0 5 7 0 0 0 0 3 0 4 0 0 0 0 0 8 0 2 6 1 0 0 0 0 0 1 6 0 0 0 7 0 0 0 0 0 26 76 19 65 20 249 VRE 01/14 0 0 4 8 0 0 0 1 0 0 0 0 0 2 0 2 9 0 0 0 0 0 4 3 0 5 3 0 0 0 0 0 7 2 0 0 0 0 8 1 2 0 0 0 0 3 1 0 33 12 99 30 18 12 41 46 26 12 18 45 273 VRE 11/13 nycticorax Nycticorax Phalacrocorax lucidus Phalacrocorax Vanellus senegallus Vanellus Tadorna cana Tadorna Circus ranivorus ayresi Sarothrura Tringa nebularia Tringa Balearica regulorum Bulbulcus ibis Charadrius tricollarisCharadrius rufa Sarothrura Ardea melanocephala Ardea Alcedo cristata Thalassornis leuconotus Plectropterus gambensis Plectropterus Vanellus armatus Vanellus Fulica cristata Gallinula chloropus Anas undulata Tyto capensis Tyto Egretta intermedia Egretta Threskiornis aethiopicus Threskiornis Porphyrio madagascariensisPorphyrio Dendrocygna viduata Himantopus himantopus Haliaeetus vocifer Microcarbo africanus Microcarbo Platalea alba Chlidonias hybrida Alopochen aegyptiaca Anas hottentota Anas erythrorhyncha Motacilla capensis Scopus umbretta Philomachus pugnax ruficollis Tachybaptus Little Grebe African Darter Anhinga rufa White-faced Duck SPECIES Cormorant White-breasted Reed Cormorant Ibis African Sacred African Spoonbill ardesiaca Black-headed Heron Egretta Night Heron Black-crowned Heron Cattle Egret Western Egret Yellow-billed Black Duck Yellow-billed Goose Spur-winged Egyptian Goose South African Shelduck Duck White-backed African Fish Eagle African Marsh Harrier Crane Crowned Grey Common Moorhen Red-knobbed Coot African Swamphen Red-chested Flufftail White-winged Flufftail Common Greenshank African Wattled Lapwing Blacksmith Lapwing Plover Three-banded Black-winged Stilt African Grass-Owl Malachite Kingfisher Little Stint Calidris minuta Ruff Hottentot Teal Hottentot Teal Red-billed Teal Anas smithii Shoveler Cape carunculatus Crane Bugeranus Wattled African Rail Rallus caerulescens Marsh Owl Asio capensis Hamerkop Hamerkop Hadeda Ibis Bostrychia hagedash Glossy Ibis Plegadis falcinellus garzetta Egretta Little Egret maccoa Maccoa Duck Oxyura nigripennisAfrican Snipe Gallinago egregia Crex African Crake crex Crex Corn Crake pusilla Porzana Crake Baillon’s flavirostris Amaurornis Black Crake glareola Sandpiper Tringa Wood Whiskered Tern Tern Whiskered Wagtail Cape cinerea Ardea Heron Grey purpurea Ardea Purple Heron ralloides Ardeola Squacco Heron Little Bittern Ixobrychus minutus VOG = Vogelvlei-Tramorne, PEN = Penny Park, HEB = Hebron, WAK = Wakkerstroom, VAN = Vanger, SEE = Seekoeivlei, BED = Bedford-Chatsworth, SUI = Elandslaagte, Suikerbosrand, VER = Verloren Vallei. Verloren MID = Middelpunt, VER = MIN = Minnehaha, Pan and adjacent sedge meadows, Valley Suikerbosrand, = Rush SUI = Elandslaagte, RUS BED = Bedford-Chatsworth, Vrederus, SEE = Seekoeivlei, VRE = Vanger, Abbreviations: = VAN during the 2013-14 survey. wetlands at respective recorded Numbers of waterbirds Wakkerstroom, = WAK HEB = Hebron, Park, PEN = Penny Vogelvlei-Tramorne, = VOG Durban Natural Science Museum Novitates 37 WHITE-WINGED FLUFFTAIL 75 25 m - - - - - grass bridge REMARKS See main text See main text See main text See main text See main text See main text See main text See main text See main text Flushed in morning airstrip, flew ca flew airstrip, on east side of wetland. Flushed in lowest arm of vlei Flushed in lowest with water depth 4-8 cm deep was burnt in the winter of 2013 paddock (no marsh in immediate vicinity) paddock Flushed in late afternoon from low sedges low Flushed in late afternoon from Bird found dead close to powerline in grassy in grassy dead close to powerline found Bird Flushed from low sedges at dawn close to R543 close to R543 sedges at dawn low Flushed from Flushed from 30-40 cm high flooded grass next to 30-40 cm high flooded grass next to Flushed from Flushed late morning from sedges (cover 80-100%) 80-100%) sedges (cover Flushed late morning from Yellow-billed