Published in the United States of America — 2019 e VOLUME 13 « NUMBER 2
AMPHIBIAN & REPTILE
CONSERVATION
IN wes to William R. Branch (1946-2018)
amphibian-reptile-conservation.org
ISSN: 1083-446X eISSN: 1525-9153
Cover photo legend, in Bill’s own words:
This remains one of my favourite shots, although it is an old slide and this is the best digital scan | have of it (but of woefully low res). It was taken about 25+ years ago, and | was driving to work when | saw these flowers in bloom beside Port Elizabeth airport. | had an image of a cobra rearing in front of them, the Port Elizabeth Snake Park had just got a beautiful Cape Cobra in from the Northern Cape, and so | asked Rob Hall to come and help manipulate the snake. | didn't have a long lens and so had to lie on my belly with a 55mm Nikkon with 1.4 convertor. | used in-fill flash, held by Rob about 1m away and to soften the deep shadow under the snake's belly. | kept shuffling forward to get a more dramatic pose and had taken several shots when the snake disappeared from the viewfinder. Rob was standing to the side holding the flash and also a snake stick to ward off the cobra. When the snake disappeared | instinctively rolled back, heard Rob shout "Shit, that was fast!", and the snake bit the camera body about 6cm from my shutter finger. A bead of venom glistened on the camera body. Looking through the lens | had lost all sense of distance and simply got too close to the snake. It remains the closest I've come to a snakebite. Technically the picture works because the snake is alert but its mouth is shut and it is not looking straight at the camera. It therefore doesn't appear too threatening, allowing viewers to admire what remains my favourite snake. Bill Branch
Official journal website: amphibian-reptile-conservation.org
Amphibian & Reptile Conservation 13(2) [Special Section]: i-—xxix (e186).
Compilation of personal tributes to William Roy Branch (1946-2018): a loving husband and father, a good friend, and a mentor
1*Werner Conradie, *7Michael L. Grieneisen, and *Craig L. Hassapakis (Editors)
'Port Elizabeth Museum (Bayworld), P.O. Box 13147, Humewood 6013, SOUTH AFRICA *School of Natural Resource Management, George Campus, Nelson Mandela University, George 6530, SOUTH AFRICA *Department of Land, Air and Water Resources, University of California, Davis, California 95616, USA *Amphibian & Reptile Conservation (amphibian-reptile-conservation.org) and Amphibian Conservation Research Center and Laboratory (ACRCL), 12180 South 300 East, Draper, Utah 84020-1433, USA
Abstract.—Personal contributions to William “Bill” Roy Branch by famly members and colleagues: Colin Tilbury, Alan Channing, Dot Hall (Pitman, Basson), Rick Shine, James B. Murphy, Luke Verburgt, Julian Bayliss, Michael F. Bates, Pedro Vaz Pinto, Kirsty Kyle, Krystal Tolley, Mzi Mahola, Brian J. Huntley, Roger Bills, Johan Marais, Mark-Oliver Rodel, Paul H. Skelton, Aaron M. Bauer, Stephen Spawls, Andrew Turner, Ernst H.W. Baard, Amber Jackson, Margaretha Hofmeyr, Jens Reissig, Harold Braack, Atherton de Villiers, Marius Burger, Mike
Raath, Werner Conradie, and Martin J. Whiting.
Keywords. Influence, contributions, farewell, African herpetology, history, researcher
Citation: Conradie W, Grieneisen ML, Hassapakis CL (Editors). 2019. Compilation of personal tributes to William Roy Branch (1946-2018): a loving husband and father, a good friend, and a mentor. Amphibian & Reptile Conservation 13(2) [Special Section]: i—xxix (e186).
Copyright: © 2019 Conradie et al. This is an open access article distributed under the terms of the Creative Commons Attribution License [Attribu- tion 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.
Received: 5 August 2019; Accepted: 5 August 2019; Published: 10 September 2019
On 14 October 2018, William Roy Branch, or simply Bill as he was known to most, passed away after a short struggle with motor neuron disease. He was not only one of South Africa’s most well-known and respected herpetologists, but also a dedicated husband, a father, a good friend, and a mentor to so many of us. We have taken this opportunity to collate personal tributes from family, friends, and colleagues, to showcase the influence Bill had on our lives and careers.
Tributes from family members
Donve Branch (Bill’s wife)
Bill was an amazing man with a huge passion for life. When we married I introduced Bill to the world of pots and potters, and he introduced me to the world of reptiles and herpetologists. Very different worlds, but they became one we both loved. Over the years I was privileged to meet and host many of you. If I sometimes looked stunned when you arrived at our door, please forgive me. Bill very often failed to tell me we would be having a guest. Together we started to collect art, succulents, and books. None of which we could afford,
Correspondence. * werner@bayworld.co.za
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but we couldn’t resist.
Bill was a family man who loved and was so proud of his three sons. When we married his generous heart took on my children and grandchildren with the same warmth. Science was his passion which he loved to share. Bill in lecture mode could not be halted. His sense of humour was legendry. A kind, gentle man but also a humble man. He never boasted of his achievements. In his later years, these qualities made him so popular with National Geographic travellers.
A man of huge intellect with a broad knowledge of all things. A kind, generous, and wonderful man. Truly a real mensch. I was so proud to be his wife. He is greatly missed.
James Vlok (Bill’s stepson)
Bill Branch was a man of passion for his craft and natural science. He was an adventurer and an explorer; a man who inspired motivation and discovery of the world around us. He could keep you interested with a keen knowledge and a sense of humour that would have you laughing and learning. He will be sorely missed by family and colleagues alike.
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Christian Vlok (Donveé’s grandson)
Gumps told me so many interesting stories about his trips. He gave me my first Masai machete and a lizard. I knew I could ask him anything and he would know the answer. I will miss my Grandpa Gumps so much.
Analeah Vlok (Donve’s granddaughter)
I loved Gumps because he taught me so many things. He taught me which plants I can or can't eat, and about snakes and frogs, which I love. I miss him and every time I go into his room I think of him.
Jenny Vlok (stepdaughter-in-law)
Bill, to look at all things herpetological on a daily basis and know that I can't ask you any more questions about it, fills my heart with such sadness. You were so patient in your explanations, always interesting and funny. With your mismatched socks and wild hair, your fancy salads and poor man's capers, hilarious Easter egg hunts with a difference, cheeky Halloween surprises and Christmas gifting, not only were you an Amazing scientist but also an inspiration, and a motivator, allowing my children to be knowledge bearers and researchers in their own environment. We love you Dear Bill, and will miss you always.
Nicole Kingston (Bill’s stepdaughter)
Bill was a rock, a voice of reason, and a safe place and so loved. I am so truly privileged to have known him, and am a better person for it. His kindness, empathy, and wicked sense of humour will not be forgotten.
Oliver Kingston (Donve’s grandson) Grandpa made me laugh lots and 1f you wanted to know anything he was the person to ask.
Will Kingston (Donve’s grandson) He was kind and knew a lot about snakes.
Tributes from friends and colleagues
Colin Tilbury
KwaZulu-Natal, South Africa
It was early in 1980. After a year as a junior medical officer at Ngwelezana Hospital in KZN [KwaZulu-Natal], I had collected a series of cases of snakebites from Atractaspis bibronii and the Mozambique Spitting Cobra. With the data in hand I had approached Alan Channing, the then- chairman of the Herpetological Association of Africa, for comments. Being more of the toad persuasion, Alan suggested that I contact his colleague Bill Branch, the incumbent curator of herpetology at the PE [Port Elizabeth] Museum, who had shown more than just a passing interest in snakes and snakebite, and might be in
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a better position to help me.
I wrote to Bill and offered to assist with any affairs of the herpetological kind from Zululand. Bill wrote back immediately, expressing a keen interest in the snakebite data and also wondering if I might be able to collect some of the local Pelusios for karyotyping. We met for the first time a few months later. Bill was visiting Durban, and Sarah and I arranged to meet him at the British Middle East Indian Sporting and Diners’ Club near the Greyville race course, to sample the local curries. A truly memorable evening (I still have intermittent diarrhoea). I think that I may also have introduced Bill to the pleasures of a good red wine—or was it vice versa?
And so began a friendship which lasted nearly 40 years.
Driven with a boundless energy and an amazing zest for life, sharp wit, wry humour, and capacity for sharing, Bill attracted people to him. Whether by active involvement or by association, he had a lasting impact on all those who encountered him. Bill adored the simple things in life, and lived his life simply. He loved the camping and field trips that were an integral part of his work and which provided him with so much satisfaction. An avid angler since childhood, he had pulled many a carp from the rivers and dams of the Eastern Cape. Bill’s laboratory and office in the Port Elizabeth Museum was always a wondrous place to visit. Beyond the entrance door which was plastered with a selection of humorous “Bill” references, a mixture of chaos and creativity, preserved snakes and lizards in piles, the air reeking with alcohol, and Bill smiling happily. Bill and Donveé’s lovely home in Port Elizabeth was in many ways an extension of his beloved office at the museum. Of the many enduring mental images that capture Bill’s essence for me, are none more so than those of Bill at work in his man-cave at home. More like a ‘control room,’ his desk surmounted with massive computer screens and surrounded on all sides—floor to ceiling—with books, paintings, and photographs (including his all-time favourite of the yellow Cape Cobra that had nearly bitten him). Shelves were packed with w.1.p. files and books with titles covering an eclectic array of topics from tadpoles to volcanoes, fossils, sunbirds, euphorbias, mesembs, and every conceivable reptile and amphibian genre.
At home, but outside his study, every nook and cranny was adorned with paintings and Donve’s beautiful pottery. Each windowsill in the house was crammed with weirdly-shaped, rare, and spiky plants. Their garden was an indigenous plant paradise with a few thorny exotics, a haven for birds and local wildlife where the largest Palystes rain spiders in the world were free to roam— although strangely I only ever saw them on the walls of the guest bedroom. Theirs was clearly a home they loved to live in and was always open to the many guests who might pop in and stay over.
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One could not know Bill and be unimpressed with his amazing intellect. Bill read—-no—he devoured books by the ton. I have never met anyone who had such a command and broad understanding of natural history. He could have been anything from botanist, ornithologist, entomologist, mammologist, geologist, physicist—you name it. The reality in fact, is that he was all of these things and many more; such was the breadth and depth of his knowledge. His intimate understanding of the intricacies of natural history, the environment, and the interconnected webs of life, filling in the dots on life’s canvas one by one—or in Bill’s case, by the dozens at a time.
In spite of his huge talents, he kept his feet firmly on the ground and freely shared his knowledge and wisdom with anyone who asked for advice or input. He was an inspirational force to anyone and everyone who had the privilege to know or work with him; a truly benevolent gentle giant and an incredibly productive scientist. The herpetological community around him was so privileged to have him as a guide and mentor. In the decade following his retirement from the museum, he worked as a specialist guide for over 50 National Geographic touring parties. These afforded him opportunities to continue to pursue herps in many iconic African locations.
As a friend, Bill was caring, insightful, non- judgmental, and always with a wonderful sense of humour just bubbling beneath the surface. As a storyteller he had few peers: in his clipped British accent with the hint of a lisp mumble and a wry smile, he would gleefully extol the excruciating agony of the many unfortunates who became the subjects of his tales. Of course, these often involved his hapless colleagues on the many field trips that he made. Quick, dry, wicked, invariably veiled in intrigue, he would construct the twists and turns of his story to extract every molecule of humour. His punch lines always immaculate.
Over the years, I spent a great deal of time outside the borders of South Africa, but Bill always found time to write and give updates on his projects and movements.
After the birth of our first child in London in July 1989, Sarah and I sent out a short notice of his birth to a few friends and relatives, making reference to ‘the discovery of a new species of the TILBURY genus found lurking in the St Helier 's Labour Ward at precisely 03h45 hours on 25 July 1989. It is wriggly, pink all over, devoid of scales and tail, and makes characteristic feeding cries every 4 hours. It weighed 3.63 kg on discovery, and has the features characteristic of the male sex. It has been named Douglas Matthew.”
I left London a week after the event and headed back to my job in Saudi Arabia. Shortly after my return to Khamis Mushayt, I received a letter from Bill:
“Dear Colin,
Congratulations on the arrival of Douglas Matthew. You must be looking forward to Sarah and DMT
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arriving at the end of the month, although I suppose that throwing all the Cerastes out to make way for the cot must be a bind. He will slow your globe-trotting down a bit, but it will only be about 12 years before he is useful in the field! Robbie and Matthew do all the hard work in the field now, so they do have their advantages.
[feel | must take exception to the new name however. Looking through the London telephone directory I came across three other references to Tilbury Douglas Matthew (usually from the poorer eastern suburbs besides the Thames). All had priority, some dating from the early 1930's. Your new name is thus pre-occupied, and according to strict nomenclatural rules (Int. Rules Zool. Nomenclature, rev. ed. London 1986; page 25, paragraph 3), becomes a strict junior homonym and is invalid. As well as afflicting the young lad with a used name, it is also incorrectly formed according to the rules governing construction of names. Being the first scientist (of truly international standing) to have spotted this error, I claim my right to propose a replacement name. I have chosen:
Tilburyanus inhirsutus arabicum Branch 1989
You will note that the Generic name is now correctly Latinized and the ending is more appropriate (being his most obvious feature for the moment!). The specific epithet also refers to the sub-adult plumage, while the sub-specific name is a traditional, uninspiring geographical allocation. Knowing that he is now correctly named, you may re-apply for birth certificates, passports and driving licences etc.”
For a man who played with snakes, Bill had a simple philosophy. Respect them and you won’t get bitten, and as far as I know, apart from a single dry bite from a Thelotornis, he never did. I remember the day that I brought a small shiny black snake all the way from the DRC [Democratic Republic of the Congo] to Bill’s home, and proudly handed him the blue cotton bag that contained the snake which I had carefully nurtured for the previous month or so. Bill gleefully but carefully opened the bag and peered inside. Then to my horror, he inserted his hand into the bag to retrieve the snake.
I said ‘Whoa! Hang on there a minute; I just want to get out my notebook and camera to record the first bite from this unknown species of Atractaspis.’ Bill pulled out his hand, the snake dangling limply between his fingers. Rigor mortis had already worn off.
“You've killed it” I said.
IN Oe ete nit excuses Do you think it is a Norwegian Blue?” (A joke that can only be appreciated by followers of Monty Python).
But it was not only herps that Bill would talk about. As much as he was a scientist, he was also a profoundly loved family man who would talk with pride as much about his loved ones as he would about his work. Give him half a chance and he would talk for hours about his
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sons Robbie, Matthew, and Tom and his new family— Nicole, Anthony, and James.
It was in 2012 that Bill first realised that he was only human after all, when he contracted malaria in Mozambique. That nearly finished him off. He eventually bounced back to his old self, but he was unable to dodge the bullet of MND [motor neuron disease]. When Bill contacted me in February 2018 to say that he had been diagnosed, it felt as if a tree had fallen on me. I had the good fortune to be able to spend several days with Bill over the last months of his life; and to be able to share memories of the good times, laugh together, discuss the iniquities of life, and to acknowledge the simple fact that we are all just fulfilling our biological destiny—albeit in different ways.
One cannot write about Bill without acknowledging the major part in his life that Donvé played, as his partner and soul mate, and in turn appreciate the huge hole that has been left behind by his passing in Donve’s life. In one conversation we had, we both agreed that it was one of the greatest privileges of life to be able to love and be loved back unconditionally. I don’t think that anyone can overestimate the enormity of this gift. She made him so happy and in the end, so sad that his Dove would have to endure the last days of his life with him in the state he was in.
Bill asked me to sign as witness to his living will to not be placed on any mechanical machine that would prolong his life. As his MND advanced, even in the late stages, in spite of his body being totally paralysed, his mind was as lively as ever; as he fought day by day to extract, utilise, and enjoy to the last moment every second that was left to him. He was immensely saddened and so disappointed that he had run out of time to complete all the many projects that he was part of or had initiated. His illness had quite literally pulled the rug out from beneath his feet. I know that Bill handed over many of these to colleagues to finish—we should make him proud. Even as he inexorably neared the end, he was so brave in facing his fate. He could still make jokes about this. He once compared himself as a likeness of the blue- headed agamid that was named after him (Acanthocercus
Fig. 1. Cover image of Hyperolius raymondi used for Frogs and other Amphibians of Africa (Photo: Bill Branch).
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branchi). Finally in the afternoon of 14 October, dulled by the ever-increasing CO, levels, he finally and peacefully breathed his last. The end to a magnificent life. His was an act we could all learn something from.
More than anyone, Bill understood and appreciated the fact that no one gets to live forever, but that everyone is hopefully gifted with the opportunity to leave a footprint embedded in the rocks of humanity—a footprint that will endure with a permanent relevance to those who follow one’s trail. Bill had big feet for such a small frame, and no doubt we will be following his prints for many a year. I can only say that I was privileged to know Bill, and even more so, to think that he might have considered me to be a friend.
The memories of Bill will be enduring and he will always be celebrated as one of the world’s leading herpetologists of our time. He will be sorely missed and long remembered.
Alan Channing
University of the Western Cape/North-West University, South Africa
I met Bill at a herp meeting while he was working at the Atomic Energy Board in the 1970s. He was hugely enthusiastic and well-read. Later, I was happy to support his application for the post of Herpetologist at the Port Elizabeth Museum, when asked by the Director. We undertook many field trips together, and for a while we formed a collaboration for funding from the forerunner of the National Research Foundation.
Although Bill and I worked on different groups, there was always a lot of friendly banter between us. His sense of humour was displayed on one field trip to northern Namibia, when he offered to cook the potatoes, while I prepared the meat. When it came time to eat, the potatoes were still crunchy. Bill's response was to explain that that was how they were cooked in Cornwall, and that it was a classical culinary procedure!
I will miss Bill's insights and our regular email exchanges. He provided a number of excellent photos for the upcoming book Frogs and other Amphibians of Africa, and was always willing to help, or offer a beer and a meal, when I was in Port Elizabeth.
as - i =
Fig. 2. Bill p otographing a lizard in southern Angola, 18 January 2009 (Photo: Alan Channing).
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Dot Hall (Pitman, Basson)
Port Elizabeth Museum (Bayworld), South Africa
A flood of memories flow through my mind when I reminisce on the very small part of Bill’s life I shared. One in particular always makes me smile. When Bill joined the Museum in 1978, he was a complete unknown. We were observing his introduction to the staff with interest: A quiet, rather serious little man? From his first day he was a regular library user. He was passionate about books. Each visit he made to it was a learning experience for me. He freely shared his knowledge and always stretched my way of thinking.
On one memorable morning, shortly after he had joined the staff, all was quiet in the library when a strange scuffing noise caught my attention. No-one was in the library, so I put this observation down to my imagination and continued working. The same noise recurred several times till I eventually decided to investigate. There was a solid counter that separated the librarian from those using the library. I peered over this counter to find Bill on all fours, crawling behind a large leguaan [varanid] holding its tail and trying to direct it around the corner to my desk. I guess it was being a little uncooperative and his full attention was required for him to achieve his goal—“Frighten this librarian out of her mind!!!” After observing the scene for a short while I decided to launch a surprise “attack” from the back and gave him a pinch on his rear end. His reaction was marvellous. The leguaan was let loose and his fright was complete.
Fig. 3. Bill Xerox-ing a puffadder to make counting of scales easier, to the disgrace of the librarian (Photo: Dot Pitman).
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We both enjoyed sharing this amusing moment. How many million more smiles has he given to the vast number of people with whom he associated?
Rick Shine
Macquarie University, Australia
I first met Bill Branch on the morning of Tuesday the 5th of September 1989, at the British Museum of Natural History (now the Natural History Museum London). Like me, he had travelled to the UK to attend the First World Congress of Herpetology, and like me, he took advantage of the opportunity to visit the British Museum of Natural History. Bill was looking for type specimens of African herps, and I was attempting to track down the reptile specimens that Charles Darwin collected in Australia during the voyage of the Beagle. As we sat and talked over lunch, I was astonished at Bill’s breadth of knowledge about the African herpetofauna, and his intimate familiarity with the scientific literature on those animals. But I had no idea that we would end up as collaborators on a major project.
Five years later, I took my first (and only!) sabbatical from the University of Sydney. My wife Terri and I had always wanted to see the famous game reserves of southern Africa, and our oldest son was about to turn 12—after which time he would have to pay a full fare on the airlines rather than half-price! So I contacted Bill about the possibility of dissecting preserved snakes in African museum collections for ecological data
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(gut contents, gonads, etc.) as I had done for snakes in Australian museums over the previous years. Bill was enthusiastic, but thought that the best collection might be in Namibia, where over 1,500 snakes that had drowned in an open canal (the Eastern Water Carrier) had been preserved by the local wildlife authorities. And with the first South African elections looming, and political unrest likely as power shifted from ‘Afrikaaners’ to native Africans, Namibia looked like a quieter, safer option than South Africa for a family with 10-year-old and 2-year- old children.
We flew to Namibia while Bill drove the museum’s Kombi-van from Port Elizabeth to Windhoek to meet us. And being Bill, he had a much better idea than staying in Windhoek to dissect the snakes—instead, we piled them into the van and took off for Gobabeb, where we could enjoy the spectacular dunes in between long hours of peering inside dead snakes. The Aussie team (me, Peter Harlow, and Jonno Webb) peered inside the innards of dead snakes and called out numbers, while Terri wrote them all down into data-sheets. Bill carefully examined every half-digested frog and reptile that came out of a snake stomach, almost always managing to ID it, even if he only had a few toes to work with. It was a happy and effective team.
After we finished the Namibian snakes, my family flew off to the USA while the rest of us drove down to Pretoria to look at MORE snakes at the national museum in Pretoria. It was a classic herp “road trip,” with frequent detours to look for specific taxa (usually, so that Bill could get a photograph for his field guide). We made an obligatory stop at Poffadder (= “Puff Adder’) one of the few towns named after a snake, near the border between South Africa and Namibia. A photograph I took on the town’s outskirts captures the relaxed joy of herpetological zealots indulging their passions (Fig. 5). We worked long hours in Pretoria, obtaining a mountain of data that eventually translated into 15 papers on the natural history of several major lineages of African snakes. We also sampled the local beer and watched World Cup soccer games at bars downtown—horrifying some of the locals who were convinced that we would be mugged as we walked the streets at night.
Throughout this first African adventure, Bill was fantastic. Extraordinarily knowledgeable, with a vast network of contacts, he made the project possible. We talked long and often about everything from fishing to the mysteries of bureaucracies and families—and especially, about snakes. Hopping off a plane and looking inside preserved specimens can generate a lot of data— but it was Bill’s long experience that enabled us to put that information into context. For many of the species about which we wrote papers, I had never even seen a live specimen—but Bill had, and his firsthand knowledge helped him to laugh off my ill-informed speculations, and keep our interpretations true to the reality of snake ecology in southern Africa. Bill was a terrific collaborator
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Fig. 5. Jonno Webb, Bill Branch, and Peter Harlow posing at the outskirts of the town of Poffadder in northern South Africa, reveling in the idea that somebody actually named a town after a snake (Photo: Rick Shine).
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and a wonderful friend. I feel privileged to have been able to work with him.
James B. Murphy
Division of Amphibians & Reptiles, National Museum of Natural History, Washington DC, USA
When we first met at a herpetological conference in the US many years ago, Bill and I noticed that our love of amphibians and reptiles, overall biological interests, and personal histories were strikingly parallel. One major difference was that Bill had completed a Ph.D. inchemistry and I barely passed my chemistry courses. Fortunately, he changed trajectories and excelled in herpetology. As we shared our stories over some beers until the break of dawn, Bill and I quickly bonded. I invited him to come to Dallas, Texas, where I was herp curator at the Dallas Zoo with a spacious guest room available in my home. As we toured the Zoo’s herp collection, Bill was delighted when he saw the large breeding group of New Caledonian Geckos (Rhacodactylus leachianus, Fig. 6). There was a particularly large and impressive male that was surplus, so I gave it to Bill—his stunned reaction and gratitude were wonderful to watch as he carefully packed the saurian to hand-carry it back to Port Elizabeth [ED note: This gecko is still alive in the Port Elizabeth
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Museum as of July 2019].