Duck flew up flushing the flufftail. Seen Seen up flushing the flufftail. Duck flew Yellow-billed Bird flew voluntarily for short distance from flooded flooded short for distance from voluntarily flew Bird Bird flew ca 80 m after flushing from low sedges; vlei vlei sedges; low ca 80 m after flushing from flew Bird J. Scott J. F. Krige F. Krige F. T. Wood T. M. Pretorius M. M. Pretorius M. D.H. Maphisa D.H. J. McCormick J. (M. Pretorius) (M. J. Cilliers et al. J. G. Davies et al. Davies G. D.W. Pietersen D.W. D. Coetzee et al . D. D. Coetzee et al. D. OBSERVER/S D. van Stuyvenberg D. S. Rautenbach et al. S. M. Drummond et al. M. Drummond et al. M. Drummond et al. M. M. van Rooyen et al. van Rooyen M. H. Smit-Robinson et al. H. Smit-Robinson et al. H. Smit-Robinson et al. H. H. Smit-Robinson et al. H. 1 1 2 1 1 1 1 3 1 1 1 1 1 1 1 1 1 3 1 1 2 1 3-4 1-2 NUMBER OF BIRDS “ ” “ ” “ ” “ ” “ ” “ ” DATE Feb 2011 Feb (m asl) APPENDIX 3. 4 Jan 2006 7 Jan 2006 7 Feb 2007 7 Feb 5 Sep 2013 8 Mar 2014 6 Dec 2012 18 Jan 2014 7 Nov 2013 7 Nov 26 Feb 2005 26 Feb 2014 13 Feb 2014 22 Feb 2015 11 Feb 2015 25 Feb 19 Mar 2005 28 Dec 2008 17 Dec 2011 21 Dec 2013 25 Dec 2013 30 Nov 2014 30 Nov not recorded Late Jan 2014 14-15 Dec 2013 Post-2005, exact date exact date Post-2005, 2008 (exact date lost)
1690 90 470 1515 1689 1695 2135 1690 2100 1880 1880 1740 1750 1675 1880 1885 1885 1885 1782 1887 1884 ca 1880 ca 1880 ca 1880 ca 1880 ca 1880 ca 1880 ca ALTITUDE
- Not recorded CO-ORDINATES 30°24’03.6”S, 29°27’13.3”E 30°24’03.6”S, 30°07’07.6”E 25°32’49.3”S, 30°07’09.9”E 25°32’38.9”S, 30°06’52.0”E 25°32’50.1”S, 30°06’59.3”E 25°32’44.1”S, 30°07’00.5”E 25°32’47.6”S, 30°07’02.3”E 25°32’51.6”S, 30°07’17.0”E 25°32’28.8”S, 29°30’58.7”E 28°14’25.0”S, 28°43’14.1”E 28°28’03.0”S, 30°06’58.2”E 25°32’44.7”S, 29°30’59.7”E 28°14’25.9”S, 31°06’32.2”E 29°36’40.0”S, 30°07’10.8”E 25°32’45.5”S, 30°07’10.8”E 25°32’45.5”S, 30°07’36.7”E 27°21’57.0”S, 29°15’57.5”E 30°05’41.2”S, 30°07’11.3”E 25°32’43.7”S, 30°07’23.3”E 25°32’38.9”S, 30°07’21.4”E 25°32’39.9”S, 30°07’20.1”E 25°32’40.4”S, 30°39’37.1”E 24°10’03.5”S, 30°07’25.9”E 25°32’36.6”S, 30°07’11.2”E 25°32’46.1”S, 27°38’3.66”S, 29°34’51.96”E 27°38’3.66”S, 25°17’14.86”S, 30°09’01.39”E 25°17’14.86”S, 30°07’13.50”E 27°21’25.11”S, 29°27’55.81”E 30°07’36.43”S,
Limpopo Free State Free State Free State Free State Free Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga Mpumalanga PROVINCE KwaZulu-Natal KwaZulu-Natal KwaZulu-Natal KwaZulu-Natal 15 km N of Kokstad Wetland Middelpunt Wetland Middelpunt Wetland Middelpunt Wetland Middelpunt Wetland Middelpunt Bedford-Chatsworth (Ingula) Wetland Qwaqwa Klerksvley, Reserve Nature Wetland Middelpunt Vallei Nature Verloren Reserve Beford-Chatsworth (Ingula) Wetland King Shaka International Airport Vallei Nature Verloren Reserve Wetland Middelpunt Wetland Middelpunt Wetland Wakkerstroom Wetland Wakkerstroom of 40 km south-west Underberg Wetland Middelpunt Wetland Middelpunt Makalali Game Lodge Wetland Ntsikeni Wetland Middelpunt Nature Seekoeivlei Reserve Sighting episodes of White-winged Flufftail in South Africa 2005 to summer of 2014-2015. White-winged Flufftail in South Sighting episodes of LOCALITY 1) 2) 3) 4) 5) 6) 7) 8) 9) 10) 11) 12) 13) 14) 15) 16) 17) 18) 19) 20) 21) 22) 23) 24)
DURBAN NATURAL SCIENCE MUSEUM NOVITATES
– INSTRUCTIONS TO AUTHORS