After the First World Congress of Herpetology in Canterbury, UK, he invited me to stay at his parents’ home nearby until we later went to Bonn, Germany, for the first Varanid Symposium held at the Museum Alexander Koenig (Fig. 7). In my view, Bill was a pretty stocky fellow, but his mother was concerned that he was not paying enough attention to proper nutrition, so she followed him for several days with handfuls of vegetables, insisting all the while that he was becoming a mere slip of a man. The scenario reminded me of a Monty Python skit.
At the varanid meeting, Bill presented a wonderful lecture on the White-throated Monitor (Varanus albigularis)—“The Regenia_ registers of Brown (1869-1909). Memoranda on a species of Monitor or Varan.” Branch covered all aspects of Alfred ‘Gogga’ Brown’s extensive observations—sex ratio, size, body proportions, hemipenial morphology, visceral fat bodies, coloration, diet, cause of death, longevity, reproduction, gestation period, egg laying, oviposition, eggs, clutch size, hatchling size, incubation period, growth, behavior, mating behavior, shedding, thermoregulation, predation, parasites, exploitation, and seasonal activity and retreats. The amount of information that Gogga had collected on his captive lizards and in wild counterparts in the late 19th century is truly astounding.
Over time, Bill sent anumber of African and Namibian reptiles for the Dallas Zoo collection, including Angulate Tortoise (Chersina angulata), Parrot-beaked Tortoise (Homopus_ areolatus), Tent Tortoise (Psammobates tentorius), Mountain Adder (Bitis atropos), Dwarf Adder (Bitis rubida), Many-horned Adder (Bitis cornuta), Cape Dwarf Chameleon (Bradypodion pumilum), Lesser Flat Lizard (Platysaurus guttatus), and Drakensberg Crag Lizard (Pseudocordylus subviridis).
In the ensuing years, we spent much time together at meetings and he shared his concern about shrinking funding for the Port Elizabeth Snake Park and Museum. Frederick William FitzSimons (born 1875) was the first Director of the Museum in 1906, and he developed the
Fig. 6. Male New Caledonian Geckos (Rhacodactylus leachianus) still alive in Port Elizabeth Snake Park (Photo: Werner Conradie).
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Fig. Z Varanid Symposium participants at Museum Alexander Koenig in 1991. Bill Branch is the seventh person from the left, in the front row.
Snake Park. His son, Vivian, assisted him and both of them published in herpetology. His younger brother, Desmond C. FitzSimons, started the Durban Snake Park. F. W. FitzSimons also wrote books on the natural history of South African mammals, including primates.
Bill was a consummate biologist whose contributions to our knowledge of African amphibians and reptiles over several decades set the high standard for herpetological work. His nominal retirement as curator of herpetology at the Port Elizabeth Museum occurred after many years of service. As far as I know, he did not free-handle venomous snakes nor put them on his head. Every time we met, I could be confident that he would cover subjects virtually unknown to me. He will be missed.
Luke Verburgt
Enviro-Insight & University of Pretoria, South Africa Bill replied to my email almost instantly and in great detail! I'd been very hesitant to contact my herpetological idol about a reptile identification query, because I guess I was afraid to disturb such an important person with possibly silly and trivial queries from me, a nobody. Yet to my delight, Bill took the time to carefully answer my questions, providing great detail and assistance. No admonishment for not having read the appropriate books/papers, and no arrogant stance regarding my lack of herping credentials! I was thrilled, and it opened up communication between us to such an extent that soon we were communicating about African herpetofauna via email quite regularly, with Bill always helpful and kind in dispensing his amazing wealth of knowledge. Like a mentor really.
I eventually met Bill in person months later in Namibia, along with Johan Marais and Aaron Bauer, while they were on a collecting field trip. It was such an honour to be sitting around the same table as these herping heroes, and I was rather star-struck. After some fieldwork with the team, I picked up on the fact that the species of Rhoptropus that we were collecting was not the one I had expected to be there according to Bill’s field guide, which was really my main source of herp
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knowledge, as it has also been for so many others. I eventually plucked up the courage and cautiously approached Bill one afternoon, to ask about this Rhoptropus situation. He laughed heartily and said, “Oh, that map is complete rubbish!" I couldn't believe it. The author of the book that I cherished above all others just told me that some of it wasn't the complete truth! And right there I learned two massive lessons from Bill about African herpetology: imperfect data from under- sampling abound, and not being afraid to question the existing understanding.
Sadly, that was the one and only time I got to be in the field with Bill, and it was far too brief. Thankfully though, we collaborated a lot after that with several resulting papers where I got to be a co-author with Bill—a huge honour! However, the greatest honour for me in this regard was having Bill as a co-author guiding my very first reptile species description, an interaction through which I learnt more than I could've ever imagined. He took what was a pretty ordinary and mundane manuscript and guided me on how to improve it to an acceptable standard, the standard which he was instrumental in setting for African herpetology.
After that I regularly reported to Bill, who was always extremely interested in my findings because I was often working in poorly sampled rural places across Africa. In his now familiar mentor role, he would encourage me to do as much useful sampling as possible, and also to think harder about why a particular species was observed in the habitat I found it in and, therefore, to consider its ecology in greater detail and gain more insight from my observations. In short, Bill made me a better herpetologist and I am forever grateful for his friendship and his mentoring.
Although I didn’t see Bill in person very often, it was always a treat to hang out with him and his fantastic sense of humour. But what I think I enjoyed the most was to hang out with him and to see him having such fun at the 2017 HAA conference at Bonamanzi, and I was even lucky enough to win a “selfie” with him! Unfortunately, I never actually received the “selfie’— but fortunately, Shivan Parusnath managed to capture the “selfie”’-taking moment perfectly, and it is my favourite photo of Bill and myself (Fig. 8).
I received the news that Bill had passed away while I was sampling in the Cabinda Province of Angola, an area of great interest for Bill. While we all had known for some time that it was an inevitability, the news of his passing came as a massive shock to me because only a few hours earlier, during his last night, Bill had somehow managed to send me a lengthy Whatsapp message, instructing me to collect as many DNA samples of certain species as I could due to the importance of the sampling locality I was in. And thinking about it now, that's just how it was always going to be for Bill, the ever-enthusiastic herpetologist and helpful mentor to the very, very end. Rest easy Bill, I miss you so much
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AANA Fig. 8. Taking a “selfie” with Bill as part of the prize for runner-up best photographer (Photo: Shivan Parusnath).
and hope that I am able to justify the effort you put into sharing your time and knowledge with me.
Julian Bayliss
Ecologist and Explorer, Wales
I first met Bill when he came to undertake a herpetological survey on Mount Mulanje in Malawi with Johan Marais and Michael Cunningham in 2005, as part of the ongoing ecological monitoring programme on Mulanje that I was coordinating. However, it was really when we met the second time, when Bill and Werner Conradie joined me on Mount Mabu in 2009, that we really got to know each other well. I had been working the mountains of northern Mozambique for several years prior to this event, and had managed to turn up several new species of snakes and chameleons, although my herp work was opportunistic (I discovered Atheris mabuensis by stepping on it!) and I needed professional assistance (Fig. 9). These discoveries attracted Bill, and we arranged for a trip to Mt. Mabu forest to collect more specimens, and also to see if we could collect specimens of a Nadzikambia chameleon that was only known from a couple of photographs taken on my previous visits. We were successful in this endeavour, and I managed to collect the first specimen of the Nadzikambia from Mt. Mabu which Bill named after me as Nadzikambia
Fig. 9. Photograph of Atheris mabuensis taken by Bill— probably the best photograph of a snake I have ever seen (Photo: Bill Branch).
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Fig. 10. The Mt. Mabu 2009 science team. Left to right: Werner Conradie, Martin Hassan, Julian Bayliss, Bill Branch, Hassam Patel, Colin Congdon, and Steve Collins (Photo:
Julian Bayliss).
baylissi. | was deeply honoured by this gesture.
The 2009 Mt. Mabu expedition proved to be a very enjoyable expedition, packed full of laughter, good company, and good food. I had also invited the butterfly crowd from the African Butterfly Research Institute (ABRI), a great bunch of eccentrics, and the stories flowed around the camp fires at night. At the end of the expedition, we all stood below a large tree on the forest camp in Mt. Mabu with Bill at centre stage (Fig. 10). This is one of my favourite photos of Bill, and it captures a moment in time where nothing outside that camp at that time really mattered. This was the start of a very good friendship with Bill (and Werner) and some great correspondents. However, one of my fondest memories of Bill was spending time with him in the Mt. Nimba forest in Liberia. It was part of an EIA on a proposed mining concession, and it was just the two of us for several days, which gave us plenty of time for chewing the fat; especially when we talked about rugby and Wales vs. South Africa or England, as I am from Wales and Bill was originally from England, and then South Africa. At that time, I had flown up from a festival in South Africa and brought with me a ‘Green Policemen’ helmet which Bill dually wore (Fig. 10, this photograph shows Bill beaming a big smile).
Bill, I will miss you greatly—you were an inspiration to me. Not only did you teach me a lot about reptiles, but you were also a professional in everything else you did. An expert and a gentleman. In the last communication I received from Bill, a couple of months before he passed, he told me ‘not to defer my dreams’—advice which is applicable to us all and advice I intend to follow.
Michael F. Bates
Department of Herpetology, Bloemfontein, South Africa
I knew about Bill soon after I started working at the National Museum in Bloemfontein in 1983, as he was then editor of the Herpetological Association of Africa’s journal. The first time I met him was at the HAA’s first
National Museum,
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Fig. 11. Bill Branch in the Nimba forests close to Nimba Mountain, Liberia (November 2011). Bill is wearing the green policeman hat I had brought with me from South Africa
(Photo: Julian Bayliss).
conference held at Stellenbosch University in 1987. I was only 25 at the time, and Bill was about 41, still quite slim and with a full head of black hair! At that time he was busy wrapping up work on the first edition of his famous reptile field guide. Even then I remember Bill having a certain charm about him and the aura of a man with a deep knowledge of his subject matter.
Over the years I visited Port Elizabeth Museum several times to examine specimens for various research projects, including some on which I collaborated with Bill. Having him all to myself and available to answer my barrage of questions was always special. However, I think my fondest memories were in the early 2010s when we spent considerable amounts of time editing the text for the Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland (published in 2014). As first and second (Bill) editors, the bulk of the editing fell on us. I would, for example, e-mail Bill the text for a species account and ask such questions as “‘is it still regarded as a subspecies” or “has anything been published about this recently.” I could count on him to respond within a day or two, and his responses were always insightful. He seemed always to be up-to-date with the latest taxonomy and the most recent literature. And so it was that we e-mailed the various sections of text back-and-forth until we were both happy. I have very good memories of those times.
Another special memory I have of Bill was in May 2018, a few months after he was diagnosed with MND, when I visited him at home in Port Elizabeth, together with Aaron Bauer and Marius Burger. By this time he was, for the most part, wheelchair-bound. Nevertheless, he was as talkative and interesting as ever, especially with regard to herpetological matters, and he also exhibited his usual great sense of humour. We spent most of the time at the computer in his study where he showed us photographs of interesting and new reptiles, and of field trips he had conducted with various colleagues over the years. Also, I brought him a copy of a recent taxonomic paper on egg-eating snakes (Bates & Broadley) that
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Fig. 12. Michael Bates (left) with Bill Branch and Darren Pietersen during the Herpetological Association of Africa’s conference in Pretoria in 2013 (Photo: W.R. Schmidt).
had just been published in the National Museum’s journal Indago. The front cover of the journal featured a montage of Bill’s excellent colour photographs of these snakes, and it gave me great pleasure to see how pleased he was with the way it turned out.
Bill had an enormous presence in the field of African herpetology. He impressed me as a very well-read man, and this was reflected in his wide and seemingly limitless knowledge of reptiles and amphibians. Bill was always willing to share PDFs of research articles and in this way he helped me on innumerable occasions. Also, I was inspired by his style of writing and attention to detail. I still think about Bill often and will miss him for several reasons, not least for the fact that his expertise was always just an e-mail away.
Pedro Vaz Pinto
Kissama Foundation, Luanda, Angola & CIBIO- InBIO, University of Porto, Portugal
I first met Bill in January 2009 in the most appropriate of places: deep in the Angolan Namib desert, in Iona National Park. We were part of a large group of scientists assembled by Brian Huntley for a biodiversity expedition in southern Angola. I remember approaching Bill after dinner in the camp site, and he was keen to see my photo files and became interested in some bush viper pictures, which led to a few engaging stories and discussions. At that point I was simply curious about reptiles, and more involved with furry or feathered creatures. The following day, I drove my Land Cruiser to where I could see Bill and his colleagues had parked their pick-up truck next to some granite boulders. I could sense some excitement in the party, so I asked Bill what they were doing. He invited me to join them and opened a little box to retrieve a tiny beautiful little gecko, one of the gems of Angolan herpetology which was not even formally described at the time: the endemic Plume- tailed Gecko, Kolekanus plumicaudus! He then showed what was special and unique about that species and chatted about other leaf-toed geckos. I was fascinated of course, and it was quite an introduction to reptiles. Over
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Fig. 13. Bill Branch processing specimens in the fading light of the Angolan Koakoveld (Photo: Pedro Vaz Pinto).
the following years we would become good friends, but looking back I’m still amazed to realize how generous he was by sharing that amazing find with someone he had just met the previous day. Other scientists would have kept their cards very close to the chest. But Bill kindly drew me towards the world of herpetology for which I’m forever indebted, but above all I believe he made me a better scientist and better naturalist. He taught me to make an effort at looking into the bigger picture, to see the multiple layers and connections that lie hidden behind the outer surface of a given ecological theme.
My best memories with Bill, without any shadow of doubt, were the days in which I was privileged enough to travel with him to some of the most remote corners, wildest places, and biodiversity hotspots in Angola. We would typically look for a scenic landscape off the beaten track and choose our camping spot. Some of the time shared with Bill, around the campfire in the Angolan desert, mountains, or forests, was memorable. Our camping expeditions were hugely stimulating scientifically, exciting and unpredictable, and very importantly, always bathed by loads of good humor! These expeditions could be physically exhausting, but soon after I was looking forward for the next trip with Bill.
Other scientists are much better prepared to praise Bill’s unique and extraordinary legacy to African herpetology. I can add that he did leave a crucial mark on Angolan herpetology, but tragically with his premature passing away, it wasn’t allowed to further crystalize during his life. He was arguably the most influential herpetologist to have worked in Angola for a sustained period, and is the main person responsible for bringing herpetology into the biodiversity agenda in modern Angola. I have no doubt that his pioneering role will be recognized in the future by young Angolan biologists. On a personal note, whenever we came across a new lizard or snake, I got used to my sons asking me ‘- Will Bill want this specimen?’, ‘- Has Bill identified this species?’, ‘- Does Bill need more specimens?’ and as result, these now rhetorical questions remain quite vivid
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Pe ws
Set
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Sy S \ : —— oh. Sa age S- Fig. 14. Bill Branch photographing a Jameson Mamba Angola with Ninda Baptista (Photo: Pedro Vaz Pinto).
in
and still drive me on my searches. There are still a lot of ‘goodies’ that we will catch for you Bill, and that’s a promise!
Kirsty Kyle
KwaZulu-Natal, South Africa
I had the good fortune of growing up at Kosi Bay with Bill Branch as a much loved family friend. Bill had gotten to know my parents, who were the resident scientist and his wife for the area. In those days, he did an almost annual foray to Zululand, what with it being such an interesting part of the country for herpetofauna. Whenever he moved through the area, with his pack of scientists, they would use our house as a base, and for my two older brothers and I this was just the best thing ever. His trips became the highlight of our year and I think he thoroughly enjoyed having three young, able-bodied slaves, ever so willing to dive after any reptile that was silly enough to stick its nose out 1n our vicinity. A friendly disagreement developed as we got older and started objecting to his pickling tendencies. In the later trips we would “not see” a lot of the more common species because dear old Uncle Bill would just pickle anything we presented him with, which was a bit hard on our budding conservationist hearts. Although we had a pretty much genetic interest in herps, I think those times with Bill were extremely formative in all three of our lives, they certainly were in mine. The fact that he was interested and enthusiastic in teaching and encouraging a little blonde thug of three years old in the ways of reptiles was amazing.
Bill was absolutely instrumental in setting me on the path I am on today. Throughout childhood it was a privilege to spend time in the field with him and just absorb all the information he so generously and freely dished out. It had a major impact on my interest in herps. I emailed back and forth with him whenever I found something interesting, and I sent him pictures of all sorts of different reptiles over the years and he would always respond in his warm, friendly, and encouraging manner, which was just amazing. My favourite memory of Bill would have to be on his last visit a few years back,
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when he proudly presented us with a beautiful pot that Donvé had made, decorated with an aloe he informed us he’d just plucked from our outdoor lizard enclosure. There were no flies on Bill and I loved that about him, he always told you the truth, even if it put him in not the best of lights. We still have the aloe in the pot.
I wish I had a picture with him from the early days because it really would be a cute one. I fondly remember parking on his lap as a very little girl, discussing whatever, feeling terribly important, with his black mop of curls and my blonde mop of curls. It would have been such a cute picture. I miss Uncle Bill, the world in general is a lot less fun without him and my Facebook is a much darker place without his frequent updates, pictures, thought processes, and quips. I hope he forgives me for specialising in amphibians instead of reptiles, and I’m incredibly grateful to have had him as a friend, as well as a mentor.
Krystal Tolley
South African National Biodiversity Institute, South Africa
I knew about Bill before I moved to South Africa in 2001, as he and Colin Tilbury had some chameleon DNA samples for my upcoming postdoc project. The project almost didn’t happen, as Bill and Colin got cold feet, but when I arrived I learned that they decided to let me give it a try. Their trust in a stranger with whom they had never worked ended up building a friendship and collaboration that lasted nearly two decades. As that project progressed and more projects arose, Bill encouraged and supported me both in a personal and a professional capacity. In fact, the entire herpetological community welcomed me, something that I was not used to, coming from the competitive world of marine biology in the northern hemisphere. Because of Bill, Colin and all the SA herpers, I felt like I had found a home that I didn’t want to leave, and Bill was instrumental in that. I cannot remember actually meeting Bill for the first time. My first distinct memories of hanging out with Bill and all the herpers is from the Port Elizabeth HAA conference in 2004. What sticks out in my mind 1s that at the concluding banquet, Bill received the Exceptional Contribution to African Herpetology Award and he was so touched by this that he wept. That spoke to his nature as a caring person who knew that strength and courage, not weakness, comes from personal relationships and bonds. And his connections with his friends are something he fostered.
I have many distinct and fond memories of Bill but strangely enough, most of them relate to our friendship, not to herpetology. When we would meet, the first things he would ask me about was how I was, how was my personal life, what was happening, was I happy? He had many wise words for me along those lines, giving advice, encouragement, and reassurances that eventually I would find my path. Then of course, the
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‘herp talking’ would start. He would go on for hours, non-stop, about snakes mainly. Most times, the topics were just beyond me. I tried to absorb what he said, but there was so much information that my brain couldn’t handle it. I do remember that a long discussion about Leptotyphlops made me realise what cool things they are, and I still hope one day to actually work on them.
I was fortunate to have the chance to visit Bill shortly before he passed away. We both knew, as did his wife Donveée, that I was there to say a final goodbye. This was indeed the last time I saw him and it was emotional for everyone, but my memory is still a good one. The same old routine was there. He asked me about my life first, and he gave me wise words and insight about life. Then he spoke about herps (including Leptotyphlops) for about four hours non-stop. The thing that was different this time, was that he often interjected the conversation with things about himself. Dreams and wishes, failures, successes, lost opportunities. He talked a lot about how it’s important not to waste time on petty or destructive things in life. But to focus time and energy on the people in your life that care about you and to never take that for granted. He spoke about the balance between the work related passions of a herpetologist, and that this has to balance with life, friends, and family. Bill was a hard worker, but he did focus on family and friends, and I don’t think he took any of that for granted. The way that his first questions always related to our personal connection and friendship, and about which analysis I was running, speaks to that. The wisest words that Bill ever said to me are: “Friendship 1s a gift. It’s a gift that others chose to give, and that you chose to accept in whatever form it takes.” Bill gave that gift to me and to so many others. That is what I will remember him for the most.
Fig. 15. Bill Branch and Krystal Tolley in south-western Angola in 2009 (Photo: Krystal Tolley).
Mzi Mahola South Africa I first met Bill when he arrived at the Museum. A year or two later, I invited him to join our Port Elizabeth Mu- seum soccer team, which was playing in the Industrial League. He didn’t play many matches, because of his
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commitments, but he was a good soccer player. A year later, I was transferred from my department to join and work with him as his research assistant. We often went to Sardinia Downs to tag, study, and monitor the move- ments, growth, and development of tortoises. After that he introduced me into his other research and study pro- gramme of other animals, such as frogs, snakes, and liz- ards.
One day we were going up the Zuurberg Mountains when he made a deal for us; “If you happen to catch a snake first, I will buy you a bottle of beer at the Zuur- berg Inn on the way back; but if I make the first catch, you will buy me a bottle.” That was fair enough for me. Bill was at the wheel of our Land Rover. We were driv- ing towards the forest at the foot of the mountain when I saw a female boomslang flying towards the forest. Bill noticed my hasty intention to open the door of the mov- ing vehicle and he quietly said, “Forget it! Boomslangs are very shy; you'll never catch it unless it is ina tree.” A few minutes afterwards, we left the Land Rover and with our hunting gear and went our separate ways. It took me less than five minutes before I heard the hissing sound of a slithering snake. I saw the disappearing tail of a rinkals entering a hole amongst rocks on a ledge. I put on my safety glasses and peeped into the hole, and saw the two shiny eyes watching the entrance. With my tools I pulled the snake out and put it in my canvas bag, and declared my victory to Bill. He didn’t believe me. It didn’t matter how many snakes he collected afterwards. I had beaten my master in his game and the bottle of beer would be a cherry on top.
We were on a trip to the Drakensburg Mountains and our first night stopover was in Centane, at my in-laws. We shared the same bedroom. At night, Bill said something, which I could not let pass unchallenged; “Kentani is the only place, in the Eastern Cape, that has no tortoises.”
“Why?” I asked, thinking that this had to do with the climatic environment.
“Africans ate all of them and left nothing to sustain these animals.”
“No! That is not true!” I protested, because I had a relationship with these people.
“What do you mean, it is not true? Dr.... (/ dont re- member his name) learnt about this when he was inves- tigating the cause of their depletion in this area, years back in the early twenties. I read his book and you can’t dispute it.”
“Well, his assumption was wrong.” I replied, confi- dently, knowing that what he was going to hear would shock him. “First of all; it 1s very, very difficult for a stranger to get information from traditional amaXhosa, because these people are known for their scepticism of strangers. If you ask them anything, they will ask, “why do you want to know?’ After that they will not share with you their knowledge; more so if you’re a stranger. In the past amaXhosa trained their children from an early age never to tell a stranger the truth, especially to white
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people.”
Bill listened quietly, without interrupting.
I continued, “Now let me tell you something that they did not tell Dr.....? AmaXhosa did not eat any creeping or crawling animals, like centipedes, lizards, snakes, crabs, frogs, locusts, ants or tortoises, hence they looked down upon Khoisan people, because of their “repulsive” diet. The Khoisans ate these animals.
Even though they were converted into Christianity, there are still some Xhosa households who hunt and kill or keep tortoises for their strong religious or cultural be- liefs. They generally believe that if they burn a tortoise Shell in a kraal with cattle, the cattle will multiply. Cat- tle are a status symbol or a bank to our people. Tortoise shells are also used as troughs to store drinking water for chickens so that they may increase. There is also a belief that if live tortoises are kept in a household, they will repel evil spirits. These beliefs surely must have been the cause of depletion of these animals in an area as conser- vative and traditional as Centane. I was told that because of their scarcity, locals are prepared to purchase and im- port them from other areas.”
“It makes sense,” Bill said and kept quiet for a long time afterwards.
Working with Bill had a very strong impact on me. He was very dedicated and committed in whatever he was doing. In Matatiele, he went out into the night to search for frogs in the river while it was raining and thundering. He didn’t allow anything to stand in his way. Many years later, after I had left P. E. [Port Elizabeth] Museum, I went on a personal and voluntary excursion of document- ing and taking pictures of bushmen paintings in the caves in the Nkonkobe and Chris Hani Municipalities. Without his basic research training I wouldn’t have embarked on this project. Bill gave me a hands-on experience in re- searching and I thank him for sharing his skill with me.
a J i} rT) f L —_, 4 (Ee » ee eS | < = ws age / \\ ‘ 3 J N
Fig. 16. Bill Branch discussing the finer points of the day’s photographic record of collections with colleagues and students, Lagoa Carumbo, May 2012. (Photo: Brian Huntley).
Amphib. Reptile Conserv.
Brian J. Huntley
South Africa
During 2009 and 2012 I had the pleasure of introducing Bill Branch and a few dozen other field biologists to the diversity of life in the deserts, montane grasslands, miombo woodlands, forests, and floodplains of the far reaches of Angola. Bill soon proved to be the hardest working and most convivial member of the teams, which comprised up to 30 biologists from ten countries.
During the first expedition to southwestern Angola, in January 2009, we camped out on the Humpata highlands and in the Namib desert. Here Bill and fellow herpetologists found multiple new records and several new species of frogs and lizards. From the faint light of dawn to the pitch darkness of night Bill would be in the field or at the makeshift laboratory tent, where a generous donation of Cuca beer from the local brewery kept spirits and laughter levels high. What impressed me most about Bill was his ability to inspire all around him— young students to ageing professors—game rangers to army generals—with fascinating stories about his cold- blooded friends.
In May 2012, when we were camped out along a Congo tributary at Lagoa Carumbo, in the far northeast of Angola, the evening’s discussions around the campfire ranged from Bill’s erudite interpretation of current species concepts to scary personal experiences of snake bites and the treatment thereof. We were eight hours drive from the closest town, and another five hours from the closest doctor. One afternoon, Bill had been out to set a trap for a black mamba that had been seen slithering down a hole on arock face. He casually told us how he had, that same morning, pulled what he thought was a harmless water snake out of the Luele River. Only when he returned to camp did he discover that it was a new elapid record for Angola — Banded Water Cobra, Naja annulata.
We had planned to visit Portugal together early in 2018 to discuss collaborative projects with colleagues at the University of Porto, but at a meeting in Cape Town that January, Bill informed me of the advice his doctor had given him that week: he should not travel. We soon learnt of the severity of his illness, but this did not slow Bill down. He was already under heavy pressure to complete his catalogue of Snakes of Angola, but did not hesitate to honour his promise of a chapter on reptiles for the synthesis volume that I was coordinating on Biodiversity of Angola. We kept up a lively correspondence to the end, his sharp wit never failing. Fittingly, given Bill’s tremendous role in inspiring young researchers in Angola, the synthesis volume includes a dedication to him.
Roger Bills
South African Institute for Aquatic Biodiversity, South Africa
I first met Bill in Marromeu, central Mozambique. We were part of a team lead by Jonathan Timberlake looking at the biota of the lower Zambezi’s delta region. I had
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driven up from Grahamstown with a bakkie and trailer, and had slept in the car over several nights due to poor road conditions and slow progress. Basically, I was exhausted. This did not get me any respite from Bill’s sharp humour and I had to quickly shape up.
For me the trip was tremendous—there was water everywhere despite it being the dry season, and an abundance of fishes and the fauna was mostly new to me. For most of the other zoologists 1t was not the best season and consequently a bit frustrating. Bill and I got on from the start and we spent time out in the dry fields with a large bulldozer that was flattening termite mounds looking for snakes and lungfishes and several days on a boat going down the Zambezi.
The boat trip down the Zambezi was supposed to be an overnight affair—down the Zambezi to the mouth, a channel through mangroves to one of the delta’s southern braids and up to the small village of Malingapanzi. Unfortunately we missed the tide and left late, and went down river on an incoming tide. It took us the whole day to get down to the mouth where we camped at a fishing village overnight. We expected to get going at first light but the local fishermen stole our rudder as they wanted payment for camping. It took our Mozambique counterparts the whole morning of negotiating and refitting the rudder before we could leave. The time however was well spent: Bill went fishing (he was a good angler) and caught our only Glossogobius giuris for the trip, and I caught a load of mud-skippers in the mangrove flats. Our delay meant we missed the tide again and going up the southern channel to Malingapanzi was against the outgoing tide. We got there late on the second day—Bill had caught one puff adder. He wasn’t very happy and did not return by boat the following day.
From all my experiences with Bill, the impressive thing about him was his resourcefulness in the field, whether collecting by himself or soliciting samples from locals, he managed to get incredible numbers of samples. Returning to camps in the evenings would invariably find Bill at a table covered with specimens that he would be fixing, photographing and taking tissue samples from. He
Fig. 17. Bill and Anton Bok at the Kalumbila Mine Camp, Mwinilunga District, North-West Province, Zambia, May 2010 (Photo: Roger Bills).
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spent long hours doing this work. On one trip to a sand mining project near Pebane, Mozambique, we fell afoul of this. Bill had been there the week before and the locals were used to giving reptiles they had caught to passing vehicles. On our drive from the airstrip to the exploration camp we were oblivious of this. After the second snake came through the window in a flimsy plastic bag, we wound up our windows and did not stop anymore!
Bill was an incredible intellect, a world-class scientist but far more importantly a great guy. It was a privilege to have spent time with him, my life 1s richer for it.
Johan Marais
African Snakebite Institute, South Africa
Back in 1980, while I was curator of Transvaal Snake Park, I met Bill during one of his visits but we barely spoke. I was a youngster cleaning snake cages and Bill was visiting Rod Patterson and Anthony Bannister. We often corresponded and I supplied Bill with a bunch of photographs for his field guide, but it was only in early 2000, on a field trip to Namibia, that we really bonded.
We did several field trips to Namibia, often with Aaron Bauer, but our trips to Niassa in northern Mozambique, Mulanje Mountain in Malawi, and southern Angola were memorable. Field trips are special as there 1s ample time to chat, especially when driving long distances. I particularly enjoyed the chats with both Aaron and Bill, and although there were endless topics discussed it was largely about reptiles. I often wound Bill up about photographing reptiles on inappropriate props like fruit and flowers that were out of place, and he accused me of taking rather poor photographs as I had a bad eye.
His wry sense of humour brightened things up on those long journeys and he was particularly good at irritating Aaron, not to mention times when he would lose specimens while photographing them! My best Bill moment: when an American missionary’s wife in Nampula asked Bill what he does for a living, he responded that he was a reptile scientist who did field work, described recently discovered reptiles, and wrote scientific papers about his discoveries. She responded:
i Po, . i —
Fig. 18. Johan Marais and Bill Branch with a Rock Monitor in Namibia. (Photo: Jackie Childers).
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“Yes but what is your real job?’
It is hard to grasp the gap that Bill has left behind, and so many of us miss the times that we could call or drop him an E-mail. He was notoriously bad at responding to E-mails so I got into the habit of numbering my questions. Needless to say, Bill would only answer those he felt like answering.
Mark-Oliver Rédel
Museum fiir Naturkunde, Germany
My first contact with Bill was in 1996. He asked for a copy of my frog book, and invited me to give a talk on West African amphibians and reptiles on the third World Congress of Herpetology in Prague, where I met him for the first time in person. Bill was organizing a session to summarize the progress in African herpetology. Thus, it was Bill ‘officially’ introducing me, my Ph.D. not yet finished, to the community of African herpetologists. We kept contact thereafter, but it took a few years until we met again.
Following a workshop to define conservation priorities for West Africa, Conservation International started a series of rapid biodiversity assessments in little known areas across the Upper Guinea forests. In early 2002, Bill and I were asked to participate on one of these RAPs, targeting the Haute Dodo and Cavally Forest reserves in western Céte d’Ivoire. He was responsible for the reptiles and I was to focus on amphibians, but of course we conducted all field work together, recorded many interesting amphibian and reptile species, ignored all CI safety rules, and had a lot of fun catching animals and talking rubbish. For Bill it was his first time being in West Africa, and his first time working in rainforests (as a ‘typical’ South African he showed up in shorts and it took me quite a bit to convince him that working in a rainforest in shorts is a very stupid idea).
Not all of the experiences were fun. In one night in the Cavally forest, we walked far from camp and encountered a few rarer species we hadn’t seen before on the trip. On our way back, we stumbled straight into the largest raid of army ants (Dorylus sp.) I ever encountered! The forest floor and all lower parts of the shrubs and trees were covered with these aggressive insects, and in seconds the ants where everywhere on and under our clothes. We just ran to leave them behind, and then had to strip naked to pull off hundreds of ants, all holding onto the skin they had successfully penetrated with their sharp mandibles. It was only when we finally finished them all off (one has to pinch off the heads of every single one) and turned again towards the campsite, that Bill realized that he had lost his glasses. We had to turn back into the ants to search for them..... A much more pleasant experience on that trip was when we found the first live caecilian, Geotrypetes seraphini, Bill had ever seen.
Caecilians were also one of the most spectacular findings, actually the first country record for the entire group, the next time we met. In the fall of 2003, Bill
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invited Johan Marais and I to survey amphibians and reptiles in the Niassa Game Reserve, in northern Mozambique. Although it was the core dry season it was an extremely successful survey, revealing 57 reptile and 31 amphibian species, including a new Cordylus, and further potentially undescribed species including the Scolecomorphus mentioned above.
Thereafter we met regularly, mostly in South Africa, but a few times in Germany as well. Bill often took me on shorter excursions across southern Africa, e.g., showing me spectacular parts of the Cape Fold Mountains or the Karoo, and I frequently visited him and his wife Donvé in their amazing house and garden in Port Elisabeth. There we had long and entertaining discussions about herpetology, science, politics, or sports, while sipping on a nice glass of wine, observing the many birds in the garden, or following a soccer or rugby match on television. We never agreed on which soccer team or player was worth supporting, and I could always bet that I would receive a derisive email after a German defeat against an English team in the Champions League. Bill was mad about some sports and missing an important rugby match was impossible, even on an excursion. Particularly memorable was when we once drove through the Karoo and he wanted to listen to a match on the radio. As the radio quality was weak, we had to finally stop and follow the broadcasting on the roadside in the desert. However, the only program Bill could find was in Afrikaans. Thus apart from the players’ names and the score, he did not understand a single word. An amazing fact about Bill was that, although he was a forceful speaker, loving to use and to play with the English language, he was completely ignorant about other languages. So he never learned Afrikaans and in other countries, other people had to cope with translations.
But Bill had encyclopedic knowledge of the natural sciences in general, and he could instantly give a lecture about southern African zoology, botany, or geology. He was easily connecting all this different knowledge into a broader, comprehensive framework and thereby developing new questions and ideas. This ability to communicate new or complex knowledge made him a very stimulating academic teacher, something which was certainly was one of the reasons why he was so popular on the National Geographic tours he was guiding in his later years. His non-protective way of openly sharing data and ideas, as well as critically and without any mercy dissecting project ideas, hopefully remains a model to all the many students and colleagues with whom he was communicating his entire life. Many of his ideas and projects now remain to be finished by others, most prominently the revision of the ‘bible’ Bill Branch’s Field Guide to Snakes and other Reptiles of Southern Africa, and the description of dozens of new reptile species he had already collected and deposited in the herpetological collection of the Port Elizabeth museum.
To me, Bill was much more than a good colleague,
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Fig. 19. Bill Branch in a sad mood after his snake stick, proudly stolen from Aaron Bauer, broke while he tried to destroy an Opuntia in the Karoo, October 2012. Bill: “What an
embarrassing death to a snake stick, killed by a plant” (Photo: Mark-Oliver Rédel).
although we did only meet occasionally. More often in recent years, he was a very good friend with whom I enjoyed discussing everything, not only science. However, the scientific discussions with him were a constant inspiration providing me with many, sometimes unusual ideas on how to interpret data or set up new projects. He introduced me to the African family of herpetologists and to Mozambique; and I am proud that I could introduce him to the West African herpetofauna and rainforests, and even convinced him (sometimes) that amphibians are not completely boring. For him, I would have even loved to see England take the World Cup in 2018. He died too early and in an unbelievably
Fig. 21. Bill Benen a Mark- Oliver Rodel wan the halite of the species named in their honour (Photo: Frank Tillack).
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, & |
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Fig. 20. Bill Branch and Mark-Oliver Rédel in July 2018 in Bill’s home in Port Elizabeth (Photo: Mark-Oliver Rédel).
cruel way. I am very happy that I could meet him one last time, shortly before his death in PE. He will always remain an unforgettable person and inspiration. His death is a great personal and scientific loss, and my thoughts are with his beloved wife Donvé and both their families.
Paul H. Skelton Wild Bird Trust, National Geographic Okavango Wilderness Project When the National Geographic Okavango Wilderness Project (NGOWP) was looking for key specialists to join them on expeditions into the unexplored highlands of Angola, Dr. Bill Branch was a first port of call. Bill was attracted into the NGOWP as an established authority of Southern African and Angolan herpetology, most especially the reptiles. He joined the founding 2015 NGOWP Expedition as part of the 'land party.' He also took part in the 2016 expedition, joining it after first enjoying an extensive journey through the escarpment reaches of Angola in the company of Dr. Pedro Vaz Pinto. Prior to this, he had visited Angola on a number of occasions, collecting and adding significantly to the herpetofaunal knowledge of the country. His collecting antics often drew curious onlookers, mostly children, who would marvel at what wonders he would bring forth from the ponds, rocks, and crevices. More significantly he both encouraged and actively mentored younger hepetologists currently active in Angola. These expeditions have resulted in several potential new species, a number of new species for Angola, and range extensions of many others.
Bill was an old friend and colleague of some of us. I personally met and knew Bill soon after he arrived in South Africa, and was working for the Atomic Energy Corporation outside Pretoria. On joiming the Port Elizabeth Museum, we became good friends and he
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Pee j | Fah fia ar * ite) 1 abit apis 2. Fig. 22. Bill oe in — (Scene for frogs (Photo: Alex Paullin).
joined me on at least two interesting expeditions that I arranged primarily for fish sampling—one was along the Lower Orange River, and the other was to Lesotho in 1988. Bill had a'nose' for reptiles. I remember vividly how he would relish the refreshment breaks on the expedition as we travelled north across the karoo, in order to sniff around the rocky kopjes and outcrops along the way. He inevitably returned with a clutch of reptiles in hand, all of which were unerringly shot with an elastic band from the hand. And then there was the spitting cobra, Naja nigricollis woodii, he caught by the tail on crossing the road, after screeching to a halt and leaping forth from the vehicle. As viewed from the vehicle, behind it was an energetic spectacle in madness—born out on arrival to realise that Bill had been spat in the eyes by the enraged reptile and, whilst he was blinded and in agony, was desperately directing his non-herpetological colleagues in the niceties of bringing a canvass bag to bear so he could insert the writhing beast. Needless to say, he succeeded and managed to wash his eyes out before he was permanently damaged. On the Lesotho trip, Bill was his amazing self and not only displayed his fly-fishing skills that I never knew he had, but also showed me the cryptic, super-jawed, Maluti River Frog (Amieta vertebralis) in its natural habitat. His calm demeanour, bubbling humour, and all-round knowledge in the field was always refreshing. Simply put, Bill was a pleasure to have around. His scientific productivity and achievements are of a top order. His passing was a great loss to our project and to the community at large.
Aaron M. Bauer
Villanova University, USA
I met Bill in 1987, during my first trip to South Africa. I had met with Alan Channing in San Francisco and he had given me a list of all of the critical herpetologists and institutions to visit in South Africa. After visits to Wulf Haacke at the then Transvaal Museum and to FitzSimons Snake Park in Durban, I made my way to Port Elizabeth via Cradock. I phoned Bill on the way (from a post office, remember no cell phones?) and he suggested that my field assistant and I stay at the camping ground on
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Fig, 23. Bill Branch in Angola bean for ean: (Photo: Alex Paullin).
Brookes Hill. A gale was blowing and we were soaked to the bone, but the next morning Bill kindly showed me around the Museum complex. Over the next day or two he took me Bradypodion hunting in Happy Valley, just down Beach Road from the Museum, showed me the introduced Lygodactylus capensis on the guard rails along the roads, and sent me off to Schoenmakerskop to look for Acontias meleagris, Homoroselaps lacteus, and other reptiles. Like everyone I met on that first trip, Bill was a critical contact if I was intending to start working in South Africa. By 1989, I was coming regularly, sometime
Fig. 24, Bill Branch in Angola with a dead on the road Vine Snake (Photo: Alex Paullin).
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Fig. 25. “Uncle Bill” enjoying the adoration of the masses (Marius Burger and Krystal Tolley) at the H.A.A. meeting in Cape Town in 2011 (Photo: Aaron Bauer).
two or three times a year, and more often than not Bill and I would go to the field together, starting a 30 year personal and professional collaboration that influenced all of my work in Africa as a whole.
I have very many fond memories of Bill. One was a 1990 trip to northern Namibia. I picked up Bill and drove with him and two of my students to a farm in Kamanjab. We stayed with the farm managers and had a wonderful time. The collecting was spectacular and mostly new to both me and Bill, who had not spent much time in Namibia before this. Every day we found additional species, in the end nearly 50 species on the farm alone, and more between Kamanjab and Palmwag. Bill had to leave before me and on his last night, after weeks of the best warthog and gemsbok, we were promised “something special,” which turned out to be a very old and very gamey goat! The next day Bill and I left the students and drove straight through to P.E. with only a short stop for a nap. Our only music in the car was The Greatest Hits of Elton John. On the trip we really got to know one another, and we both got so sick of Elton John that we couldn’t listen to his music for years.
Other fond memories are of our multi-year projects in the Little Karoo and later the Richtersveld. In those days there were very few visitors to the Richtersveld, and Bill and I both enjoyed the solitude of the park, evenings by the fire along the Orange River, and finding two Bitis xeropaga only meters away from one another. I can also mention a magical trip to the Kaokoveld along with Johan Marais and my Villanova colleague, Todd Jackman. We were in the bed of the Munutum River and all of a sudden we were surrounded by a herd of giraffe. Even Bill, always ready for the good photo opportunity, was temporarily awestruck by the scene. I also spent many memorable weeks with Bill in the States. One trip was to the South Carolina coast just after a hurricane. Despite some serious close calls with disaster, the loss of one of Bill’s cameras, and hundreds of mosquito bites, Bill was pleased to catch a baby alligator and to have
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had the chance to be in the field with Whit Gibbons, a great herpetologist and ecologist, and an author whose writings Bill admired. On another trip, we drove 10,000 km from coast to coast and back in the US with my students and postdocs. At 3,700 m we saw a herd of elk and Bill managed to get most of his body outside of our moving van to get the perfect shot. I think all of these fond memories are united by the common theme of sharing with Bill the feeling of how lucky we are to have a vocation we love and that lets us enjoy spectacular animals in amazing places in the company of our friends.
Bill was the face of South African herpetology, indeed of African herpetology. His interests were wide-ranging and he had a mind for details when it came to all things herpetological. He was also a master naturalist who knew his birds and his plants, as well as the history of natural history exploration in Africa. He was also down-to-earth. Even the most novice of herpetologists was welcome to call him Bill, not Dr. Branch. Although he could, and often did, go on for hours about something in a quite serious tone, anyone who spent much time with Bill knew that he had a wicked sense of humor, and conversations with him could swing between debates about the International Code of Zoological Nomenclature one minute to a hectic exchange of friendly insults the next. That he was known as “Uncle Bill” to many speaks volumes about how comfortable we all felt with Bill. My relationship with him was somewhat different. Years ago I told Bill that I thought of him as the older brother I never had and indeed, the last words Bill spoke to me were “Be well, little brother.” We are all both better herpetololgists and better people for having known Bill.
Stephen Spawls
Although Bill and I had corresponded since the early 1980’s, we didn’t meet until 1987, when Bill drove up to Botswana and stayed a few days with us at Moeding College, Otse. It was an exciting visit. Bill came in a white windowless Volkswagen Kombi, which was the same type of vehicle that had been used by the South Africa Defence Force on their 1985 raid into Botswana. Consequently, the Botswana security forces had tracked the vehicle, and as Bill drove out of our college he was stopped by the soldiers, who went through the vehicle. Finding nothing, the military concluded that Bill had cached his weapons at my house which was then searched! After this inauspicious start, my wife and I subsequently stayed with Bill in Port Elizabeth. We went on an amazing safari, to the Addo Elephant National Park, to Graaff-Reinet, and thence into the Karroo, where we stayed at the Karroo National Park headquarters with Bill’s friend, the warden Harold Braack. We returned via the Swartberg and Oudtshoorn.
Being in the field with Bill in some of his favourite country was an amazing experience; he knew the land, the customs, and the animals, and gave freely of his expertise. We found many spectacular species that were totally
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new to me, including Hewitt’s Ghost Frog, Heleophryne hewitti, the Giant Ground Gecko, Chondrodactylus angulifer, and the Blue-spotted Girdled Lizard, Ninurta coeruleopunctatus. But we weren’t lucky all the time. At one point Bill and I drove for several hours at night on the tarred road near Beaufort West, hoping to find a Horned Adder, Bitis caudalis, but we saw virtually nothing. As we returned to the park, near midnight, there on the road was a snake, and we leapt out with great excitement but it turned out to be only a Herald Snake, Crotaphopeltis hotamboeia. One morning Bill and I were pursuing a sand lizard, Pedioplanis, which was sheltering 1n a clump of bush. As it appeared near my feet I dove at it but missed, and it fled to another clump. Bill clapped his hands on his head. ‘It’s obvious you’ve always collected by yourself’, he said exasperatedly, ‘you should have just shuffled it towards me, not leapt at it without telling me.’
On that trip, we also learnt of each other’s shared enthusiasm for bird-watching. As we drove across country, Bill directed me to a side road. ‘I’ve got a surprise for you,’ he told me, as we took the diversion. We went a few miles and then Bill told me to pull up and get my binoculars; and there, in a grassy area below the road, were a pair of Blue Cranes, the first I had ever seen.
In 1991, I went to work in Ethiopia, and Bill wrote to me in 1992, suggesting we might work together on a book on Africa’s dangerous snakes. Blandford Press showed interest in the project, and in 1993 Bill came and spent a few weeks with us in Ethiopia, doing field work and working on the book. We made several field trips, one was to the highlands east of the Rift Valley, to a town called Dodolla where a specimen of Bitis parviocula, the spectacular Ethiopian Mountain Viper, had been collected, still the only specimen known from east of the rift valley. As we ascended the rift valley wall, up through dense broad-leafed forest, we became increasingly excited; this looked like Bitis parviocula country. Then as we approached Dodolla, we emerged on the plateau, and found ourselves on a vast open grassland, as bald as a billiard table. Bill sighed and looked at me. ‘Listen, matey’ he said (Bill and I were both born in North London, he at Finsbury Park, me in Muswell Hill, and sometimes in the field we were just two Londoners together), ‘we’re looking for a forest viper, and as far as habitat goes, we’ve just gone from the sublime to the ridiculous.’ But that day we did find some spectacular frogs, including Paracassina kounhiensis, Mocquard’s Mountain Kassina.
The following week, down in Awash National Park, we had some remarkable luck; in one afternoon and evening we got a North-east African Carpet Viper (Echis pyramidum) under a rock right outside our room, on the road in the dark we found a Kenya Sand Boa and two species of egg-eater, and as we drove back to the lodge, we caught a huge Atractaspis fallax on the road, an adventure that Bill described as being ‘like trying to subdue a spiked manhole cover.’ At Lake Langano, we
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caught a small Egyptian Cobra (Naja haje) on the road. Back in Addis Ababa, I was teaching one morning and Bill worked in the garden, creating a small set on top of a rather nice garden table to photograph the cobra. Unfortunately, he incorporated several biggish rocks in the set and in moving these around, he managed to thoroughly gouge the polished surface of the table. My wife went ballistic, but Bill turned on the charm and managed to persuade her that it was all part of the great scientific endeavour, and he took us out for a meal as well. The following day I found Bill crawling around in the canna lilies when I got home; one of the frogs he was photographing had sprung into the flowerbed and escaped.