PUBLICATION POLICY: The Durban Natural Science Synonymies are cited as follows: Agulhasia davidsonii King 1871: 109, Museum Novitates is approved by the South African Department of pl. 11, figs 1-8; Dall 1873: 180; Thompson 1927a: 166, fig. 52; 1927b: 13, National Education for SAPSE funding. It publishes papers dealing with pl. 1, fig. 1, text-figs 1-3. Note that commas are not used to separate the natural sciences, with an emphasis on the geographical area of author and date, paginations are indicated by colons, lower case is used southeastern Africa. Authorship is open to persons not directly for plates and figures, plates/figures are abbreviated pls/figs (note no associated with the Museum. Each submission will be sent to at least full stop at the end) and a plate/figure is abbreviated pl./fig. (note the full two referees for review. Only original material that has not been stop). All references cited in synonymies should be included in the submitted for publication elsewhere may be submitted. Contributions reference list. should be written in English. The first reference to species should be followed by the author of the species, e.g. Lanius minor (Gmelin). The names of new taxa formally PRESENTATION OF MANUSCRIPTS: The electronic described in the paper should not be included in the title. submission of manuscripts by means of email to the Editor is expected, and this includes all text, tables and figures wherever possible, provide TABLES AND FIGURES: Tables must be numbered in the their e-mail addresses. Each full paper, but not short communications order that they are to appear and are mentioned in the text. Each table (<2 000 words including references but excluding words in figures, should be on a separate page with the table number and title at the top. tables and appendices), must start with a short summary (<200 words) Note the arrangement of horizontal lines in tables (no other lines should which describes the main findings and conclusions. Five to ten keywords appear) and that entries in the body of the table should be centred should be provided for full papers. Provide a short, descriptive title and (except for those in the first column and those involving decimal points an abridged title for the running head. Where relevant, the title should which should be aligned with the decimal points). Avoid duplication of mention the English and scientific names of the study organism/s, material in the text. All illustrative material should be arranged as figures include the order and family names in parentheses, and make reference and numbered consecutively. Line illustrations, lettering, numbers and to the geographical area covered. Number all pages consecutively symbols must be bold and large enough for up to 50% reduction. through to the end of the references. Hand-drawn lettering on figures is unacceptable. Maps should usually include a scale-bar, co-ordinates and an indication of orientation. Each CONVENTIONS: Authors should carefully study the latest figure should be on a separate page and legends must be provided. Give edition of the Novitates for guidance as to the conventions to be a key for symbols in illustrations. Photographs will be considered if they followed in the text, tables, figures, titles, legends, references, etc. are essential to the text and must be of high contrast and allow Follow these exactly. Give the scientific name at the first mention of a reduction. Where figures involve multiple illustrations/photographs, species both in the summary and in the text. Thereafter, either the these should be labelled alphabetically, commencing in the top lefthand English or scientific name may be used. The English names of species, corner. Authors must design all tables/figures with the page size of the except for birds, should not normally be captialised. Usage of names should follow standard texts. Scientific names (except for taxa higher Novitates in mind. than genus) and foreign words must be in italics. Metric units and their international symbols are used throughout, and a full-stop is used for the REFERENCES: References are listed in alphabetical order of the decimal point. The 24-hour clock is employed (02h30 and 14h30) and first author’s name in the following form: dates should be written: 7 October 1952. Use hyphens as sparingly as BURNHAM, K.P., ANDERSON, D.R. & LAAKE, J.L. 1980. Estimation of possible. Abbreviations such as loc cit., op