We didn’t always get on well. Bill had a very relaxed attitude towards deadlines, and often preferred to go into the field rather than knuckle down. He once told me how his publishers (Struik) ‘had flown him to Cape Town’ to finish his field guide, and a fellow herpetologist, who overheard this, said ‘What you mean, Bill, is that Struik made you fly there, sat you down in their offices and said you weren’t leaving until you got it finished.’ Bill laughed and admitted it; and in one of his books he thanks his editor for ‘making ridiculous deadlines seem acceptable.’ Our work on the Dangerous Snakes of Africa book was complicated. Bill was in South Africa, I was in Ethiopia, and there was no e-mail in those days. We used to send stuff to each other by courier. As the deadline for the delivery of the manuscript approached, Bill had a lot of the snakebite stuff still to do and wasn’t getting it done. With two weeks to go and the publishers muttering angrily about penalty clauses (the production was catalogued, and tied into a publicity/release schedule), I rang Port Elizabeth to be told that ‘Dr. Branch had left on an extended safari to Zambia, and would not be back for a few weeks.’ In a panic, I managed to get hold of Dr. Colin Tilbury, who stepped into the breach and wrote virtually all the snakebite stuff in short order. The manuscript went in on time, but it led to a furious row between Bill and I over the order of our names on the cover. But eventually we got over it, and in fact in 2017, we agreed to do a revision of the Dangerous Snakes book.
The last time I met Bill was in 2014, in Bagamoyo, Tanzania, where we were part of a team assessing Tanzania’s reptile biodiversity. We went out one evening and I climbed a tree to catch a sleeping Spotted Bush Snake. Bill watched thoughtfully. ‘I’m past climbing trees’, he told me. ‘In fact, ’'m past climbing over anything. Last time I was in Namibia with Aaron Bauer, a lizard ran under quite a low fence and neither Aaron nor I could get over it.’ He laughed, ‘It’s my fondness for prawn curry.’ On that conference, Bill talked with great enthusiasm of a projected book. ‘I really want to do a big book’ he told me, ‘covering the natural history of Africa’s snakes, along the lines of Harry Greene’s book.’ He showed me some ideas and pictures on his computer; his ideas were mind-stretching and holistic; he saw
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Fig. 26. Bill Branch photographing some lilies at Lake Langanao, Ethiopia (Photo: Steven Spaw!ls).
the snakes in the landscape as part of the interlocking whole ecosystem, and his accompanying pictures were, as always with Bill’s photographs, spectacular. Nobody else has photographed the African herpetofauna like Bill.
We started work on the revision of the Dangerous Snakes book in early 2017. Bill sent me some draft material, a list of important references (he had an encyclopaedic knowledge of the literature on African reptiles), and a stunning portfolio of pictures. But in late 2017, Bill cautiously wrote to tell me he was having mobility issues, and wasn’t sure how this might affect our project. And in early 2018, to my shock, I heard from Bill that he had been diagnosed with motor neurone disease. But he remained full of optimism; and said that he fully intended to do his bit; cheerfully pointing out that Stephen Hawking had lasted many years with the same affliction. But it was not to be. The disease quickly took hold. Tragically, Bill died on the 14th October 2018. His untimely death is a major loss to African herpetology. And I hope that our book, which should be published in mid-2020, will be a suitable monument to Bill. Few herpetologists have reached both the public and their fellow scientists with such verve and accuracy as Bill did.
Ss
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Fig. 28. Bill Branch and others at Bagamoyo, Tanzania, in 2014.
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Fig. 27. Bill Branch admiring an old tank near Dodolla, Ethiopia (Photo: Steven Spawls).
Andrew Turner
CapeNature, Western Cape, South Africa
I first met Bill Branch at an HAA meeting, I think the Stellenbosch meeting of 1998 or thereabouts. He was the top South African herpetologist in my mind because of his comprehensive treatment of the reptiles of the region (his lesser interest in amphibians did not bother me, as his emphasis on the snakes more than made up for this). He was always interested in other people’s experiences, especially regarding observations on distributional occurrence, and encouraged documenting this valuable form of data. Knowing that someone like Bill, who came from a rather different background, could switch to making a career of herpetology—and a rather exciting and enjoyable career at that—was a great inspiration for me to continue my professional herpetological interest. His photography was also inspirational and keeps me (and many others) clicking away.
Bill was a great raconteur, and his stories of herping gone wrong were particularly amusing. One story in particular, although I don’t remember that exact mechanics of it, involved Bill trying to catch a Cordylus under a rock that was being lifted by a colleague. (I shall not mention his name but he did have an extensive snake collection at one point). Bill shoots his right hand under the rock to catch the Cordylus but notices at the same time there is a second Cordylus under the rock so shoots in his left hand too, catching both of them. But then a third Cordylus runs out from under the rock and said colleague catches that one by dropping the rock on both Bills hands!
Bill really set the scene for getting the full picture of South African herpetological diversity, and did a good job of placing this in an African and global context. He travelled widely and shared his great photographs, and was always wondering how the various species fitted together. His fondness for the small adders was totally understandable, and he did a good job of discerning their subtle (and probably recent) divergence and highlighting the need for conservation of several of these species.
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Fig. 29. Bill doing what he did best: geeing everyone up to maximise income from the HAA auction! (Photo: Andrew Turner).
Ernst H.W. Baard
CapeNature, Western Cape, South Africa
My career as a herpetologist with CapeNature (then Cape Department of Nature and Environmental Conservation) started in January 1983. My first task was to sort out and process a few thousand specimens of frogs, lizards, snakes, and tortoises collected and collated by my predecessor, John Comrie Greig (Greig and Burdett, 1976. Patterns in the Distribution of Southern African Terrestrial Tortoises) and, at the time, colleagues, Richard Boycott and Atherton de Villiers.
After writing (yes, there were no emails those years) to the curator of the Port Elizabeth Museum, Dr. Bill Branch, and the curator of herpetology at the South African Museum in Cape Town, Dr. Geoff McLachlan, about depositing the specimens (roughly divided into the Western and Eastern Cape), we got positive responses from both curators. Atherton and myself proceeded, and we completed the task of sending, among others, the whole Greig and Burdet wet and dry tortoise collection, and several hundred “eastern” Cape lizards, snakes, and frogs to Bill at the Port Elizabeth Museum. Bill’s epic paper on the lizards of the Cape Province (Branch, William. 1981. An Annotated Checklist of the Lizards of the Cape Province) made a huge impact on my knowledge of the lizards of the Cape, and together with FitzSimons 1943 (Lizards of South Africa) guided us through the process.
An incident that stood out during this time, was the discovery in the Jonkershoek collection of an un- identified many-spotted lizard in a small bottle, collected in 1973. It was beautifully preserved and, fortunately, with a geographical location in the Groot Winterhoek Mounatins down to seconds South and East (this was before GPS). It took me a few days using FitzSimons 1943 to identify the lizard as “Lacerta” australis, and we were very excited about this discovery. One unsuccessful collection trip to the locality (by Boycott, De Villiers, and Baard) was undertaken in February 1983, followed by Atherton and I managing to collect two more specimens
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at the same locality in April 1983. Imagine our joy, since these were as far as we could establish, only the 6th and 7th specimens known to herpetology of this “elusive” species. This information was shared with Bill and he encouraged us to publish a note which we promptly did, asking him to co-author (De Villiers, Baard and Branch. 1983. ‘Lacerta’ australis: additional material). Bill was always supportive of any further investigations and readily responded to queries on the herpetological collection.
In later years, Bill got back to us and was very excited about some of the tortoise specimens we sent him, including some of the largest individuals of some species he had encountered. He then published a short note in the Journal at the time, honouring Atherton and I with co- authorship (Branch, Baard and De Villiers. 1990. Some exceptionally large Southern African chelonians).
I only met Bill for the first time at my first HAA Conference in Stellenbosch a year or two later, and was really honoured to make his acquaintance. His paper on angulate tortoise ecology in the Eastern Cape (Branch 1984. Preliminary observations on the ecology of the Angulate Tortoise) had a huge impact on my career, since this paper shaped my thoughts and guided my research and attempts at understanding the ecology of the geometric tortoise of the Western Cape; having completed my research in 1990. For a young herpetologist like me at the time, it was almost natural to think: What would Bill do in this case? or How would Bill approach this topic?
Bill’s astonishing knowledge of lizards, tortoises and snakes, snake venom, snakebite, etc. was really something to behold, and few herpetologists could keep up with him. I fondly remember Bill at conferences communicating with all and sharing his knowledge. The best story I remember him telling, was about the evening in the veld around the fire. After Marius Burger latched a Pseudocordylus crag lizard to his (Marius’) earlobe, Bill kept on touching the lizard which wouldn’t let go of Marius’ ear; with the lizard biting down harder and harder, Bill spent an hour or so enjoying Marius’ agony and futile attempts to get the lizard to let go of his ear!
William R. (Bill) Branch was a legend of his generation and time. Not only was he a brilliant scientist, excellent herpetologist, and I believe, a great bird enthusiast, but also somebody one could look up to. His contribution to South African and global herpetology will go down in history as exceptional, ground-breaking, and outstanding, and will stand the test of time like with other greats; FitzSimons, Broadley, etc. His contribution to the written and peer-reviewed herpetological science and popular literature is unsurpassed, and it is my honour to have known him.
Amber Jackson
Cape Town, South Africa
Uncle Bill’s Bible was a well-used field guide by the time I met the man himself. I had even sent him a few
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Fig. 30. DNA sampling of Spek’s Hinged Tortoise, Kinixys spekii (Photo: Amber Jackson).
specimens of Leptotyphlops years before when I was a student. I finally met Bill as an awe struck herpie requesting he sign my copy of his field guide before a very serious meeting at my new place of work. He wrote: “Wow, Uncle Bills bible!!! 167 species out of date, but what else is there.” Out of nowhere he then spouted a lecture about the Galapagos and island biogeography, and held up the meeting for 20 minutes in the process. All eyes on me, I left with my signed copy and some knowledge I never requested but was all the better for knowing. Little did I know, our first meeting was an accurate precursor for the years that followed. Thanks to numerous development EIA’s Bill and I travelled to Lesotho, Augrabies, and Mozambique (multiple times), with me always as his self-proclaimed assistant.
One of my favourite memories with Bill is lying in the dark, on a rocky shelf at the top of the Augrabies paleo falls, staring at the stars and waiting for the geckos to come back out after our disturbance. The stars were incredible! I later caught him a Pachydactulus atorquatus, without breaking the skin, and received an exclamation of ‘I could kiss you.’ He didn’t, and ran off with his prize. At the time, I was naively more excited that 1t meant we could go to bed before our dawn wakeup call in four hours. Bill, 40 years my senior, put me to shame with his energy levels.
We got along at first because I was eager to learn, and he was eager to teach. Then one day, I called him a bastard for one of the anti-feminist comments he used to purposefully provoke me with, to which he laughed,
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Fig. 31. Bill reading science in 45 °C heat (Photo: Amber Jackson).
sighed, and said “finally!” All pretenses over with, we were then friends.
Most of my time with Bill was spent learning, so much so that my brain stopped being able to absorb any more information and saturated by the end of the long day. The knowledge he possessed was impressive, diverse, and felt insurmountable. He taught me plenty about herpetology and science in general. His enthusiasm was contagious and witnessing his studiousness in the field was impressive, with his daily diary and specimen processing. But one thing that stands out is the things he probably never intended to teach me, for example: How being reserved doesn’t have to impact negatively on your life, how euthanizing something as cute as a bush baby can make a huge contribution to science and someone’s career, or how LED lights can make fresh produce more appealing. How just because your life starts in one place doesn’t mean you have to stay there. How you can be a jack of all trades and a master of one. How you can make mistakes. That apologies are important. How careers can be diverse and often unfold. That one of the biggest joys is to love, freely and openly.
To the man that “was by turns (and somehow all at once) relaxed, intense, sincere, self-mocking, modest, confident, serious, and funny.” Kim Stanley Robinson
To the man that could make an economist understand biodiversity by using economic terms.
To the man that could answer the question: “Is a penguin a fish or a bird?” politely and honestly.
To the man that could provoke an entire lecture with a simple question: ‘What’s the odd one out?’
To the man that believed in love, in science and in the unknown.
To the man that studied cancer but fell in love with a cobra while fishing.
To kind, funny, and sometimes forgetful ‘Bum in the Butter’ Bill.
I think about you often, your teachings, your adventures,
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Fig. 32. Bill with a shoftshell terrapin he caught (Photo: Craig Weatherby).
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our last day together, and all the days before then. I couldn’t have asked for a better mentor, teacher, and dearest friend. I will treasure you always. Thank you for believing in me.
Margaretha Hofmeyr
University of the Western Cape, South Africa
I first met Bill Branch in October 2000, when my colleague Alan Channing invited him along on a field trip to Namibia for the UWC Zoology Honours students. I had most students with me in my husband’s Kombi Synchro, while Bill was a passenger in Alan’s Land Rover. We stopped at Springbok’s Caravan Park for the first night, where I booked accommodation in chalets. Because Bill joined the party at the last minute, I knew there would not be a bed for him and had some concerns about sleeping arrangements. For one or other reason, perhaps because Bill was idolised by all herpetologists, I expected him to be rather arrogant, but he quickly won me over when he made a bed for himself in the trailer Alan took along. The trailer was quite short, but so was Bill; fortunately the trailer was rather wide, because so was Bill. The sight of him surfacing the next morning from his trailer bed will always stay with me. Yes, he might have been arrogant at times, but he was always a great sport, and teased the students to distraction on this trip.
I always feel dishonest calling myself a herpetologist, because my field of expertise is restricted to tortoises and terrapins. Yet, on this trip, as herpetologists do, we went on night drives to look for herps (never tortoises) on the roads. One of the nights while staying at Klein Aus, everybody squeezed into my Kombi to search for exciting things on the roads. While driving through a narrow stretch of road between two fences, a springbok ram materialised in the road before my husband’s car. I switched the main lights off within seconds, but it was still too late. The springbok ran straight into the Kombi, broke his neck, and put quite a dent into the front of the car. All the girls were crying but we had to deal with the situation. Alan and Bill dragged the springbok out of the road and then we had the unfortunate task of driving to the owner’s house to report the incident. His only
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reaction was that it was the only springbok he had on the farm. This was an unpleasant experience for all of us but also created a bond, because Bill mentioned it many times to me in ensuing years.
At conferences, I would get annoyed with fellow herpetologists for teasing Bill about his lisp, yet, he always laughed at their jokes. To me, the ability to laugh at yourself reflects true character, and Bill had that. I have many fond memories of Bill and always regarded him as the ultimate herpetologist and naturalist in South Africa. His expertise stretched so much wider than reptiles and amphibians. He may not have been an expert on every animal or herp group, but his knowledge was astounding. He was also willing to share his expertise and helped many young scientists to find their way. I may not be described as a young scientist, but when I switched from large mammals to tortoises, Bill knew much more than I did, and he was willing to share. Over the years we co- authored several papers and it was always a pleasure to work with him in a professional capacity. South Africa, Africa and the World are now deprived of one of their top intellectuals, and an exceptional person—we salute you Bill.
Jens Reissig
Ultimate Creatures, Gauteng, South Africa
The first time I met Bill was during a high school field trip to northern KwaZulu-Natal around the year 2000. At that stage, I was rather shy and having had a very keen interest in reptiles since my early childhood I of course knew exactly who he was, however never made any contact with him. Many years had passed until I crossed paths with him again at the Herpetological Association of Africa’s Conference at the National Zoological Gardens in Pretoria, South Africa. From this point on, we stayed in contact and he was always willing to help wherever he could. Unfortunately, he was extremely busy while I was compiling my book on the Girdled Lizards and their Relatives in 2013 and 2014, so that he was not able to assist me with it in any way. He did however end up reviewing the book for me. The book review ended up being published in Herpetological Review, 2015, 46(2): 1-7.
After having received the tragic news of Bill’s diagnosis, I decided to go and visit him at his home in Port Elizabeth on the 20th of April 2018. Even though one could see that he was battling his illness, he still tried to be upbeat about life and could not stop talking about reptiles and a (to me) hidden passion of his, Orchids. We sat for hours on his patio talking about various reptile projects, his Orchid collection, birding, and some of his many field trips into Africa. After spending quite some time with us, one could see that he was tiring and we decided to say our goodbyes and I left. It was the best day I had ever spent with him. My favourite email I have ever received from him was received on the 12th of June 2015 and stated: “Dear Jens. Here are the proofs of my review of your excellent book which should appear in
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.—
Fig. 33. 2017 Herpetological Association of Africa’s Conference in Bonamanzi, South Africa. From left to right: Werner Conradie, Tyrone Ping, Prof. William Branch, Luke Verburet, Dr. Michael Bates, Johan Marais, Prof. Graham Alexander, Prof. Aaron Bauer, Jens Reissig, Coleen Tiedemann, Dr. Colin Tilbury, and Dr. Victor Loehr (Photo: Andre Coetzer).
Herp Review this month. Hope you're happy with it and all goes well. Best wishes. Bill”
Professor W.R. Branch, your passing has left a massive hole in so many people’s lives and in African Herpetology as a whole. Your knowledge and sense of humour will be greatly missed by anyone who ever had the privilege to know you and who’s life you may have affected in some way or another. Africa has lost two great herpetologists way too soon and in relatively quick succession. Till we meet again!
Harold Braack
South Africa
Bill Branch first came into my life, I think, in 1974, at an HAA conference held at Skukuza in the Kruger National Park (KNP). It might have been the first such get-together. At that time, I was doing the herpetology survey of the KNP, so it was indeed fortuitous to meet Bill Branch, as well as Don Broadley and Carl Gans.
In 1976, I was transferred to the Bontebok National Park and so I started species surveys and checklists. I wanted to know what it was that I was supposed to be looking for, and so Bill and I started working together through various National Parks and adjacent areas. But it was not only herps. We looked at succulent plants and birds as well.
What Bill gave me was the confidence to do the surveys. In him, I had a partner with whom I could share my passion for conservation and protection of all the inhabitants of those areas. He was totally enthusiastic and this rubbed off on us all. Above all, he was a good friend.
I have many fond memories of Bill, but share only a few here.
The pepper ticks at Addo National Park were a vast irritation to Bill. We picked them up every time we ventured on a collecting trip. He hated them. I wiped them off with paraffin. But Bill had a different solution.
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“Oh no,” he said. “I go home. I get undressed completely then lie down in the nude on the kitchen counter. My family has to pick them off. “
Bill Duellman and Bill Branch stayed with us in the Karoo for several days. To separate the two, we called Duellman “Bill” and Branch “Billikins.” Bill B. was not totally enamoured with the solution.
Bill and I did a night road survey in the Richtersveld. After several hours we arrived at Paradyskloof. We lay down flat on our backs for a while counting passing satellites, then later scratching among the rocks where we saw the largest Hadogenes that either of us had seen. Then Bill went to the little pool to find a Strongylopus springbokensis. Ka-splash, splash. Bill fell in the pond. He sat huddled in the bakkie on the way home.
Spending a long, long time trying to catch a lizard in Richetrsveld, Bill suddenly ran back to the bakkie. Out he came with a revolver loaded with dust shot. “Ka- boom!” he shot the thing—we had our specimen.
He had excellent repartee and a quick lucid mind. How many of us remember his response during a frog meeting at Stellenbosch? The chair said we should be democratic in the course of the meeting. Bill’s immediate response “Thank You, Mr. Mugabe.”
Bill also enjoyed fishing, especially for carp. We spent some time along the banks of the Orange and Breede Rivers doing just that. Didn’t catch much, but those were relaxing times.
I best remember Bill as a man who was a dear friend. To all of us, he revealed the treasure chest of our vast herpetological wealth—and, more, he opened it up for us to see and explore. He followed his passion with a radiant glee which he passed on to us.
Bill, my friend, I salute you for being a friend, a guide, and an explorer who found and revealed.
Atherton de Villiers
CapeNature, Western Cape, South Africa
I have good memories of Bill that date back to when his career 1n herpetology started at Port Elizabeth Museum, and have always admired his enthusiasm and vast knowledge of reptiles and amphibians. It is well known that one of his greatest achievements was his Field Guide to the Snakes and other Reptiles of Southern Africa. This landmark publication opened up the world of reptiles to countless numbers of people, and it was a pleasure to contribute information and images for one of the most important herpetological publications in southern Africa. I share with you all the huge loss of Bill to herpetology, biodiversity conservation, and life in general.
Marius Burger
North-West University, South Africa
Try as I may, I just can’t seem to pinpoint the precise memory of actually meeting Bill for the first time. ?'m quite shocked by this realisation. I presume that it was sometime during 1987 when I was a young (20 y/o)
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Fig. 34. Ozzy Osbourne meets Elvis Presley. My all-time favourite photo of Bill and I (Photo: John Measey).
nature conservation student in Grahamstown, and I vaguely remember something about visiting him at his office at Port Elizabeth Museum (PEM). The only definite lead that I have to go on is a specimen of Karoo (ex-Namaqua) Plated Lizard (Gerrhosaurus typicus) that I collected in November 1987 on the Karoo Nature Reserve in Graaff-Reinet. At the time, this record represented an eastern distribution range extension of 211 km. Whoopee! I would have hurriedly taken the specimen to Bill at PEM, with my tail wagging in excitement. Yes, what a joy it always was to make some sort of new herpetological discovery that even The Bill Branch would find somewhat noteworthy. And so it came to pass that my first ‘scientific publication’ was a short note in the Journal of the Herpetological Association of Africa (Burger 1988). The truth be known, Bill actually wrote the damn thing. But this was my official introduction to the Herpetological Association of Africa, and it marks the approximate start of a very lekker 30- year friendship with Bill.
An article published in Zootaxa on 24 October 2018 demonstrated that the African Slender-snouted Crocodile (Mecistops cataphractus) is in fact comprised of two superficially cryptic species, and thus M. leptorhynchus from Central Africa was resurrected as a valid species (Shirley et al. 2018). The first thought that crossed my mind when I read this paper was “Fok, Bill didnt get to see this!”, because Bill had died ten days earlier. Bill would have loved the news that Mecistops is monotypic no more, and perhaps (probably) he even knew that this was in the pipeline. Fast-forward nine months to July 2019 (i.e., right now as I’m writing this), and I’m still experiencing Where-TF-is-Bill moments on an almost daily basis. Bill was my Google Herps. Whenever I needed photos of far-flung African reptiles to be identified, the oh-so-convenient Google Herps would
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Fig. 35. Bill emerging from swamp in Gabon (2002) after checking funnel traps that he had set with the hope of catching
an African Parachanna (Photo: Marius Burger).
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usually be my first check. When the complexities of taxonomy would bewilder my brain, Bill had the knack of explaining it in a way that I could sort of comprehend. And so for me, it is not only very sad, but also utterly inconvenient and totally kak that Bill died.
It always intrigued me that a Pom could arrive in Africa with some sort of medical doctorate degree, something to do with foetal rabbit liver metabolism and primary liver cancer, only to end up perusing a career of chasing reptiles and amphibians. Like, how the hell did that happen?! Anyway, it’s a good thing that it turned out the way it did. Well, so says I, because I have derived much joy and intellectual enrichment from the times hanging out with Bill. To South Africans, Bill was loved and respected as our local herp guru. He was of course also internationally renowned for his herpetological contributions, and the momentum that he built up over the decades will have him publishing papers for a long while after he clocked out.
Fig. 36. Bill receiving medical attention during a biodiversity survey of Loango National Park (2002). If I remember correctly, it had something to do with removing ticks from a place where the sun don’t shine (Photo: Carlton Ward Jr.).
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This may be somewhat of a narcissistic trait of mine, but I like it that Bill liked me. I would often purposefully say and do socially objectionable stuff in Bill’s presence, for the reward of his approval and appreciation of my crudeness. Now that I think about it, this kind of behaviour was probably akin to a son showing off in front of his father for attention and approval. In a book that collated a collection of interviews on how to become a herpetologist (Li Vigni 2013), Bill wrote the following: “T also never miss a chance to be with Marius Burger, just to re-emphasize how sane I am.” Whenever Bill acknowledged me in a publication, I would smile, and feel all warm and gushy inside for receiving his praises. In the acknowledgments section of Tortoises, Terrapins & Turtles of Africa (Branch 2008), Bill wrote: “A special thanks to Marius Burger, whose tortoise photography is Just too good.” Well, just imagine the grin on my face for that bit of flattery. He then went on to say: “...ifhe could only look after his camera lenses as well as his hair...” in The Dangerous Snakes of Africa (Spawls and Branch 1995), and Bill included a thanks to a certain Marias (sic) Birger (sic) for companionship and advice. If that was indeed me that he was referring to, then I say ditto to that.
Whilst on a fieldtrip with Olivier Pauwels and Bill Branch in Gabon, the three of us shared a shipping container that was modified into a bedroom of sorts. I retired to bed late one evening, with Olivier and Bill giggling away like preteen girls. The reason for their hysterics was that they had planted a condom half-filled with Condensed Milk in my bed. How silly 1s that! Anyway, I never noticed said condom in my bed and managed to fall asleep in spite of the spurts and snorts of laughter. The next day whilst checking our trap arrays they told me of their really funny prank, all the while grinning from ear to ear as they awaited my reaction. Instead of shock and dismay, I replied with a calm reminder that a cleaning team was making our beds each morning and just imagine what their take would be on discovering this soggy item in one of our beds. I watched as their expressions gradually turned from smile to mild alarm, as the two of them slowly processed and realised the gravity of this scenario. Now it was my turn to laugh.
I’m not a spiritual kind of guy, and thus I won’t be saying things like RIP. old friend or check you on the other side. But ja, Bill was for sure a significant component of my life. Iam very chuffed to have had him as a friend.
Mike Raath
Director, Port Elizabeth Museum Complex (now Bayworld), 1987-1995
I first met Bill Branch in the early 1980s, when I was at Wits University as head of the Bernard Price Institute for Palaeontological Research. I had been invited by Prof. Brian Allanson of Rhodes University’s Zoology Department to present a short course on the evolution of the Class Reptilia at my much loved Alma Mater.
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At that early point in my career, I only knew of Bill by reputation, and had never met him personally. I felt flattered that he had taken the time and trouble to travel from Port Elizabeth to Grahamstown to listen to my ramblings. I little realised then that he and I would meet up again in a different context several years later, when I was fortunate enough to be appointed Director of the Port Elizabeth Museum Complex, as it was then called before it got its trendy current name of “Bayworld.’
Bill was a much-respected member of the research staff of the Complex, having charge of one of the most comprehensive and important herpetological collections in the country, building on the solid legacy of its original founder, the legendary F.W. FitzSimons, almost a century before. He was one of the stars of our research team, regularly producing work that was published in some of the world’s top peer-reviewed scientific journals. But in addition, he was a prolific writer of popular articles and books aimed at the general reader that spread his expert knowledge to a much wider general readership. I remember one envious member of our research staff calling him “the Naas Botha of our research team” in terms of earning brownie-points for research output (only those who know something about South African rugby in the 1980s will understand that comment! ).
One of the things that defined Bill was his off-the- wall unconventionality. I remember how audiences at his various public presentations would shudder in shock, horror, and jaw-dropping disbelief when he demonstrated his go-to technique for distinguishing between identical sibling species of toads—by licking them! And, by Jove, it worked!
I respected Bill as a person and as a scientist. But to be candid, I have to say that he and I did not get on that well personally. He suffered neither fools nor administrators gladly, so as his director I guess I failed on both counts! But as a scientist committed to his discipline there was no faulting him. He was single-mindedly devoted to his collection and his research, often to be found in his lab or office over weekends or public holidays when most others on the staff were taking what they rightly regarded as a well-earned rest.
One particular Saturday morning remains starkly and darkly etched in my memory, when Bill received an urgent call in his lab mid-morning from the Snake Park. One of the Snake Handlers, Mr. Nimrod Mkalipi, had been bitten by a Puff Adder at the end of the daily live snake demonstration, and he was in dire distress. Bill dropped everything and rushed to Nimrod’s side, administering antivenom and applying appropriate emergency first aid. Tragically, though, it was to no avail and Nimrod succumbed on the scene despite Bill’s urgent, expert, and devoted efforts. Medical opinion afterwards held that it was anaphylactic shock that took Nimrod’s life, and that nothing other than immediate on-site specialised medical intervention would have had any chance of preventing it. That event shocked us all. It is a dark memory that I
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I si ee ee ee Fig. 37. Bill walking off into the early morning light to photograph some Welwitschias in south-western Angola in 2009 (Photo: Werner Conradie).
carry with me to this day, but I applaud Bill Branch for his swift reaction and his valiant and urgent attempts to save the life of a fellow staff member. We all had much to learn from that tragedy.
Werner Conradie Port Elizabeth Museum, South Africa The first time I became aware of Bill was when I attended my very first HAA conference at Bayworld in 2004, which he organised. I remember the occasion very clearly, as the only options available at the icebreaker were beer, and Coca-Cola. I was too shy to speak to him then—he was the famous Bill Branch and I was, after all, just a lowly student. I met Bill again at the 2006 HAA held at Potchefstoom, and this time I was on the organising committee. At this event I recall fondly Bill’s talk on ‘guts and gonads,’ which of course went well over its allotted time. However, it wasn’t until I finished university completely that we would have what would turn out to be a bit of a prophetic chance encounter. While on a December break, just before I would start a new job as a Physical Science high school teacher, me and my now wife walked into him while strolling around the museum. I introduced myself to him, finally having a moment in his direct eyesight after all this time, and with a curt nod paired with a brief “nice to meet you,” he disappeared through the door to his lab and office. Never would I have guessed that less than six months after this, I will be sitting in front of him for an interview for the job of Assistant Herpetologist. I must have impressed Bill one way or the other (couldn’t have been pure desperation), as two days later I received a call that I got the job. I finished my contract at the school as fast as I could, and with great excitement walked straight into the museum the very same day. Bill looked at me in utter shock and sent me home, saying I should come back in the New Year... I guess he wasn’t prepared for my quick start!
For the first year at the museum I had to learn all the ropes. Now this is a very steep uphill battle for
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Fig. 38. Bill holding a Meller's Chameleon (7rioceros melleri), looking radiant despite a very tiring hike during the summit of Mt. Mabu in 2009 (Photo: Werner Conradie).
an Afrikaans-speaking seun that knew his frogs, but no reptiles. Up to that point in time, the only reptile ever caught by me was a harmless Common Brown Water Snake. The first challenge I faced was getting into a conversation with Bill. Because of his British heritage, he mumbled a lot and this made me struggle to understand his pronunciation of scientific names. After many “conversations,” I often went back to my office and paged through his field guide to prepare for the next engagement. Bill, however, never forced his reptilian inclinations on me. Once, he walked into my office and promptly asked me what I wanted to specialise in, to which my response was tadpoles. He dryly remarked that the only thing they are good for is fish bait. As it turns out, I never did work on tadpoles that much...
I went into the field with Bill for the first time as part of a multi-collaborative expedition to Angola in 2009. Bill didn’t have to bring me along, he could have kept all the new places and specimens to himself, but I will ever be grateful as it was on that trip that I fully came to understand and realise what my responsibilities as a museum herpetologist include: New discoveries! I joined Bill on two more consecutive trips to Mount Mabu, Mozambique in 2009 and Lagoa Carumbo, Angola in 2011. It was at this stage that Bill assisted me with my first-ever species description, and just as I was starting to bask in the glow of his knowledge, he clearly thought he had trained me enough and turned off the light. It seemed I was on my own: Bill expected me to swim. It was up to
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me to show him that I could. We wouldn’t go on another field trip together again until 2015, again to Angola. By this time, Bill had retired, and I was keeping the fort on my own. During the trip, around the campfire one evening, Bill told me that he can now rest in peace, knowing the Port Elizabeth Museum herpetology collection is in good hands. Thank you, Bill.
I worked with Bill for more than ten years, but I only really started to get to truly know him and his family when he was unfortunately diagnosed with motor neuron disease. It was a devastating experience to see your mentor and friend fade away in front of your eyes. Bill was determined to follow his passion to the very end, and his determination was amazing to behold. Bill has taught me life lessons that I will cherish forever. He was truly a one-of-a-kind man. He is and will be missed.
Martin J. Whiting Department of Biological Sciences, University, Sydney, Australia I first met Bill in 1994, just as I was about to start my Ph.D. working on flat lizards (Platysaurus). I was based at the Transvaal Museum in Pretoria, and he arrived to Slice up snakes as part of a project on the ecology of African snakes with Rick Shine, Jonathan Webb, and Peter Harlow. Shortly after meeting him, he told me about the Augrabies flat lizard system, which ended up being the subject of my Ph.D. and many happy field seasons. I owed him a huge debt, without realising it at the time! And as it turned out, our discussions about flat lizards led to a collaboration that continued until his death. Everyone that meets Bill is immediately struck by how warm and caring he is. It’s hard to describe, but he had a personality that immediately drew you in. And I think that’s why he had such an impact on so many people. He was particularly giving and helpful to young aspiring herpetologists, and I very much appreciated his friendship and advice as a young Ph.D. student fresh on the herpetological scene in South Africa. A few years into my Ph.D., he invited me on a field trip to a remote area of northern Mozambique to survey the vertebrates of the Moebase region, the site of a proposed titanium mine (sadly). I really appreciated this gesture, because he could have invited any number of far more qualified people! His son Tom was also on the trip, to survey birds. Little did I know that this would become such a memorable trip, and that I would have experiences I still talk about to this day. There is nothing like a field trip to really get to know someone, and that trip forged a lifelong friendship. With Bill, there was never a shortage of stimulating conversation on a wide range of topics beyond herpetology. His love for natural history was
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infectious. The only thing Bill spoke about with more passion was his wife Donvé. While on that same field trip to Mozambique, Bill set out to find a clay pot that was representative of the region, to take back to Donveé. I should mention that Donvé is an award winning potter, so this was the perfect gift! Our fixer couldn’t quite understand what a westerner would want with a clay pot, but we examined quite a few, before buying one from a surprised local villager. Actually, I also acquired one which has survived multiple moves in South Africa and a final move to Australia. (How could I not buy one after hearing Bill wax on about Donveé and her pottery!)
Bill was larger than life and made a huge impact on African herpetology. It’s hard to accept that he’s gone, but he will never be forgotten. He will certainly be missed by many. I am currently working on finishing a phylogeny and revision of the Platysaurus with Scott Keogh and Mitzy Pepper, a project that Bill and the late Don Broadley were both involved in, and he will certainly be in our thoughts as we put together the final touches.
Fig. 39. Bill and his son Tom while passing through a village during our 1997 Mozambique trip (scanned from a slide). To this day, that field trip ranks as one of my most memorable (Photo: Martin Whiting).
egeerie BA ee io ee at ie Fig. 40. Bill in action during our 1997 Mozambique trip [scanned from slides] (Photo: Martin Whiting).
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Literature Cited
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Li Vigni F (Editor). 2013. A Life for Reptiles and Amphibians, Volume 1. Chimaira, Frankfurt, Germany. 495 p.
Branch B. 2008. Tortoises, Terrapins & Turtles of Africa. | Shirley MH, Carr AN, Nestler JH, Vliet KA, Brochu Struik Publishers, Cape Town, South Africa. 128 p. CA. 2018. Systematic revision of the living African
Burger M. 1988. Geographical distribution: Gerrhosaurus Slender-snouted Crocodiles (Mecistops Gray, 1844). typicus. Journal of the Herpetological Association of Zootaxa 4504(2): 151-193.
Africa 35: 36.
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Spawls S, Branch B. 1995. The Dangerous Snakes of Africa. Southern Book Publishers, Halfway House, Johannesburg, South Africa. 192 p.
Werner Conradie holds a Masters in Environmental Science (M.Env.Sc.) and has 12 years of experience with the southern African herpetofauna. His main research interests focus on the taxonomy, conservation, and ecology of amphibians and reptiles. Werner has published numerous principal and collaborative scientific papers, and has served on a number of conservation and scientific panels, including the Southern African Reptile and Amphibian Relisting Committees. Werner has undertaken research expeditions to various countries including Angola, Botswana, Lesotho, Malawi, Mozambique, Namibia, South Africa, Zambia, and Zimbabwe. He is currently the Curator of Herpetology at the Port Elizabeth Museum (Bayworld), South Africa.
Michael L. Grieneisen spent much of his childhood searching for and observing herps in the Appalachian Mountains of Pennsylvania, USA. He obtained a B.S. in Biology and Chemistry from Shippensburg University in Pennsylvania, and a Ph.D. in Biology from University of North Carolina, Chapel Hill, on a National Science Foundation graduate fellowship. Mike’s Ph.D. and post-doc work (at University of Nevada, Reno) investigated the hormones that turn caterpillars into butterflies. Over the past 12 years at University of California, Davis, Mike has authored journal articles in fields as diverse as nanotechnology, climate change, biodiversity, scientometrics, environmental science, and reduced-risk pest control practices in California. Mike is a freelance editor, co-editor of Amphibian & Reptile Conservation, and he is compiling metadata for the theses and dissertations on amphibians and reptiles produced worldwide. The compilation currently includes over 54,000 theses, completed from 1803 to the present at institutions in well over 100 countries, and is expected to be made available sometime in 2020. He also has an extensive collection of world banknotes which feature herps in the design.
Craig L. Hassapakis is the Founder, Co-editor, and Publisher of the journal Amphibian & Reptile Conservation (official journal website: amphibian-reptile-conservation.org), which was founded in 1996, and a former editor of FrogLog (www.amphibians.org/froglog/). Craig has been an instructor (first grade through college), non-profit and governmental volunteer at Public Library of Science (PLoS), Co-group Facilitator, Genome Resources Working Group, IUCN/SSC Amphibian Specialist Group (ASG), and is a member of the IUCN/SSC Amphibian Specialist Group. His interests include biodiversity, evolution, systematics, phylogenetics, taxonomy, conservation, and behavior of amphibians and reptiles. Craig is instrumental in developing and establishing “Amphibia Bank: A genome resource cryobank and network for amphibian species worldwide.” His professional memberships include: Society for the Study of Amphibian and Reptiles (SSAR), Herpetologists’ League (HL), International Society for Biological and Environmental Repositories (ISBER), and International Society for the History and Bibliography of Herpetology (ISHBH).
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Amphibian & Reptile Conservation 13(2) [Special Section]: 1-28 (e181).
A herpetological survey of western Zambia Gabriela B. Bittencourt-Silva Department of Life Sciences, Natural History Museum, London, SW7 5BD, UNITED KINGDOM
Abstract.—A list of 60 species of amphibians and reptiles found during a six-week survey in western Zambia is presented. Two species of amphibians are newly reported for Zambia: Amietia chapini and an undescribed species of Tomopterna, previously known to occur in the Democratic Republic of Congo and in Namibia, respectively. Some of the material collected could not be confidently identified to species level because of the taxonomic complexity and uncertainty of some groups (e.g., Phrynobatrachus, Ptychadena), even with the use of DNA barcoding. This list is a small contribution to the growing knowledge of Zambian and African herpetology.
Keywords. Amphibians, barcode, checklist, reptiles, Southern Africa, undescribed species Citation: Bittencourt-Silva GB. 2019. Herpetological survey of western Zambia. Amphibian & Reptile Conservation 13(2) [Special Section]: 1-28 (e181). Copyright: © 2019 Bittencourt-Silva GB. This is an open access article distributed under the terms of the Creative Commons Attribution License [At-
tribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced,
are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org. Submitted: 2 September 2018; Accepted: 4 March 2019; Published: 6 August 2019
Introduction
Zambia is a landlocked southern African country considered part of the Zambesiaca area, which also includes Botswana, Malawi, Mozambique, parts of Namibia (Caprivi), and Zimbabwe (Poynton and Broadley 1991). Zambia is located on the main central African plateau where elevations range from 1,200 m to 1,500 m and the vegetation 1s dominated by miombo woodland (Phiri 2005).
Very little has been published on the herpetofauna of Zambia since the first then comprehensive reports from the early 1900’s (see reviews in Haagner etal. 2000; Pietersen et al. 2017). Poynton and Broadley’s compendium Amphibia Zambesiaca (Poynton and Broadley 1985a,b, 1987, 1988, 1991) and Channing (2001) reported on the distribution of Zambian amphibians, while Broadley (1971) presented an initial treatise of the reptiles and amphibians of Zambia, and Broadley et al. (2003) provided an updated atlas and field guide to the snakes of Zambia. Similarly, Haagner et al. (2000) and more recently Pietersen et al. (2017) have made important contributions to Zambian herpetology. Broadley (1991) presents a comprehensive list of reptiles and amphibians from the Mwinilunga District, northwestern Zambia, including records from museum collections dating from 1957. However, except from the extreme north-west (Hillwood Farm), the herpetofauna of western Zambia remains very poorly studied, with only a few regional checklists (e.g., Broadley 1991; Pietersen et al. 2017).
Correspondence. g. bittencourt@nhm.ac.uk
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Currently, there are 189 species of reptiles recorded for Zambia according to The Reptile Database (Uetz et al. 2018) and 181 (two crocodile, 10 chelonian, 78 lizard, and 91 snake species) according to Pietersen et al. (2017). The number of amphibian species varies substantially according to different sources. Pietersen et al. (2017) report 86 species of amphibians for Zambia, while AmphibiaWeb (2018) reports 87 species, and a search in the Amphibian Species of the World 6.0 database (ASW; Frost 2018) returns 104 species. This disparity between databases is possibly due to the fact that the ASW includes non-confirmed occurrences. An example is the caecilian Boulengerula, which is expected to occur in Zambia based on its known distribution but is as yet unreported there.
Herein I present a checklist of species collected during a six-week herpetological survey in western Zambia.
Materials and Methods Study site and sampling
The survey was carried out in April and May 2014 encompassing protected as well as non-protected areas of western Zambia (Fig. 1, Table 1). Figure 2 shows the different vegetation types surveyed, comprising miombo woodlands (dominated by Brachystegia spp.), dry evergreen forests (dominated by Cryptosepalum sp.), riverine forests (mushito) and grassy wetlands (dambo). During the whole survey period there were only four
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30
sinker mr. 10
Elevation (m) M200 °° -15 Mi 430
|_| 660
890
1120
[) 1350
{| 1580
|_| 1810
|_| 2040
[__] 2270
[| 2500
Fig. 1. Map of Zambia showing sites surveyed for herpetofauna. The star indicates the capital (Lusaka).
days of rain and average temperatures were 30 °C during the day and 15 °C at night. The main sampling methods were acoustic and visual encounter surveys (diurnal and nocturnal). Entomologists participating in the expedition opportunistically collected some specimens with the use of sweep nets and small pitfall traps (500 ml cups). All specimens collected were euthanized with 20% benzocaine (applied on the skin or in the mouth). Samples of thigh muscles were taken and stored in absolute ethanol before the specimens were fixed in 10%
formalin and transferred to 70% industrial methylated spirit for long-term storage. All specimens are deposited in the herpetological collection of the Natural History Museum in London, United Kingdom (see Appendix 1).
Species identification Identification keys for Amphibia (Channing 2001;
Poynton and Broadley 1985a) and Reptilia (Branch 1998; Broadley 1971: Broadley et al. 2003) were used to
Table 1. Localities in Zambia surveyed during this study. Protected areas are indicated by shaded green. NP: National Park; HQ:
Headquarters. Locality District Chavuma FR Chavuma Hillwood Farm Ikelenge Itezhi-Tezhi, Kafue NP Itezhi-Tezhi Lukwakwa Kabompo Livingstone, Maramba Lodge Livingstone Mayukuyuku, Kafue NP Mumbwa Nanzila Plains, Kafue NP Itezhi-Tezhi Ngonye Falls Camp Shangombo Nkwaji Mwinilunga Sioma Ngwezi NP Shangombo Sioma Ngwezi NP (HQ) Shangombo
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Longitude Latitude Elevation (m) -13.07006 22.92880 1070 -11.26316 24.32782 1400 -15.77340 26.01151 1040 -12.66084 24.43697 1150 -17.89120 25.85821 900 -14.91533 26.06311 1010 -16.28138 25.91676 1030 -16.66139 23.57280 930 -11.56567 24.52605 1300 -16.89873 23.59847 1010 -16.66953 23.56743 1000
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Fig. 2. Habitats surveyed in western Zambia. (A) Dambo and Cryptosepalum dry forest in Lukwakwa, (B) Dambo in Nanzila Plains, (C) Miombo woodland in Nanzila Plains, (D) Margin of the Zambezi River at Ngonye Falls.
assist with the identification of specimens. Some species identifications presented here are tentative because some groups have complex and difficult taxonomies (e.g., Hyperolius, Hemisus, Phrynobatrachus, and Ptychadena). Most samples were barcoded to help species identification (see details below). Despite its known limitations (e.g., Deichmann et al. 2017; Hebert and Gregory 2005; Meier et al. 2006), DNA barcoding is generally, and sometimes very, helpful. The Basic Local Alignment Search Tool (BLAST; Altschul et al. 1990) was used to search the GenBank repository and identify the closest matches for each sample. As there are not many 16S rRNA sequences of Zambian reptiles and amphibians openly available for comparison, percentage of sequence similarity presented here should be interpreted with caution and while taking the possibility of geographic isolation or isolation by distance into account. Private databases were also used for sequence comparisons (D. Portik and B. Zimkus). Snakes were identified primarily using morphological characters.
Genetic analysis
Given the large amount of 16S rRNA sequence data available in GenBank for African amphibians and reptiles, this gene was selected for DNA barcoding. Total genomic DNA was extracted using a Qiagen DNeasy kit (Venlo, Netherlands) following the manufacturer’s protocol for purification of total DNA from animal tissues. A fragment (ca. 500 bp) of the 16S rRNA
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mitochondrial gene was amplified using the primers 16S H3062 (CCGGTTTGAACTCAGATCA) and 16SB FROG (CGCCTGTTACCAAAAACAT) [modified from Palumbi et al. 1991]. Polymerase chain reaction (PCR) was performed using Illustra PuReTaq Ready-To-Go PCR Beads (GE Healthcare Life Sciences) for 35 cycles of 1 minute with annealing temperature at 51 °C. Single strand sequencing reactions and electrophoresis were carried out by the molecular lab team at the Natural History Museum in London, United Kingdom. All sequences generated are available in GenBank under accession numbers MK464267—MK464483.
DNA sequences were trimmed in Genelous v.7 (Kearse et al. 2012) with a maximum of low-quality bases of 20. Uncorrected pairwise distances (p-distances) of the 16S sequences were calculated for some groups in PAUP* (Swofford 2001). For Phrynobatrachus, a maximum likelihood (ML) analysis with non- parametric bootstrapping was carried out with RAxML v.8.2 (Stamatakis 2014). The alignment was generated in Geneious using the Auto algorithm of MAFFT v.7 (Katoh et al. 2002), inspected visually and poorly aligned regions were eliminated using the GBlocks Server v.091b (Castresana 2000). Evolutionary models were evaluated using Automated Model Selection (using a Neighbor Joining tree) in PAUP*. The best fitting model (GTR + GAMMA) was selected according to the Akaike Information Criteria (AIC). Tomopterna marmorata was used for rooting.