cit., ibid. and idem. are not density from line transect sampling of biological populations. Wildlife used. Quotations should be accompanied by author, date and page Monographs 72: 1-200. number, e.g. (Gresbach 1888: 76), and the relevant reference should be BLAAUW, F.E. 1897. A monograph of the cranes. E.J. Brill: Leiden. included in the reference list. BLACKMAN, J.G. 1978. The swamp. In: LAVERY, H.J. (ed.), Expedition north, pp. 147-183. Richmond Hill Press: Victoria, Australia. SYSTEMATIC ACCOUNTS: Papers that include systematic Journal titles should be given in full; do not abbreviate these. Note the descriptions should present these under the heading ‘Systematics’. The use of hanging indents. Follow these conventions exactly when description of species (and subspecies) should be preceded by a higher presenting reference lists. taxonomy including at least family, genus and type species, and should take the following form: PROOFS AND REPRINTS: Electronic proofs will be e-mailed Family EXOGYRINAE Vialov, 1936 to authors and their prompt return is essential. The cost of changes by Genus Aetostreon Bayle, 1878 the author at the proof stage will be charged to the author. Authors will Type species: Gryphaea latissima Lamarck, 1801; by the subsequent be e-mailed pdf reprints of papers. designation of Douvillé, 1879. This higher taxonomy should be followed, in order, by the name of the ADDRESS FOR SUBMISSIONS species under discussion, list of figures, synonymy list, Diagnosis, Type David Allan, Editor, Durban Natural Science Museum Novitates, material (including holotype, allotypes and paratypes), Referred material, Durban Natural Science Museum Locality data, Habitat/Ecology, Description, Discussion and Distribution. P.O Box 4085, Durban 4000, South Africa; Morphometric information should be presented in the form of a table. e-mail: [email protected] DURBAN NATURAL SCIENCE MUSEUM NOVITATES VOLUME 37 2014
Editorial - South African natural science museum journals - the past 50 years (1964 - 2013): going down swinging or adjusting to the right fighting weight? David G. Allan ……………………………………………………………………………………………………...…….. 1-6
The Cainozoic palaeontology and stratigraphy of KwaZulu-Natal. Part 5. The False Coral Limestone Formation. Stratigraphy and fauna. Michael Robert Cooper ………………………………………………………………………………………….…….... 7-24
Effects of hydrocortisone and sodium methohexital on growth rate of Chrysomya chloropyga weidemann (Diptera: Calliphoridae): developmental and behavioural indications of presence of drugs. K.A. Williams & M.H. Villet …………………………………………………………………………………………...... 25-29
Observations on the African grass lizards Chamaesaura Fitzinger (Reptilia: Sauria: Cordylidae) in Swaziland with emphasis on fire impacts on populations in Malolotja Nature Reserve. Richard C. Boycott …………………………………….……………………………………………………………….... 30-39
Moult of the Grey-headed Gull Larus cirrocephalus in South Africa. A.M. McInnes, D.G. Allan & L.G. Underhill ……………………………………………………………………...... 40-46
Decrease of Coqui Francolin Peliperdix coqui (Aves: Phasianidae) in southern KwaZulu-Natal, South Africa. G.B.P. Davies ……………………………………………………………………………………………………………... 47-53
A major waterbird breeding colony at Lake Urema, Gorongosa National Park, Moçambique. G.B.P. Davies …………………………………………………………………………………………..………………..... 54-57
White-browed Sparrow-Weaver Plocepasser mahali (Aves: Ploceidae) in KwaZulu-Natal, South Africa. H. Dieter Oschadleus ……………………………………………………………………………………..…………….. 58-61
Recent records of the White-winged Flufftail Sarothrura ayresi (Aves: Sarothruridae) in South Africa, including details of a survey of high-altitude wetlands in 2013-14 G.B.P. Davies, H.A. Smit-Robinson, I.M.M. Drummond, B. Gardner, S. Rautenbach, D. Van Stuyvenberg, C. Nattrass, M. Pretorius, D.W. Pietersen & C.T. Symes ……………………………………………………………. 62-75
ISSUED: JUNE 2015 PRICE: R35.00 ISSN-0012 723X