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Results
A total of 40 species of amphibians (anurans) belonging to nine families and 13 genera, and 20 species of reptiles from nine families and 17 genera (14 lizards, five snakes, one tortoise) were recorded during this survey (Appendix 1). Among the localities surveyed, Hillwood Farm had the highest species diversity (n=23), followed by Nkwaji (n=15). Different from all other areas surveyed, which are characterized by a combination of miombo woodland, dambo and/or dry forest, both localities mentioned above have riverine or swamp forest, locally known as mushitos
(Fig. 2). Species accounts
All collected specimens and their respective vouchers are listed below. Voucher numbers in bold refer to specimens identified solely based on morphology (i.e., no tissue sample available).
Amphibia Order Anura Arthroleptidae
Arthroleptis stenodactylus Pfeffer, 1893
Shovel-footed Squeaker
Material. LUKWAKWA: BMNH 2018.5826, BMNH 2018.5827 (Fig. 3A), BMNH 2018.5828-29; NKWAJI: BMNH 2018.5830. Comments: Found in leaf litter in Cryptosepalum forest, in dambo and at edges of mushito. Arthroleptis stenodactylus as currently understood is widely distributed from Angola to Tanzania, and from Kenya to South Africa. This taxon clearly represents a species complex, possibly two ecologically distinct forms (see comments in Pickersgill 2007). All specimens listed above have white venters without any dark markings, large inner metatarsal tubercles and a dark line on each side running from the snout over the tympanum to the shoulder. Sequence similarity with A. stenodactylus from Malawi is 98% (GenBank accession numbers FJ51098— 99).
Arthroleptis xenochirus Boulenger, 1905
Plain Squeaker
Material. LUKWAKWA: BMNH 2018.5811 (Fig. 3B), BMNH 2018.5812—13; HILLWOOD FARM: BMNH 2018.5814—20; NKWAJI: BMNH 2018.5821-— 25. Comments: Specimens were found in leaf litter in Cryptosepalum and mushito. All specimens have a very small tympanum and relatively large inner metatarsal tubercle (when compared to A. xenodactyloides). The closest match on GenBank (94%) is to A. xenodactyloides from Malawi (FJ151103). There is no sequence of A. xenochirus available for comparison.
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Bufonidae
Schismaderma carens (Smith, 1848)
Red Toad
Material. CHAVUMA FR: BMNH 2018.5729; ITEZHI- TEZHI: BMNH 2018.5724—28 (Fig. 3C). Comments: Specimens were found in miombo woodland. This species 18 widely distributed in Zambia. The BLAST result show 100% sequence similarity with S. carens from South Africa (KF665176, AF220913).
Sclerophrys gutturalis (Power, 1927)
Guttural Toad
Material. CHAVUMA FR: BMNH_ 2018.5703; LIVINGSTONE: BMNH 2018.5702; LUKWAKWA: BMNH 2018.5705—06, BMNH 2018.5707, MAYUKUYUKU: BMNH_ 2018.5701 (Fig. 3D); NKWAJI: BMNH 2018.5704. Comments: This species is found in miombo and Cryptosepalum forest. These specimens lack the typical red infusions on their thighs, although this could be due to preservation. Specimen identification was confirmed using DNA barcoding (100% sequence similarity with AF220876, from Botswana).
Sclerophrys lemairii (Boulenger, 1901)
Yellow Swamp Toad
Material. HILLWOOD FARM: BMNH 2018.5723; LUKWAKWA: BMNH_ 2018.5715—22 (Fig. 3E). Comments: One male was found in a pond at night and eight individuals were found in dambo near the Cryptosepalum forest. About six males were calling during the day and two couples were observed in amplexus. The species exhibits dynamic sexual dichromatism, where males undergo a temporary color change (from dark green to bright yellow), depending on the breeding period; females are reddish, especially the parotid glands (Bittencourt-Silva 2014; Conradie and Bills 2017).
Sclerophrys pusilla (Mertens, 1937)
Southern Flat-backed Toad
Material. ITEZHI-TEZHI: BMNH _ 2018.5709-10; MAYUKUYUKU: BMNH_ 2018.5708 (Fig. 3F); NKWAJI: BMNH — 2018.5711-12. Comments: Specimens identification were confirmed using DNA barcoding (100% sequence similarity) and non- morphometric morphological characters following Poynton et al. (2016). Specimens were found in miombo woodland.
Hemisotidae Hemisus cf. guineensis Cope, 1865 Guinea Snout-burrower
Material. HILLWOOD FARM: BMNH _ 2018.5801 (juvenile); LUKWAKWA: BMNH 2018.5800 (Fig. 3G);
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“te a . »
Fig. 3. Amphibians of western Zambia. (A) Arthroleptis stenodactylus (BMNH 2018.5827), (B) Arthroleptis xenochirus (BMNH 2018.5811), (C) Schismaderma carens (BMNH 2018.5728), (D) Sclerophrys gutturalis (BMNH 2018.5701), (E) Sclerophrys lemairii, (F) Sclerophrys pusilla (BMNH 2018.5708), (G) Hemisus cf. guineensis (BMNH 2018.5800), (H) Hemisus cf. guineensis (BMNH 2018.5799), (1) Hemisus marmoratus (BMNH 2018.5713), (J) Hemisus marmoratus (BMNH 2018.5714), (KX) Hyperolius dartevellei (BMNH 2018.5681), (L) Hyperolius major (BMNH 2018.5675), (M) Hyperolius marginatus (BMNH 2018.5667), (N) Hyperolius nasicus (BMNH 2018.5666), (O) Hyperolius parallelus (BMNH 2018.5689), (P) Hyperolius parallelus (BMNH 2018.5697), (Q) Hyperolius quinquevittatus, (R) Kassina senegalensis.
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SIOMA NGWEZI NP: BMNH 2018.5799 (juvenile; Fig. 3H). Comments: One juvenile (BMNH 2018.5801) was found dead in a pitfall trap set for dung beetles near mushito. One large individual (snout-vent length 42.2 mm) from Lukwakwa was found buried in sandy soil under a log between Cryptosepalum forest and dambo. The specimen from Sioma Ngwezi is a juvenile. The BLAST search shows 95% sequence similarity with H. guineensis from the Republic of the Congo (K Y080117— 19). According to Channing and Broadley (2002), Hemisus barotseensis, which is endemic to western Zambia, differs from H. guineesis and H. marmoratus in body proportions. The presence of a bright yellow vertebral stripe and small yellow spots on the back agree with the description of H. barotseensis and it is possible that these specimens are that species. If confirmed, this would represent an extension both north and south from its current known range.
Hemisus marmoratus (Peters, 1854)
Marbled Snout-burrower
Material. MAYUKUYUKU: BMNH 2018.5713 (Fig. 31); SIOMA NGWEZI NP: BMNH 2018.5714 (Fig. 3J). Comments: Juvenile individuals found at night in sandy soil in miombo woodland. Hemisus marmoratus is widely distributed in sub-Saharan Africa, excluding rainforest, and it mainly inhabits savannahs but can also be found in gallery forests. The BLAST search shows the closest match on GenBank (94%) is H. marmoratus (AY 531831). Table 2 shows the p-distance for both species of Hemisus presented here and highlights a limitation of the use of this measure for species delimitation.
Hyperoliidae
Hyperolius dartevellei Laurent, 1943
Dartevelle's Reed Frog
Material. CHAVUMA FR: BMNH 2018.5681 (Fig. 3K); HILLWOOD FARM: BMNH _ 2018.5683-—84; LUKWAKWA: BMNH — 2018.5682. Comments: Specimens found in miombo woodland and edge of mushito basking on vegetation during the day. According to Channing et al. (2013) the snout profile of H. dartevellei is truncated instead of shark-like or rounded, but none of the specimens listed above have truncated snouts. However, the BLAST results show 98-99% sequence similarity with samples of H. dartevellei from Ikelenge, north-western Zambia (JQ863650, JQ863653, JQ863673, JQ863676—-78, JQ863704, JQ863708, JQ863718, JQ863750, JQ863753—-54, JQ863756—-S9, KY080197, KY080199).
Hyperolius kachalolae Schiotz, 1975
Kachalola Reed Frog
Material. HILLWOOD FARM: BMNH 2018.5676, BMNH 2018.5677-80. Comments: Juvenile specimens collected during the day on vegetation near a stream in
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mushito forest. In life, the overall coloration was green with a faint canthal and dorsolateral line, consisting of small spots. The green color faded after preservation, although the line is still visible. This agrees with the description provided by Schietz (1975). The sequenced individual shows 98% similarity with H. kachalolae from northern Zambia (D. Portik, pers. comm. ).
Hyperolius major Laurent, 1954
Material. HILLWOOD FARM: BMNH 2018.5675 (Fig. 3L). Comments: One male found on top of a leaf (1.5 m from the ground) calling at night. Schietz (1999) reports this as a savannah species from north-western Zambia and eastern Democratic Republic of the Congo (DRC), however, this specimen was found in a forest patch (mushito). The closest matches from DNA barcoding (97%) are H. kuligae and H. langi (D. Portik, pers. comm. ). Schiotz (1999) states that these species found in west and central Africa are very similar in morphology and dorsal pattern, and may be conspecific. In contrast, Kohler et al. (2005) treat H. kuligae as a western and H. langi as an eastern Central African form. However, the color pattern (especially the post orbital marking) differs substantially from Laurent’s description of the type material of H. /angi, whereas this specimen agrees with the morphological description of H. major provided by Schiotz (1999). There is no DNA sequence of H. major available for comparison.
Hyperolius marginatus Peters, 1854
Margined Reed Frog
Material. LUKWAKWA: BMNH 2018.5667 (male; Fig. 3M); NANZILA PLAINS BMNH 2018.5668, BMNH 2018.5674 (juveniles). Comments: Specimens found near ponds in miombo woodland. There is 100% sequence similarity with H. marginatus from Zambia (D. Portik, pers. comm. ).
Hyperolius nasicus Laurent, 1943
Pointed Long Reed Frog
Material. NANZILA PLAINS BMNH _ 2018.5666 (Fig. 3N). Comments: This single specimen was collected during the day while resting on vegetation ca. 1 m above the ground in dambo. The specimen fits the morphological description of the species provided by Channing et al. (2013): when viewed in profile the snout has a shark-like tip, and the first, third, and fifth toes have one phalanx free (or nearly free) of webbing (see Fig. 4), distinguishing it from all the other species. The closest match on GenBank (98-99% sequence similarity) is Hyperolius inyangae (JQ863674, JQ863683-—84), which is only known from the Eastern Highlands of Zimbabwe and Malawi (Channing et al. 2013). Although Channing et al. (2013) provide accession numbers for the genetic material of H. nasicus, no sequences could be found on GenBank under this species name. This could be due to sequence mislabelling and the accession numbers
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Fig. 4. Details of head profile and webbing of Hyperolius nasicus (BMNH 2018.5666). (A) Profile of head, (B) webbing of right foot, and (C) schematic representation of webbing. Scale bar represents 1 mm.
associated with H. invangae may actually be from H. nasicus.
Hyperolius parallelus Giinther, 1858
Angolan Reed Frog
Material. HILLWOOD FARM: BMNH 2018.5687-88, BMNH 2018.5689 (Fig. 30), BMNH 2018.5690-94, BMNH 2018.5695; NKWAJI: BMNH_ 2018.5696, BMNH 2018.5697 (Fig. 3P), BMNH 2018.5698—5700. Comments: Specimens found near ponds in miombo woodland. These specimens show a color variation similar to the alborufus group (see Schiotz 1999). Hyperolius parallelus is a taxonomically complex group due to its considerable color polymorphism. According to the BLAST search, the closest match (98% sequence similarity with JQ513623, JQ513626, and JQ513625) is H. angolensis from Angola (see Conradie et al. 2012; Frost 2018).
Hyperolius quinquevittatus Bocage, 1866
Five-striped Reed Frog
Material. NKWAJI: BMNH 2018.5685—86 (Fig. 3Q). Comments: Juveniles collected during the day while resting on vegetation in mushito. BLAST results show 99% sequence similarity with H. quinquevittatus from Ikelenge, north-western Zambia (GenBank accession number JQ863752).
Kassina senegalensis (Dumeéril and Bibron, 1841) Bubbling Kassina
Material. CHAVUMA FR: BMNH = 2018.5810; HILLWOOD FARM: BMNH 2018.5802-03 (Fig. 3R); NKWAJI: BMNH 2018.5804; SIOMA NGWEZI NP: BMNH 2018.5805—09. Comments: Specimens found in
Table 2. Uncorrected pairwise distances (p-distances) for a fragment of the 16S rRNA gene for Hemisus spp. Distances between
conspecific populations are inside boxes.
Taxon ID 1 2 3 4 5 1 Hemisus marmoratus AY326070 2 Hemisus marmoratus DQ283430 3 Hemisus marmoratus AY531831 4 Hemisus marmoratus KY 176997 5 Hemisus marmoratus AY948749 6 Hemisus marmoratus KX492610 7 Hemisus marmoratus KM509138 8 Hemisus marmoratus KY 176998 9 Hemisus marmoratus BMNH 2018.5713 10 Hemisus marmoratus BMNH 2018.5714 11 Hemisus guineensis KY080117 12 Hemisus guineensis KY080118 13. Hemisus guineensis KY080119 14 Hemisus guineensis KY080120 15 Hemisus cf guineensis BMNH 2018.5799 16 Hemisus cf guineensis BMNH 2018.5800
17. -Hemisus cf guineensis BMNH 2018.5801
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93
100
97
100
Namibia (GU457565)
100 | Lindi, Tanzania (KY177049) Beira, Mozambique (DQ022361)
Tomopterna marmorata: Zambia (BMNH_2018.5792)
0.05
Ethiopia (FJ829292) Ethiopia (FJ829296) Ethiopia (FJ829295) Ethiopia (FJ829297) Uganda (FJ829298) Ghana (FJ769126) Ghana (FJ769127) Céte d'Ivoire (GU457566) Guinea (EU718726)
Sioma Ngwezi NP (BMNH_2018.5864) Sioma Ngwezi NP (BMNH_2018.5865) Ngonye Falls Camp (BMNH_2018.5836) Ngonye Falls Camp (BMNH_2018.5855) Mtunzini, South Africa (DQ019605)* South Africa (DQ347303)
of western Zambia
Itezhi-Tezhi, Kafue NP (BMNH_2018.5848) Itezhi-Tezhi, Kafue NP (BMNH_2018.5849) Itezhi-Tezhi, Kafue NP (BMNH_2018.5850) Nanzila Plains, Kafue NP (BMNH_2018.5851) Nanzila Plains, Kafue NP (BMNH_2018.5852) Mayukuyuku, Mayukuyuku, Mayukuyuku, Mayukuyuku, Mayukuyuku, Mayukuyuku, Mayukuyuku, Maramba Lodge, Livingnstone (BMNH_2018.5853) Maramba Lodge, Livingnstone (BMNH_2018.5854) Lake Malawi Park, Malawi (FJ889462)
Kakamega Forest, Kenya (FJ889464)
Kakamega Forest, Kenya (FJ889463)
Tatanda Village, Tanzania (DQ283414)
Northern Territory, Rwanda (FJ829290)
Northern Territory, Rwanda (FJ829291)
Kigoma Region, Tanzania (FJ829299)
Tanzania (FJ829289)
lringa, Tanzania (FJ829293)
Kakamega Forest, Kenya (FJ889463)
Kakamega Forest, Kenya (FJ889464)
Kigoma Region, Tanzania (FJ829300)
Mangochi District, Malawi (FJ889462)
Niassa Game Reserve, Mozambique (FJ829303) Niassa Game Reserve, Mozambique (FJ829301) Niassa Game Reserve, Mozambique (FJ829302) Morogoro Region, Tanzania (FJ829294)
Kafue NP (BMNH_2018.5837) Kafue NP (BMNH_2018.5838) Kafue NP (BMNH_2018.5839) Kafue NP (BMNH_2018.5840) Kafue NP (BMNH_2018.5841) Kafue NP (BMNH_2018.5842) Kafue NP (BMNH_2018.5843)
Haplotype Group A**
Haplotype Group B**
| Sp.2
| Phrynobatrachus natalensis
Nkwaji (BMNH_2018.5863) Nkwaji (BMNH_2018.5858) Nkwaji (BMNH_2018.5860) Hillwood Farm (BMNH_2018.5866) Nkwaji (BMNH_2018.5856) Nkwaji (BMNH_2018.5859) Nkwaji (BMNH_2018.5871) Nkwaji (BMNH_2018.5872) Nkwaji (BMNH_2018.5869) Nkwaji (BMNH_2018.5868) Hillwood Farm (BMNH_2018.5867) Nkwaji (BMNH_2018.5870)
Sp.3
Fig. 5. Maximum likelihood phylogenetic tree inferred from nucleotide sequence data from mitochondrial 16S rRNA of Phrynobatrachus natalensis. Numbers above branches are non-parametric bootstrap support values. Specimen vouchers or GenBank accession numbers are shown in parentheses. Colored polygons highlight the clades comprising specimens from this study. (*) Nearest sample from type locality of Phrynobatrachus natalensis; (**) Haplotype groups A and B in Zimkus and Schick (2010).
miombo woodland near temporary ponds. All sequences closely match K. senegalensis (98%, GenBank accession number AF215445).
Phrynobatrachidae (see Table 3 for inter- and intra- specific p-distances; Fig. 5 shows maximum likelihood tree for this group)
Phrynobatrachus cf. parvulus (Boulenger, 1905)
Small Puddle Frog
Material. HILLWOOD FARM: BMNH 2018.5873-78; LUKWAKWA: BMNH 2018.5889 (Fig. 5A); NKWAJI: BMNH 2018.5882 (juvenile), BMNH 2018.5879-80, BMNH 2018.5883—88. Comments: All specimens were found during the day in dambo. Males have a dark throat (BMNH 2018.5882, BMNH 2018.5886—87). While the specimens of P. mababiensis listed below have the venter immaculate (creamy), these specimens have the venter white with dark speckles. Additionally, these specimens show a more well-defined band on the thigh (which runs from knee to knee) and, in most specimens, a light
Amphib. Reptile Conserv.
line runs along the tibia-fibula and thigh (parallel to the band) and joins a vertebral line above the vent (see Fig. 6A). According to Du Preez and Carruthers (2017), the presence of the latter feature distinguishes P. parvulus from P. mababiensis. However, this character is present on both species and therefore cannot be used to separate them (see Poynton and Broadley 1985a. Pietersen et al. (2017) report P. parvulus for Ngonye Falls, approximately 25 km from Sioma Ngwesi NP, but unfortunately there is no voucher specimen. Poynton and Broadley (1985a) and Marques et al. (2018) provide discussions of the literature on the identifications of P. mababiensis and P. parvulus. The barcode is very inconclusive given that the closest hits on GenBank (92-95%) include samples of an unidentified species of Phrynobatrachus from Gabon (KP247505), one from the Republic of the Congo (KY080354), and P. keniensis (JX564885) and P. scheffleri (FJ889479), both from Kenya. Poynton and Broadley (1985a) suggest P. parvulus tends to be associated more with upland and forest conditions than
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Table 3. Uncorrected pairwise distances (p-distances) for the 16S rRNA gene for Phrynobatrachus spp. Distances between conspecific populations are inside boxes. Dotted-line box indicates P. natalensis group.
1 Phrynobatrachus cf. parvulus BMNH 2018.5873
1
2
3
2 Phrynobatrachus cf. parvulus BMNH 2018.5874 0.01
3 Phrynobatrachus cf. parvulus BMNH 2018.5880 0.04 0.05 - 4 Phrynobatrachus mababiensis FJ889461 0.11 O12 0.12 5 Phrynobatrachus mababiensis BMNH 2018.5831 0.11 O11 0.12 6 Phrynobatrachus mababiensis BMNH 2018.5832 0.11 O11 0.12 7 Phrynobatrachus natalensis DQO019605 0.14 O15 0.14 8 Phrynobatrachus natalensis BMNH 2018.5855 0.14 O15 0.14 9 Phrynobatrachus sp. 1 BMNH 2018.5838 0.16 O17 0.17 10 Phrynobatrachus sp. 1 BMNH 2018.5839 0.16 O17 0.17 11 Phrynobatrachus sp. 3 BMNH 2018.5856 0.15 016 0.15 12 Phrynobatrachus sp. 3 BMNH 2018.5858 0.15 016 0.15 13 Phrynobatrachus sp. 2 BMNH 2018.5864 0.17 O17 0.18 14 Phrynobatrachus sp. 2 BMNH 2018.5865 0.17 O17 0.18
P. mababiensis. The localities where these specimens were found are all upland and either inside or near forest, therefore I refer them to P. cf. parvulus. Phrynobatrachus mababiensis FitzSimons, 1932
Dwarf Puddle Frog
Material. MAYUKUYUKU: BMNH 2018.5831-32, BMNH_ 2018.5881; NANZILA PLAINS: BMNH 2018.5833; SIOMA NGWEZI NP: BMNH 2018.5834— 35. Comments: All specimens are juveniles and were found in dambo both during the day and at night. According to Poynton and Broadley (1985a), it 1s not easy to distinguish P. mababiensis from P. parvulus based on external morphology, but they suggest that some characters usually serve to separate them (1.e., labial and subtympanic markings, and shape of tarsal tubercle). The usual well-marked black and white barring on the upper and lower jaws is rather faint on these specimens. Zimkus and Schick (2010) suggest that there 1s cryptic diversity within the P mababiensis group. The closest match on GenBank is P. mababiensis (FJ889461; 99% sequence similarity) from eastern Zambia, which belongs to a population that is sister to the clade containing P. ukingensis and P. ungujae (see Zimkus and Schick 2010).
Phrynobatrachus natalensis (Smith, 1849)
Snoring Puddle Frog
Material. NGONYE FALLS: BMNH _ 2018.5855. Comments: The overall external morphology resembles P. natalensis. The BLAST search shows 98% sequence similarity to P. natalensis from Mtunzini, South Africa (DQ347303), which is near Durban, the type locality of P. natalensis (see Table 3 and Fig. 5).
Phrynobatrachus sp. | Material. I[TEZHI-TEZHI: 50; LIVINGSTONE:
z)
BMNH BMNH
2018.5848— 2018.5853—54;
Amphib. Reptile Conserv.
4 5 6 7 8 9 10 11 12 13 14
0.00 0.00 0.00
0.14 013 013
0.14 013 013
0.17 0.16 0.16}
017 0.16 0.16}
015 014 0.14} 0.09
0.05 0.08 0.08 7
0.05 0.08 0.08 0.00 |
MAYUKUYUKU: BMNH 2018.5837-38 (Fig. 6B), BMNH 2018.5839-42 (Fig. 6C), BMNH 2018.5843, BMNH 2018.5844—-47,; NANZILA PLAINS: BMNH 2018.5851-52. Comments: Specimens were found in dambos. One juvenile was found ona beach of the Zambezi River at Ngonye Falls. The closest match on GenBank (99% sequence similarity) is P. natalensis DQ283414 from Tanzania (see Fig. 5). Zimkus and Schick (2010) suggest that there are two species of P. natalensis in East Africa, and these Zambian populations are more similar to the central and southern populations corresponding to Haplotype group B. It further corresponds to Zimkus et al. (2010) P. natalensis Clade E. See further comments in Discussion.
0.16 015 0153
0.17 0.16 0.16:
0.16 }
0.17 0.16
Phrynobatrachus sp. 2
Material. NGONYE FALLS: BMNH2018.5836; SIOMA NGWEZI NP: BMNH 2018.5864—-65. Comments: Morphologically, these specimens resemble P. natalensis in terms of size, toe webbing and overall color pattern. However, they present silver/white spots around the vent (and ventral part of the thigh in BMNH 2018.5836). The closest match on GenBank is P. natalensis from Tanzania (95%; DQ283414). Although their range overlaps with the Southern African geographic zone (populations A and B) in Zimkus et al. (2010), these populations form a southern Zambian clade, which is a sister group of the eastern and western African clades (see Fig. 5). These findings allude to further cryptic diversity in the group.
Phrynobatrachus sp. 3
Material. HILLWOOD FARM: BMNH 2018.5866—67 (Fig. 6D); NKWAJI: BMNH 2018.5856-57, BMNH 2018.5858-60, BMNH 2018.5861 (Fig. 6E), BMNH 2018.5862-63; BMNH 2018.5868-69 (Fig. 6F), BMNH _ 2018.5870-72. Comments: All specimens
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rs a An «spi ae ie i
: Nee
a
Fig. 6. onan fe western Zaria, (A) re of one (BMNH 2018.5889), (B) Phrynobatrachus natalensis (BMNH 2018.5838), (C) Phrynobatrachus natalensis (BMNH 2018.5842), (D) Phrynobatrachus sp. 1 (BMNH 2018.5867), (E) Phrynobatrachus sp. 1 (BMNH 2018.5861), (F) Phrynobatrachus sp. 1 (BMNH 2018.5869), (G) Xenopus poweri (BMNH 2018.5659), (H) Xenopus pygmaeus, (1) Ptychadena anchietae (BMNH 2018.5730), (J) Ptychadena cf. mossambica (BMNH 2018.5759), (IX) Ptychadena porosissima (BMNH 2018.5766), (L) Ptychadena porosissima (BMNH 2018.5769), (M) Ptychadena porosissima (BMNH 2018.5770), (N) Ptychadena taenioscelis (BMNH 2018.5785), (O) Amietia chapini (BMNH 2018.5664), (P) Pyxicephalus cf. adspersus (BMNH 2018.5791), (Q) Tomopterna sp. (BMNH 2018.5797), (R) Chiromantis xerampelina (BMNH 2018.5798).
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are morphologically similar to P. natalensis, however the BLAST search shows sequence similarity with P. natalensis from South Africa (GenBank accession number DQ347303) varying between 90-91% (see Fig. 5). These populations form a northern Zambian clade, sister to southern, western, and eastern African clades. Further investigation is needed to resolve the taxonomical status of this group.
Pipidae
Xenopus poweri Hewitt, 1927
Power’s Clawed Frog
Material. HILLWOOD FARM: BMNH 2018.5654—58; NKWAJI: BMNH 2018.5659 (Fig. 6G). Comments: Specimens collected from a pond in miombo area. The BLAST search shows 99% sequence similarity with X. poweri from Ikelenge, north-western Zambia (GenBank accession number KP345253). Furman et al. (2015) removed this name from the synonymy of X. petersii and reassigned the West African populations of X. /aevis to this species.
Xenopus pygmaeus Loumont, 1986
Bouchia Clawed Frog
Material. HILLWOOD FARM: BMNH 2018.5651—52 (Fig. 6H), BMNH 2018.5653. Comments: Species identification was based on their relatively small size and the presence of a fourth claw. Specimens were found ina small pool formed in a car track next to a riverine forest (mushito). Recently, Wagner et al. (2013) presented the first record of Xenopus pygmaeus for Zambia, representing a significant range extension (about 1,300 km). This species belongs to the fraseri subgroup and it was previously known to have its southernmost distribution in the northern part of the DRC. The DNA barcode corroborates the morphological identification (99% sequence similarity with KF738291).
Ptychadenidae
Ptychadena anchietae (Bocage, 1868)
Anchieta's Ridged Frog
Material. ITEZHI-TEZHI: BMNH_ 2018.5735-36; MAYUKUYUKU: BMNH 2018.5730-31 (Fig. 61), BMNH = 2018.5732-34. Comments: Specimens collected near water in miombo woodland. The closest match (99%) is P. anchietae (AY517610) from Tanzania.
Ptychadena grandisonae Laurent, 1954
Grandison's Ridged Frog
Material. NK WAJI: BMNH 2018.5737-43. Comments: Two juveniles and four males found in a pond in miombo. This series fits the description for P. grandisonae in Poynton and Broadley (1985a). The results of the BLAST search show that the closest match (98%) is
Amphib. Reptile Conserv.
11
Ptychadena sp. (GenBank accession number KF 178892) from Gabon.
Ptychadena cf. guibei
Material MAYUKUYUKU: BMNH_ § 2018.5764; SIOMA NGWEZI NP: BMNH 2018.5765. Comments: Specimens identified following the key in Poynton and Broadley (1985a). Foot length of BMNH 2018.5765 is slightly less than half the body length and thus, according to the key, this specimen would be mossambica or cotti (i.e. schillukorum). The BLAST result shows 96% sequence similarity with P. porosissima (KY177058) from Kenya. There is no sequence of P. guibei available for comparison.
Ptychadena mapacha Channing, 1993
Mapach Ridged Frog
Material. MAYUKUYUKU: BMNH 2018.5772 (male). Comments: Specimen collected near water in miombo woodland. Following Poynton and Broadley (1985a), this specimen should be assigned to P cotti, now a synonym of P. schillukorum. However, this specimen also fits the description of Ptychadena mapacha (not included in the key), except for the white spots on the posterior face of the tibia and a thin tibial line that are present in the holotype (CAS 160535). The closest match (89%) is P. porosissima (GenBank accession number KY177058) from Kenya, and the sequence similarity with P. schillukorum (KY177060) is 82%. There is no sequence of P. mapacha available for comparison. Although P. schillukorum is \isted in the AmphibiaWeb database as occurring in Zambia, there is no reference to the literature confirming this claim. Pietersen et al. (2017) provided an unconfirmed record of P. mapacha for Sioma Ngwesi. This record represents the northernmost record of this species.
Ptychadena cf. mossambica (Peters, 1854)
Mozambique Ridged Frog
Material. ITEZHI-TEZHI: BMNH 2018.5753, BMNH 2018.5754-57; MAYUKUYUKU: BMNH 2018.5763; SIOMA NGWEZI NP: BMNH 2018.5758—59 (Fig. 6J), BMNH 2018.5760, BMNH 2018.5761. Comments: The key in Poynton and Broadley (1985a) points to P. mossambica, except for the skin folds that are not continuous in these specimens. The authors note that P. mossambica shows an east-west cline in size and degree of webbing, where material from western Zambia tends to be smaller (maximum SVL 28.9 mm) than the series from Mozambique (maximum SVL 52.5 mm). The average SVL of this series is 34 mm. Most specimens have a pair (sometimes two) of large dark blotches on the scapular region. The closest match on GenBank (93-94%) is Ptychadena cf. mossambica from coastal Tanzania (KY177057). These specimens may be referrable to PR mapacha Channing, 1993, for which there is no available sequence data. Ptyvchadena mapacha
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has recently been recorded in Ngonye Falls, south-west Zambia by Pietersen et al. (2017), confirming Channing’s (2001) prediction about its distribution.
Ptychadena nilotica (Seetzen, 1855)
Nile Grass Frog
Material. LIVINGSTONE: BMNH — 2018.5781; NANZILA PLAINS: BMNH 2018.5773-76, BMNH 2018.5777—80. Comments: Specimens collected at night near water in miombo woodland. Sequence similarity with P. nilotica is 98% with KFO27211 and 99% with KX836515 from the DRC. For further discussion about this species see Dehling and Sinsch (2013) and Zimkus et al. (2016).
Ptychadena obscura (Schmidt and Inger, 1959) Material. HILLWOOD FARM: BMNH 2018.5766-67 (Fig. 6K), BMNH 2018.5768. Comments: These small specimens (SVL ranges from 21.8 to 22.8 mm) fit the description of P. obscura in Poynton and Broadley (1985a). The specimens have a pair of dark marks on the scapular region. The results of the BLAST search show higher similarity (97-98%) to P. broadleyi (GenBank accession number MH300600-02). This is an unexpected finding considering that P. broadleyi is only known to occur in the Mulanje Mountain and the Zomba Plateau, in Malawi. These specimens from Zambia have a light triangle on the snout distinguishing them from P. broadleyi. Hence, the barcoding results should be interpreted with caution.
Ptychadena oxyrhynchus (Smith, 1849)
Sharp-nosed Grass Frog
Material. HILLWOOD FARM: BMNH 2018.5783; NANZILA PLAINS: BMNH 2018.5782. Comments: Specimens collected near water in miombo woodland. Sequence similarity with P oxyrhynchus from Kwambonambi, South Africa (GenBank accession number AF215403) is 99%.
Ptychadena porosissima (Steindachner, 1867) Three-striped Grass Frog
Material. HILLWOOD FARM: BMNH 2018.5769 (Fig. 6L), BMNH 2018.5770 (Fig. 6M), BMNH 2018.5771. Comments: This species, common in miombo woodland near water bodies, was present in large numbers at Hillwood Farm. Sequence similarity with P. porosissima (GenBank accession number KF0O27212) from Rwanda is 98%.
Ptychadena cf. taenioscelis Laurent, 1954
Stripe-legged Grass Frog
Material. HILLWOOD FARM: BMNH 2018.5784—-86 (Fig. 6N). Comments: Adult specimens found in pond. These specimens have been identified using the key in Poynton and Broadley (1985a). The closest match on GenBank (95%) is P. taenioscelis from the Republic of the
Amphib. Reptile Conserv.
Congo (GenBank accession number KY080397). Perret (1979) assigned records of taenioscelis from west and central Africa to pumilio Boulenger. There seems to be some confusion in the literature regarding the taxonomy of these species, and a review of the group is needed.
Ptychadena upembae (Schmidt and Inger, 1959) Upemba Ridged Frog
Material. HILLWOOD FARM: BMNH 2018.5750— 52 (last number is a juvenile); NKWAJI: BMNH 2018.574446 (juveniles), BMNH 2018.5747-49 (last number is a juvenile); SIMA NGWEZI NP: BMNH 2018.5762. Comments: Following the key in Poynton and Broadley (1985a), this series should be assigned to Ptychadena upembae. The BLAST search shows that the closest match (96%) is Ptychadena aff. porosissima (GenBank accession number DQ525940) from Tanzania, but it is important to note that there is no sequence of P. upembae available for comparison.
Pyxicephalidae
Amietia chapini (Noble, 1924)
Chapin's River Frog
Material. HILLWOOD FARM: BMNH 2018.5660-61, BMNH 2018.5662—63, BMNH 2018.5664 (Fig. 60), BMNH 2018.5665. Comments: Specimens found near streams in miombo woodland. The result of the BLAST search shows that A. chapini is the closest match to the specimens collected at Hillwood Farm (sequence similarity with A. chapini of 96-98%). All specimens have long legs (tibiofibula ~0.6 of snout-vent length), as noted by Noble (1924). I note that the specimens described as A. chapini by Channing et al. (2016) differ from the specimens listed above in coloration — the latter being darker. If these specimens are confirmed to be A. chapini, this will be the first record of this species for Zambia, but their presence in Zambia is not surprising considering the proximity (ca. 380 km) between the currently known populations from southern DRC (Kundelungu National Park) and Hillwood Farm.
Pyxicephalus cf. adspersus Tschudi, 1838
Giant Bullfrog
Material. SIOMA NGWEZI NP: BMNH 2018.5787- 91 (Fig. 6P). Comments: All individuals collected are juveniles. Hence, identification is tentative and based on the geographic range of the species. The BLAST shows 95% sequence similarity to Pyxicephalus cf. adspersus (DQ347304) and P. edulis (DQ022366).
Tomopterna marmorata (Peters, 1854)
Marbled Sand Frog
Material. LIVINGSTONE: BMNH — 2018.5792. Comments: Juvenile found in a small pond at night. Skin of dorsum and venter is smooth; venter is pale
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Table 4. Uncorrected pairwise distances (p-distances) for the 16S rRNA gene for Zomopterna spp. Distances between conspecific
populations are inside boxes.
1 Tomopterna sp. "Shankara" AY255095 7 Tomopterna sp. BMNH 2018.5793 Tomopterna sp. BMNH 2018.5794 Tomopterna sp. BMNH 2018.5795 Tomopterna sp. BMNH 2018.5796
Tomopterna delalandii DQ283403 0.06 0.06
2
3
4
5
6 Tomopterna sp. BMNH 2018.5797 7
8 Tomopterna damarensis KX869909 0.07 0.06 9
Tomopterna cryptotis JX564898 0.06 0.06 10 Tomopterna marmorata BMNH 2018.5792 0.08 0.08 11. Yomopterna marmorata AY 255084 0.08 0.08
with a few dark markings under the throat; tympanum indistinguishable; undivided sub-articular tubercle on first finger. The closest match on GenBank (100%) is Tomopterna marmorata (AY 255084) from Zambia.
Zomopterna sp.
Material. SIOMA NGWEZI NP: BMNH 2018.5793— 97 (Fig. 6Q). Comments: All individuals are juveniles and were found at night close to a light-trap set to collect beetles. The main morphological characters of these specimens are: dorsal and ventral skin smooth; immaculate venter; presence of a ridge below tympanum; tympanum indistinguishable; and undivided sub-articular tubercle on first finger. The closest match on GenBank (99%) is Tomopterna sp. “Shankara” (AY255095), an undescribed species from Namibia (Dawood et al. 2002). Table 4 shows the p-distances among the closest matches from GenBank.
Rhacophoridae
Chiromantis xerampelina Peters, 1854
African Grey Treefrog
Material. MAYUKUYUKU: BMNH 2018.5798 (Fig. 6R). Comments: This is a widespread species found in miombo woodland. One adult individual found at night on a tree near the campsite.
Reptilia
Order Squamata
Agamidae
Agama armata Peters, 1855
Northern Ground Agama
Material. NGONYE FALLS CAMP: BMNH 2018.2751
(Fig. 7A). Comments: One juvenile found basking on a log by the Zambezi River. Sequence similarity is
Amphib. Reptile Conserv.
0.06 0.06 0.06 0.06
0.07 0.07 0.07 0.07 0.06 0.05
4 5 6 7 8 i) 10 11
0.05 0.05 0.05 0.05 -
0.02 -
0.06 0.06 0.06 0.06 0.03 0.02 - 0.07 0.07 0.07 0.07 0.06 0.05 0.06 -
00 [aon] -
98% with A. armata ZFMK 84990 (GenBank accession number GU128447).
Chamaeleonidae
Chamaeleo dilepis Leach, 1819
Flap-necked Chameleon
Material. CHAVUMA FR: BMNH 2018.2755 (Fig. 7B), BMNH 2018.2756. Comments: Specimens were found in miombo woodland on shrubs above | m. There are currently seven subspecies in this group and based on their distributions, these specimens represent C. dilepis quilensis (Uetz et al. 2018). Sequence similarity is 99% with Chamaeleo dilepis from Matema, Tanzania (GenBank accession number AY927272).
Gekkonidae
Hemidactylus mabouia (Moreau de Jonnes, 1818) Common Tropical House Gecko
Material. I[TEZHI-TEZHI: BMNH 2018.2742, BMNH 2018.2740; NANZILA PLAINS: BMNH 2018.2741. Comments: Common species found in a variety of habitats, including heavily degraded ones, though not found in forests. The closest matches from GenBank are Hemidactylus mercatorius (AY 863034) and H. mabouia (AY 863038), both showing 94% sequence similarity.
Lygodactylus chobiensis Fitzsimons, 1932
Chobe Dwarf Gecko
Material. ITEZHI-TEZHI: BMNH 2018.2743 (female; Fig. 7C), BMNH 2018.2744 (male) Comments: Specimens found on tree-trunks in miombo. Identification follows the key provided by Broadley (1971). The rostral is excluded from the nostril and the male has dark forward-directed chevron marks on the throat. The female is yellow and white underneath (Fig. 7D). The closest match from GenBank (95% sequence similarity)
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Fig. 7. Reptiles of western Zambia. (A) Agama armata (BMNH 2018.2751), (B) Chamaeleo dilepis (BMNH 2018.2755), (C}H(D) Lygodactylus chobiensis (BMNH 2018.2743), (E) Pachydactylus punctatus, (F) Ichnotropis capensis (BMNH 2018.2750), (G) Meroles squamulosus (BMNH 2018.2753), (H) Trachylepis cf. albopunctata (BMNH 2018.2765), (I) Trachylepis varia (BMNH 2018.2769), (J) Zyphlacontias rohani (BMNH 2018.2761), (IK) Crotaphopeltis hotamboeia (BMNH 2018.2776), (L) Philothamnus hoplogaster (BMNH 2018.2775), (M) Rhamnophis aethiopissa ituriensis (BMNH 2018.2772), (N) Thelotornis kirtlandii (BMNH 2018.2760), (O) Atractaspis congica (BMNH 2018.2274), (P)-(R) Kinixys spekii.
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is Lygodactylus chobiensis from Zimbabwe (GenBank accession number GU593456).
Lygodactylus angolensis Bocage, 1896
Angola Dwarf Gecko
Material. NKWAJI: BMNH 2018.2767 (male), BMNH 2018.2766 (female). Comments: Specimens were identified following the key provided by Broadley (1971). In both specimens, the mental has a pair of lateral clefts, resulting from fusion with a large postmental. The male has nine preanal pores, which distinguish it from L. bradfieldi. The closest match from GenBank (90% sequence similarity) is Lygodactylus sp. from East Africa (GenBank accession numbers GU593448-50).
Pachydactylus punctatus Peters, 1854
Speckled Thick-toed Gecko
Material. I[TEZHI-TEZHI: BMNH 2018.2757, BMNH 2018.2758 (Fig. 7E), BMNH 2018.2759. Comments: Specimens found at night in dry leaf litter in miombo woodland. All specimens have the dorsum covered with sub-uniform granules. The closest matches from GenBank (93% sequence similarity) are Pachydactylus punctatus (AF449120) and P. scherzi (AY123379). As the latter is only known from Namibia (Bauer and Branch 1995), I assign these specimens to P. punctatus.
Gerrhosauridae
Gerrhosaurus bulsi Laurent, 1954
Laurent’s Plated Lizard
Material. HILLWOOD FARM: BMNH 2018.2754. Comments: One juvenile collected by the farm scouts in dry miombo. The BLAST search shows the closest match (98%) as Gerrhosaurus bulsi from Angola (KF717381). Broadley (1971, 1991) referred the population of Gerrhosaurus from Ikelenge (north-western Zambia) to multilineatus but this was later contested by Haagner et al. (2000). Bates et al. (2013) consider G. bulsi a valid species and discuss the taxonomic problems regarding G. multilineatus.
Lacertidae
Ichnotropis capensis (Smith, 1838)
Cape Rough-scaled Lizard
Material. CHAVUMA FR: BMNH = 2018.2746; LUKWAKWA: BMNH 2018.2749, BMNH 2018.2747; NANZILA PLAINS: BMNH_ 2018.2750, BMNH 2018.2748 (Fig. 7F); SIOMA NGWEZI NP: BMNH 2018.2745. Comments: All specimens are juveniles and were found in miombo woodland. Sequence similarity is 99% with [. capensis from Katima Mulilo, Namibia (GenBank accession number JX962898).
Meroles squamulosus (Peters, 1854) Savanna Lizard
Amphib. Reptile Conserv.
Material. NANZILA PLAINS: BMNH _ 2018.2753 (Fig. 7G); SIOMA NGWEZI NP: BMNH 2018.2752. Comments: Two adult males found in miombo. Sequence similarity 1s 96% with Meroles (Ichnotropis) squamulosus from Laela, Tanzania (GenBank accession number JX962897).
Scincidae
Panaspis cf. wahlbergi (Smith, 1849)
Snake-eyed Skink
Material. CHAVUMA FR: BMNH 2018.2738, BMNH 2018.2739. Comments: Specimens found during the day in leaf litter. The closest match from GenBank (98%) is Panaspis sp. (KU236726), from Katanga, DRC. Medina et al. (2016) provide a molecular phylogeny of this genus, which suggests that there is cryptic diversity within P. wahlbergi.
Trachylepis cf. albopunctata (Bocage, 1867)
Angolan Variable Skink
Material. ITEZHI-TEZHI: BMNH — 2018.2762; MAYUKUYUKU: BMNH_ 2018.2765 (Fig. 7H), BMNH_ 2018.2763. Comments: Specimens found during the day. The BLAST search shows 99-100% sequence similarity with sequences from 7! varia clade B (accession numbers MG605651—59), which was recently assigned to Trachylepis cf. albopunctata by Marques et al. (2018).
Trachylepis damarana (Peters, 1870)
Damara Skink
Material. SIOMA NGWEZI NP (HQ): BMNH 2018.2764. Comments: Morphologically similar to 7. varia group. However, both its distribution and sequence similarity (99%) match T. damarana (see Weinell and Bauer 2018).
Trachylepis wahlbergii (Peters, 1869)
Wahlberg’s Striped Skink
Material. ITEZHI-TEZHI: BMNH 2018.2769 (Fig. 71); LUKWAKWA: BMNH = 2018.2768, BMNH 2018.2770; NK WAJI: BMNH 2018.2771. Comments: Specimens found during the day on rocks (Itezhi-Tezhi), dambo (Lukwakwa), and inside a tree trunk at Nkwaji. According to Broadley (2000) these specimens fall in the distribution range of 7. wahlbergii. The most similar sequence from GenBank is 7’ wahlbergii from Zambia (99%, accession number DQ234810).
Typhlacontias rohani Angel, 1923
Rohan's Blind Dart Skink
Material. SIOMA NGWEZI NP (HQ): BMNH 2018.2761 (Fig. 7J). Comments: One specimen was found buried in sand and collected by Errol Pietersen. The closest match on GenBank (90%) is 7. punctatissimus (DQ316889). There is no sequence of 7. rohani available
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Fig. 8. Drawings of Rhamnophis aethiopissa ituriensis (BMNH 2018.2772). Head and anterior of body in dorsal, left lateral, and ventral views. Scale bar 10 mm. Drawings by Ed Wade.
for comparison. Colubridae
Crotaphopeltis hotamboeia (Laurenti, 1768) White-lipped Herald Snake
Material. ITEZHI-TEZHI: BMNH 2018.2776 (Fig. 7K); HILLWOOD FARM: BMNH 2018.2773; SIOMA NGWEZI NP: BMNH 2018.2777. Comments: All individuals collected are juveniles and were found at night in miombo woodland near rocks. The most similar sequence on GenBank is C. hotamboeia from Malawi (99%, accession number AY611816).
Philothamnus hoplogaster (Gunther, 1863)
Green Water Snake
Material. NANZILA PLAINS: BMNH 2018.2775 (Fig. 7L). Comments: Specimen found while eating tadpoles and juveniles of Kassina sp. in a temporary pond during the day. This species is similar to other Philothamnus but usually smaller (Marais 2004). The closest match on GenBank (99%) is P. hoplogaster from Mozambique (accession number FJ913484).
Rhamnophis aethiopissa ituriensis (Schmidt 1923) Large-eyed Green Tree Snake
Material. HILLWOOD FARM: BMNH 2018.2772 (Fig. 7M; Fig. 8). Comments: Specimen found in leaf litter of riverine forest during the day. Broadley (1991) provided the first record of this species for Zambia. Based on distribution, this form represents the subspecies R. a. ituriensis from Niapu in the DRC (see Eimermacher 2012).
Amphib. Reptile Conserv.
Thelotornis kirtlandii (Hallowell, 1844)
Forest Vine Snake
Material. HILLWOOD FARM: BMNH 2018.2760 (Fig. 7N). Comments: One juvenile was found at night resting on green vegetation in riverine forest (mushito). Species was identified using the key in Broadley (2001) and the following characters were observed: top of the head, including temporal region, is uniform green; rostral and nasals are strongly recurved onto top of snout; supralabials are white with small green spots.
Lamprophiidae
Atractaspis congica Peters, 1877
Congo Stiletto Snake
Material. HILLWOOD FARM: BMNH 2018.2274 (Fig. 70). Comments: One relatively large specimen found in moist leaf litter inside a patch of mushito at night. Broadley and Blaylock (2013): 232 expanded the description of A. bibronii to accommodate the condition of 19 midbody scales of A. congica (19-21 scales at midbody). This specimen exhibits erratic counts of 19+17, the latter count predominating after the 84" ventral.
Order Testudines Testudinidae
Kinixys spekii Gray, 1863
Speke’s Hinge-back Tortoise
Comments: One specimen (Figs. 5P—R) was found in mushito at Nkwaji. The carapace was hinged between the 7" and 8" marginals. The specimen was photographed and released.
Discussion
This is a non-comprehensive list of the herpetofauna of western Zambia. The survey was conducted shortly after the general breeding season for amphibians and reptiles in this part of the world, and consequently most specimens collected are juveniles. For the same reason, most species were not active, which made the search for them more challenging. However, some species of amphibians were active during the survey. Phrynobatrachus were heard calling during the day and at night, and at least ten males of Sclerophrys lemairii were calling during the day in Lukwakwa (see Bittencourt-Silva 2014). The presence of juveniles of /chnotropis capensis and adults of Meroles squamulosus in sympatry is explained by their staggered life cycles (see Broadley 1967, 1979).
DNA barcoding is an important tool for identifying candidate species. However, there are a number of caveats. For instance, for amphibians, Vences et al. (2005a) propose a tentative 16S rRNA threshold at 5% for interspecific sequence divergence but also
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highlight the broad overlap of intra- and interspecific divergence values (see Vences et al. 2005b, p. 1,865) that complicates the establishment of threshold values. Using DNA barcoding alone can potentially lead to simplistic diagnoses of putative species. Another issue with DNA barcoding relates to the taxonomic accuracy of public DNA databases (e.g., GenBank, BOLD, EMBL). Misidentified sequences are not uncommon (Bridge et al. 2003; Vilgalys 2003), which reinforces the importance of vouchering all sequences deposited. Table 2 shows that the intra-specific p-distances within H. marmoratus are considerably large in some cases. This could be partly due to geographic distances, given that the specimens are from Ghana, Guinea-Bissau, Kenya, and Tanzania. A taxonomic review of this group is clearly necessary. Nonetheless, genetic data may be crucial in cases where species are genetically different but morphologically largely conserved. An example is the mongrel frogs from Mozambique and Malawi, which have ca. 5% interspecific divergences but are in general phenotypically indistinguishable (Conradie et al. 2018).
Some of the taxa reported here could not be assigned to currently recognized species based on DNA barcoding and/or external morphology. For instance, based on p-distances, some of the Phrynobatrachus specimens represent putative new species (Table 3). Zimkus and Schick (2010) suggest that there are at least two species currently identified as Phrynobatrachus natalensis in East Africa, and another two clades are reported from western and southern Africa (Zimkus et al. 2010). The phylogeny presented here indicates that there may be more species of this group in western Zambia (see Fig. 5). Species identified here as P. cf. parvulus may be a new species. These results corroborate the conclusions of Zimkus and Schick (2010) and Zimkus et al. (2010) that a taxonomic review of the genus Phrynobatrachus is needed. Similarly, the Zomopterna population found in Sioma Ngwezi NP could represent an undescribed species (Table 4) previously reported from Namibia (see Dawood et al. 2002). These taxa deserve further investigations by specialists. It is often the case that original species descriptions lack diagnostic details, including illustration of characters, and/or type material may be lost or in poor condition, all of which can contribute to inconclusive or even incorrect species identification. Additional datasets (e.g., bioacoustics, ecology) often provide important information and can solve some of these taxonomic conundrums.
The genus Ptychadena currently comprises 56 species, some (possibly many) representing species complexes (e.g., Zimkus et al. 2016). Nineteen species have been reported in Zambia (see genus account in Frost 2018), eleven of which were recorded during this survey. The lack of an updated key to the Ptychadena of Zambia makes the species identification process challenging. Similarly, barcoding is not helpful when there are no reference sequences available. A search of
Amphib. Reptile Conserv.
GenBank for 16S sequences of Ptychadena shows that 41% of currently recognized species are not represented, and 22% of the sequences available are either pending confirmation or identified only to genus. The taxonomy of this group is clearly in need of attention.
Except for a few areas—on the extreme north-west of the country and along the Zambezi river—most of Zambia remains poorly studied. Recently, Channing and Willems (2018) described a new species of Ptychadena from the northern part of the country, and a new cryptic species of Polemon (Squamata: Lamprophiidae) described from the DRC and Uganda is likely to occur in Zambia (Portillo et al. 2019). The list of species provided here adds new points to the map of the Zambian herpetofauna.
The herpetofauna of Zambia is mostly contained in the Zambezian biogeographical core, with only the south-western region forming part of the South African core (sensu Linder et al. 2012). Not surprisingly, many species found during this survey also occur in the DRC (e.g., Channing et al. 2016), Angola (Conradie et al. 2016) and Namibia (Dawood et al. 2002), including Amietia chapini, recorded here for the first time from Zambia. Four species of amphibians that Pietersen et al. (2017) expected to occur near Ngonye Falls are now confirmed to occur at Sioma Ngwezi NP (Kassina senegalensis, Phrynobatrachus mababiensis, Ptychadena porosissima, and Pyxicephalus adspersus). The still very incomplete knowledge of the Zambian herpetofauna remains the main obstacle to our understanding of its biogeography and the conservation statuses of its constituent species.
Acknowledgements.—| thank the entomologists Hitoshi Takano, Lucia Chmurova, and Lydia Smith for support in the field. I am grateful to Errol Pietersen for assisting with logistics and collection of specimens at Ngonye Falls, and to Darren Pietersen for helping with identification of some reptiles and providing valuable comments on the manuscript. The expedition to Zambia would not have been possible without the support of Richard Smith, to whom I am extremely grateful. I thank Werner Conradie and Mark Wilkinson for their valuable comments on the manuscript. Ed Wade kindly provided the scale counting of Atractaspis congica and the drawings of Rhamnophis aethiopissa ituriensis. | thank Simon P. Loader for his invaluable support. I appreciate the help of Dan Portik and Breda Zimkus with identification of Hyperolius and Phrynobatrachus, respectively. Permits to collect and export specimens were issued by the Department of Veterinary Services, Ministry of Livestock and Fisheries Development (ICS#08417).
Literature Cited Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ. 1990. Basic local alignment search tool. Journal of
Molecular Biology 215: 403-410. AmphibiaWeb. 2018. Available: https://amphibiaweb.
August 2019 | Volume 13 | Number 2 | e181
Herpetological survey of western Zambia
org. [Accessed: 17 July 2018].
Bates MF, Tolley KA, Edwards S, Davids Z, Da Silva JM, Branch WR. 2013. A molecular phylogeny of the African plated lizards, genus Gerrhosaurus Wiegmann, 1828 (Squamata: Gerrhosauridae), with the description of two new genera. Zootaxa 3750: 465-493.
Bauer AM, Branch WR. 1995. Geographic variation in western populations of the Pachydactylus punctatus complex (Reptilia: Gekkonidae). Tropical Zoology 8: 69-84.
Bittencourt-Silva GB. 2014. Notes on the reproductive behaviour of Amietophrynus lemairii (Boulenger, 1901) (Anura: Bufonidae). Herpetology Notes 7: 611-614.
Branch WR. 1998. Field Guide to the Snakes and other Reptiles of Southern Africa. Struik, Cape Town, South Africa. 399 p.
Bridge PD, Roberts PJ, Spooner BM, Panchal G. 2003. On the unreliability of published DNA sequences. New Phytologist 160: 43-48.
Broadley DG. 1967. The life cycles of two sympatric species of /chnotropis (Sauria: Lacertidae). African Zoology 3: \-2.
Broadley DG. 1971. The Reptiles and Amphibians of Zambia. The Puku, Number 6. Kenya Litho, Nairobi, Kenya. 143 p.
Broadley DG. 1979. A field study of two sympatric annual lizards (genus Ichnotropis) in Rhodesia. African Zoology 14: 133-138.
Broadley DG. 1991. The herpetofauna of northern Mwinilunga District, northwestern Zambia. Arnoldia Zimbabwe 9: 519-538.
Broadley DG. 2000. A review of the genus Mabuya in southeastern Africa (Sauria: Scincidae). African Journal of Herpetology 49: 87-110.
Broadley DG. 2001. A review of the genus 7helotornis A. Smith in eastern Africa, with the description of a new species from the Usambara Mountains (Serpentes: Colubridae: Dispholidini). African Journal of Herpetology 50(2): 53-70.
Broadley DG, Blaylock R. 2013. Snakes of Zimbabwe and Botswana. Edition Chimaira, Frankfurt am Main, Germany. 387 p.
Broadley DG, Doria C, Wigge J. 2003. Snakes of Zambia: An Atlas and Field Guide. Edition Chimaira, Frankfurt am Main, Germany. 280 p.
Castresana J. 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17. 540— ao2.
Channing A. 2001. Amphibians of Central and Southern Africa. Comstock Publishing Associates, Ithaca, New York, USA. 470 p.
Channing A, Broadley DG. 2002. A new snout-burrower from the Barotse floodplain (Anura: Hemisotidae: Hemisus). Journal of Herpetology 36: 367-372.
Amphib. Reptile Conserv.
Channing A, Willems F. 2018. A new grass frog with rupicolous tadpoles from northern Zambia (Anura: Ptychadenidae). Zootaxa 4462: 349-366.
Channing A, Hillers A, Lotters S, Rodel M-O, Schick S, Conradie W, Roédder D, Mercurio V, Wagner P, Dehling JM. 2013. Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species. Zootaxa 3620: 301-350.
Channing A, Dehling JM, Lotters S, Ernst R. 2016. Species boundaries and taxonomy of the African river frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa 4155: 1-76.
Conradie W, Bills R. 2017. Wannabe Ranid: Notes on the morphology and natural history of the Lemaire’s Toad (Bufonidae: Sclerophrys lemairii). Salamandra 53: 439-444.
Conradie W, Branch WR, Measey GJ, Tolley KA. 2012. A new species of Hyperolius Rapp, 1842 (Anura: Hyperoliidae) from the Serra da Chela mountains, south-western Angola. Zootaxa 3269: 1-17.
Conradie W, Bills R, Branch WR. 2016. The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola. Amphibian & Reptile Conservation 10(2) [Special Section]: 6-36 (e126).
Conradie W, Bittencourt-Silva GB, Farooq HM, Loader SP, Menegon M, Tolley KA. 2018. New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozambique inselbergs. African Journal of Herpetology 67: 61-85.
Dawood A, Channing A, Bogart JP. 2002. A molecular phylogeny of the frog genus Tomopterna in southern Africa: examining species boundaries with mitochondrial 12S rRNA sequence data. Molecular Phylogenetics and Evolution 22: 407-413.
Dehling JM, Sinsch U. 2013. Diversity of Ptychadena in Rwanda and taxonomic status of P. chrysogaster Laurent, 1954 (Amphibia, Anura, Ptychadenidae). ZooKeys 356: 69-102.
Deichmann JL, Mulcahy DG, Vanthomme H, Tobi E, Wynn AH, Zimkus BM, McDiarmid RW. 2017. How many species and under what names? Using DNA barcoding and GenBank data for west Central African amphibian conservation. PLoS ONE 12: 0187283.
Eimermacher TG. 2012. Phylogenetic systematics of Dispholidine colubrids (Serpentes: Colubridae). Ph.D. Dissertation, The University of Texas at Arlington, Arlington, Texas, USA. 109 p.
Frost DR. 2018. Amphibian sSpecies of the World: an online reference. Version 6.0. American Museum of Natural History, New York, New York, USA. Available: —http://research.amnh.org/vz/herpetology/ amphibia/index.php [Accessed: 10 November 2018].
Furman BLS, Bewick AJ, Harrison TL, Greenbaum E, Gvozdik V, Kusamba C, Evans BJ. 2015. Pan-African phylogeography of a model organism, the African
August 2019 | Volume 13 | Number 2 | e181
Bittencourt-Silva
clawed frog Xenopus laevis. Molecular Ecology 24: 909-925.
Haagner GV, Branch WR, Haagner AJF. 2000. Notes ona collection of reptiles from Zambia and adjacent areas of the Democratic Republic of the Congo. Annals of the Eastern Cape Museum |: 1-25.
Hebert PDN, Gregory TR. 2005. The promise of DNA barcoding for taxonomy. Systematic Biology 54: 852— 859.
Katoh K, Misawa K, Kuma K, Miyata T. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30: 3,059-3,066.
Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, et al. 2012. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1,647—1,649.
Kohler J, Scheelke K, Schick S, Veith M, Lotters S. 2005. Contribution to the taxonomy of hyperoliid frogs (Amphibia: Anura: Hyperoltidae): advertisement calls of twelve species from East and Central Africa. African Zoology 40: 127-142.
Linder HP, de Klerk HM, Born J, Burgess ND, Fyeldsa J, Rahbek C. 2012. The partitioning of Africa: statistically defined biogeographical regions in sub- Saharan Africa. Journal of Biogeography 39: 1,189- 1,205.
Marais J. 2004. A Complete Guide to the Snakes of Southern Africa. Struik Nature, Cape Town, South Africa. 740 p.
Marques MP, Ceriaco LMP, Blackburn DC, Bauer AM. 2018. Diversity and distribution of the amphibians and terrestrial reptiles of Angola. Atlas of historical and bibliographic records (1840-2017). Proceedings of the California Academy of Sciences 65(Supplement 1): 1-501.
Medina MF, Bauer AM, Branch WR, Schmitz A, Conradie W, Nagy ZT, Hibbitts TJ, Ernst R, Portik DM, Nielsen SV. 2016. Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution 100: 409- 423.
Meier R, Shiyang K, Vaidya G, Ng PKL. 2006. DNA barcoding and taxonomy in Diptera: a tale of high intraspecific variability and low identification success. Systematic Biology 55: 715-728.
Noble GK. 1924. Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909-1915. Part 3, Amphibia. Bulletin of the American Museum of Natural History 49(2): 147-347.
Palumbi SR, Martin A, Romano S, McMillan WO, Stice L, Grabowski G. 1991. The Simple Fool’s Guide to PCR. Version 2.0. Privately published document
Amphib. Reptile Conserv.
compiled by S. Palumbi. Department of Zoology, University of Hawaii, Honolulu, Hawaii, USA.
Perret JL. 1979. Remarques et mise au point sur quelques especes de Ptychadena (Amphibia, Ranidae). Bulletin de la Société Neuchdteloise des Sciences Naturelles 102: 5-21.
Phiri P. 2005. A Checklist of Zambian Vascular Plants. SABONET, Pretoria, South Africa. 169 p.
Pickersgill M. 2007. Frog Search, Results of Expeditions to Southern and Eastern Africa. Frankfurt Contributions to Natural History. Editions Chimaira, Frankfurt am Main, Germany. 574 p.
Pietersen DW, Pietersen EW, Conradie W. 2017. Preliminary herpetological survey of Ngonye Falls and surrounding regions in south-western Zambia. Amphibian & Reptile Conservation 11(1) [Special Section]: 24-43 (e148).
Portillo F, Branch WR, Tilbury CR, Nagy ZT, Hughes DF, Kusamba C, Muninga WM, Behangana M, Greenbaum E. 2019. A cryptic new species of Polemon (Squamata: Lamprophiidae, Aparallactinae) from the Miombo woodlands of Central and East Africa. Copeia 107: 22-36.
Poynton JC, Broadley DG. 1985a. Amphibia Zambesiaca 2. Ranidae. Annals of the Natal Museum 27. 115-181.
Poynton JC, Broadley DG. 1985b. Amphibia Zambesiaca 1. Scolecomorphidae, Pipidae, Microhylidae, Hemisidae, Arthroleptidae. Annals of the Natal Museum 26: 503-553.
Poynton JC, Broadley DG. 1987. Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae. Annals of the Natal Museum 28: 161-229.
Poynton JC, Broadley DG. 1988. Amphibia Zambesiaca 4. Bufonidae. Annals of the Natal Museum 29: 447— 490.
Poynton JC, Broadley DG. 1991. Amphibia Zambesiaca 5. Zoogeography. Annals of the Natal Museum 39: 221-277.
Poynton JC, Loader SP, Conradie W, Roedel M-O, Liedtke HC. 2016. Designation and description of a neotype of Sclerophrys maculata (Hallowell, 1854), and reinstatement of S. pusilla (Mertens, 1937) (Amphibia: Anura: Bufonidae). Zootaxa 4098: 73-94.
Du Preez L, Carruthers V. 2017. Frogs of Southern Africa: A Complete Guide. Struik Nature, Cape Town, South Africa. 519 p.
SchiotzA. 1975. Treefrogs of Eastern Africa. Steenstrupia, Copenhagen, Denmark. 232 p.
Schiotz A. 1999. Treefrogs of Africa. Andreas S. Brahm, Frankfurt am Main, Germany. 350 p.
Stamatakis A. 2014. RAXxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1,312-1,313.
Swofford DL. 2001. PAUP*: Phylogenetic Analysis Using Parsimony (and other methods) 4.0.65. Sinauer Associates, Sunderland, Massachusetts, USA.
Uetz P, Freed P, HoSek J. 2018. The Reptile Database.
August 2019 | Volume 13 | Number 2 | e181
Herpetological survey of western Zambia
Available: http://www.reptile-database. org [Accessed: 15 February 2019].
Vences M, Thomas M, Bonett RM, Vieites DR. 2005a. Deciphering amphibian diversity through DNA barcoding: chances and challenges. Philosophical Transactions of the Royal Society B: Biological Sciences 360: 1,859-1,868.
Vences M, Thomas M, van der Meijden A, Chiari Y, Vieites DR. 2005b. Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology 2: \—12.
Vilgalys R. 2003. Taxonomic misidentification in public DNA databases. New Phytologist 160: 4-5.
Wagner P, Wilms TM, Rodder D, Schmitz A. 2013. A great leap—the first record of Xenopus pygmaeus (Anura: Pipidae) from south of the Congo Basin. Salamandra 49: 206-210.
Weinell JL, Bauer AM. 2018. Systematics and phylogeography of the widely distributed African
Amphib. Reptile Conserv.
20
skink Trachylepis varia species complex. Molecular Phylogenetics and Evolution 120: 103-117.
Zimkus BM, Schick S. 2010. Light at the end of the tunnel: insights into the molecular systematics of East African puddle frogs (Anura: Phrynobatrachidae). Systematics and Biodiversity 8: 39-47.
Zimkus BM, Rodel MO, Hillers A. 2010. Complex patterns of continental speciation: Molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus). Molecular Phylogenetics and Evolution 55: 883-900.
Zimkus BM, Lawson LP, Barej M, Barratt CD, Channing A, Dash KM, Dehling JM, Du Preez L, Gehring P-S, Greenbaum E, et al. 2016. Leapfrogging into new territory: How Mascarene ridged frogs diversified across Africa and Madagascar to maintain their ecological niche. Molecular Phylogenetics and Evolution 106: 254—269.
Gabriela Bittencourt is a Brazilian evolutionary biologist with research experience in natural history, evolution, ecology, and biogeography, and a particular focus on amphibians. Gabriela has more than 15 years of herpetological laboratory and fieldwork experience in the Neo- tropics, Africa, and Asia. Her research has focused on understanding phylogenetic relation- ships and biotic distribution patterns of amphibians. Gabriela is currently a Research Assistant in the Herpetology Group at the Natural History Museum, London, United Kingdom.
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Appendix 1. List of amphibians and reptiles found in western Zambia, including species vouchers, GenBank accession numbers, and locality information. Museum acronym: BMNH — Natural History Museum, London, United Kingdom. GPS datum WGS-84.
Species
AMPHIBIA: ANURA ARTHROLEPTIDAE Arthroleptis stenodactylus Arthroleptis stenodactylus Arthroleptis stenodactylus Arthroleptis stenodactylus
Arthroleptis stenodactylus
Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus
Arthroleptis xenochirus
Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus Arthroleptis xenochirus BUFONIDAE
Sclerophrys gutturalis Sclerophrys gutturalis Sclerophrys gutturalis Sclerophrys gutturalis Sclerophrys gutturalis Sclerophrys gutturalis Sclerophrys gutturalis Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys lemairii Sclerophrys pusilla Sclerophrys pusilla Sclerophrys pusilla Sclerophrys pusilla
Voucher ID
BMNH 2018.5826 BMNH 2018.5827 BMNH 2018.5828 BMNH 2018.5829 BMNH 2018.5830 BMNH 2018.5814 BMNH 2018.5815 BMNH 2018.5816 BMNH 2018.5817 BMNH 2018.5818 BMNH 2018.5819 BMNH 2018.5820 BMNH 2018.5811 BMNH 2018.5812 BMNH 2018.5813 BMNH 2018.5821 BMNH 2018.5822 BMNH 2018.5823 BMNH 2018.5824
BMNH 2018.5825
BMNH 2018.5703 BMNH 2018.5705 BMNH 2018.5706 BMNH 2018.5707 BMNH 2018.5702 BMNH 2018.5701 BMNH 2018.5704 BMNH 2018.5723 BMNH 2018.5715 BMNH 2018.5716 BMNH 2018.5717 BMNH 2018.5718 BMNH 2018.5719 BMNH 2018.5720 BMNH 2018.5721 BMNH 2018.5722 BMNH 2018.5709 BMNH 2018.5710 BMNH 2018.5708
BMNH 2018.5711
Amphib. Reptile Conserv.
Field ID GenBank Locality Latitude SL2109 MK464479 ~~ Lukwakwa -12.66084 SL2121 | MK464478 ~~ Lukwakwa -12.66084 SL2123, MK464477_ ~— Lukwakwa -12.66084 SL2128 = MK464476 ~~ Lukwakwa -12.66084 SL2221 MK464475 — Nkwaji -11.57728 SL2145 =MK464471 — Hillwood Farm -11.26316 SL2146 =MK464470 — Hillwood Farm -11.26316 SL2147, MK464469 — Hillwood Farm -11.26316 SL2148 = MK464468 — Hillwood Farm -11.26316 SL2152. MK464467 — Hillwood Farm -11.26690 SL2153. MK464466 — Hillwood Farm -11.26690 SL 2246 MK464465 — Hillwood Farm -11.26690 SL2122 MK464474 ~~ Lukwakwa -12.66084 SL2124 MK464473 ~~ Lukwakwa -12.66084 SL2125 MK464472 ~~ Lukwakwa -12.66084 SL 2200 MK464464 Nkwaji -11.60592 SL 2201 MK464463 = Nkwaji -11.60592 SL 2202 MK464462 = Nkwaji -11.60592 SL 2203. MK464461 Nkwaji -11.60592 SL2204 MK464460 = Nkwaji -11.60592 SL2102 Chavuma FR -13.07006 SL2108 = MK464294 ~~ Lukwakwa -12.66084 SL2107 MK464293 ~Lukwakwa -12.74275 SL 2111 Lukwakwa -12.66084 SL 2069 MK464296 Maramba Lodge, Livingstone -17.89120 SL2025 MK464297 = Mayukuyuku, Kafue NP -14.91533 SL2190 MK464295 Nkwaji -11.60592 SL 2245 Hillwood Farm -11.26690 SL 2112 Lukwakwa -12.66084 SL2113 Lukwakwa -12.66084 SL 2114 Lukwakwa -12.66084 SL 2115 Lukwakwa -12.66084 SL 2116 Lukwakwa -12.66084 SL 2117 Lukwakwa -12.66084 SL 2118 Lukwakwa -12.66084 SL 2119 Lukwakwa -12.66084 SL 2047. MK464291 _ Itezhi-Tezhi, Kafue NP -15.77340 SL 2053. MK464290 _ Itezhi-Tezhi, Kafue NP -15.77340 SL2012 MK464292. Mayukuyuku, Kafue NP -14.91533 SL2189 MK464289 = Nkwaji -11.60592 21
Longitude
24.43697 24.43697 24.43697 24.43697 24.53960 24.32782 24.32782 24.32782 24.32782 24.31666 24.31666 24.31666 24.43697 24.43697 24.43697 24.55448 24.55448 24.55448 24.55448 24.55448
22.92880 24.43697 24.28436 24.43697 25.85821 26.06311 24.55448 24.31666 24.43697 24.43697 24.43697 24.43697 24.43697 24.43697 24.43697 24.43697 26.01151 26.01151 26.06311 24.55448
Altitude
1063 1063 1063 1063 1291 1356 1356 1356 1356 1308 1308 1308 1063 1063 1063 1244 1244 1244