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# Mountain Reedbuck: A Fifty-Year Personal Retrospective

by

L. R. Irby

*******

Published by

L. R. Irby at Smashwords

Copyright 2019 L. R. Irby

Smashwords Edition, License Notes

Thank you for downloading this free e-book. You are welcome to share it with your friends. This book may be reproduced, copied and distributed for non-commercial purposes, provided the book remains in its complete original form. Thank you for your support.

Disclaimer: This is a personal discourse on the biology of mountain reedbuck. In it, I have included numerous references from a wide variety of sources. I have attempted to give credit for ideas developed by other authors, but the conclusions presented in this book are my own. I have not intentionally distorted the views of other individuals, but for any mistakes in facts or attribution, I apologize.

Dedication: To the field biologists who devote their careers to collecting basic data needed to understand and manage wild animals.

# Table of Contents

Chapter 1 – Introduction

Chapter 2 - Background

Chapter 3 – Information Available in 1969

Chapter 4 – Grand Design

Chapter 5 - Behavior Patterns

Chapter 6 - Social Organization

Chapter 7 – Carcass Collections

Chapter 8 - Captive Animals

Chapter 9 - Habitat Use

Chapter 10 - Census Attempts

Chapter 11 - Limiting Factors

Chapter 12 - New Information 1976-2019

Chapter 13 - Status of Study Areas 2019

Chapter 14 - Future Prospects

Chapter 15 - Calling It a Day

Literature Cited

# Chapter 1 \- Introduction

My first independent research experience began in 1969. I was a cocky graduate with a B.S. in Fish and Wildlife Management from Texas A&M University. I was also commissioned as a second lieutenant in the U.S. Army Field Artillery in May 1969. Because I was able to get a deferment from the army for graduate school, I was selected for an informal exchange program hatched out by Dr. Jim Teer, my undergraduate advisor, and Professor Waldo Meester, a faculty member at the University of Pretoria. I arrived in South Africa in July 1969 with two suitcases and more self-confidence than biological knowledge and was given basically free rein to describe the natural history of a small, inconspicuous antelope, the mountain reedbuck ( _Redunca fulvorufula_ ). Not much was known about the species in 1969. I spent nearly three years in the field collecting data and most of five more years in graduate school trying to make sense of the data I collected. Altogether, this species consumed nearly eight years of my life, and I'm not sure I had a good handle on the critter when I finally finished my Ph.D. I published 10 or 12 scientific papers on the beast (see Irby in Literature Cited section), but in retrospect, they were only okay to "fair-to-middling." None of them revolutionized biology. Now, after retiring from an academic career built largely on a search committee's opinion of my early work on reedbuck, I have the time to try to objectively examine my conclusions and see how this species fared in the past half century.

I will use my work as a jumping off point, but a large part of this book is dependent on information collected by other (and likely smarter) grad students and agency biologists in Africa. The world has definitely changed in the past fifty years. In 1969, manual searches of libraries at Texas A&M and the University of Pretoria yielded fewer than 100 articles relevant to mountain reedbuck. A half-second search on Google in 2019 yielded over 20,000 references on " _Redunca fulvorufula_." When I arrived in Pretoria, mountain reedbuck were regarded as a minor player in the world of African large mammals. They were small, had unimpressive horns, lived in small groups, and had a disjunct distribution associated with moderately specific habitats. No one considered them crucial to conservation in Africa, and no one was particularly worried about their survival. The situation has changed. In 2017, they were listed by the International Union for Conservation of Nature (IUCN) as endangered. The major threats were identified as habitat loss/degradation and hunting. Legal hunting is regulated, but illegal hunting has increased dramatically in the last 50 years. Mountain reedbuck are vulnerable to uncontrolled take because they are distributed unevenly in pockets of suitable habitat. Small localized populations of most species can be easily destroyed, and mountain reedbuck are slow to re-colonization lost ground. If the lost habitat is distant from extant populations, natural re-colonization may be virtually impossible in many areas given current human land uses.

Throughout this book, I have used my own photographs where possible, rather than the much better photos posted on the Internet. When I did use images from other sources, I retrieved them from public sites (primarily Wikimedia – http//:commons.wikimedia.org/wiki) and have credited the photographers/organizations that produced them. All of the tables and graphs in the book were taken from data in my dissertation (Irby 1976). Conclusions based on my data and on papers by other biologists are my own. I tried to report information from sources beyond my dissertation as accurately as possible. For any errors, I will apologize in advance.

# Chapter 2 - Background

Mountain reedbuck (Fig. 1) were first recognized as a distinct species in 1782 and were formally named " _Redunca fulvorufala_ , Allamand 1815" in the early nineteenth century (Sclater 1900). They were placed in the Kingdom Animalia, the class Mammalia, the order Artiodactyla, sub-order Ruminantia, infraorder Pecora, and family Bovidae This system, still in use today, dates back to the eighteenth century (Linnaeus 1735) and reflects the inherent tendency of humans to try explaining things by putting them in boxes. In general, species with a few important common characteristics would be grouped in higher order categories (i.e. Animalia includes species that share characteristics such as multi-cellular bodies, membrane-bounded organelles, and flexible cell walls. This category includes organisms as different as jellyfish and elephants). Lower-order categories group organisms that share more and more characteristics. This system worked reasonably well during the 18th and 19th century for vertebrates. As scientific knowledge accumulated and technology improved, scientists realized that most of the important characteristics used to define groupings in the Linnaean system reflected the influence of genes inherited from common ancestors. Classification of invertebrates. plants, and the phyla containing "simpler" single and multi-cellular organisms does not lend itself to a system based on physical characteristics. This system does not work well at all for lower levels of classification of self-replicating entities that fall in the "not-quite-life" category, such as viruses and prions.

Figure 1. Southern mountain reedbuck ram and ewe in the Orhigstad Dam Nature Reserve,

South Africa, 1970.

In the Linnaean classification system, the mountain reedbuck has been placed in the Order Artiodactyla because they have limbs modified to enhance running, teeth specialized for clipping and grinding plants, and a digestive tract modified for extracting nutrients from plant material. Using the widely (although not universally) accepted classification scheme provided by Nowak (1999), these modifications include reduced numbers of digits, usually two or four rather than the ancestral five digits associated with ancient mammals, elongation of lower leg bones, reductions in numbers of bones in feet/ankles, teeth with high grinding ridges, and digestive-tract modifications that allow them to better process a low-quality plant diet. The sub-order Ruminantia is characterized by a complex digestive system, including a three or four-chambered stomach, which allows animals in this sub-order to digest coarse plant material with the help of, to use obsolete semi-technical terms, bacterial and protistan symbionts. Species in Ruminantia typically tear off plant parts using lower incisors and a cartilaginous pad on the upper jaw. Molars and premolars are used to break plant material into fragments which are then swallowed and retained in a fermentation chamber known as the rumen. Rumen contents can then be regurgitated for more mechanical treatment with the teeth and re-swallowed. Regrinding of the relatively large initial bites taken by the animals creates small particles that can be easily attacked by symbionts living in the digestive tract. This action leads to post-feeding behavior of repeated regurgitation and re-chewing that has come to be known as "chewing the cud" or "rumination." Animals that practice this behavior have come to be known as "ruminants," presumably because an animal appears to be considering the state of the world (i.e. ruminating) as it chews and re-chews stomach contents between feeding bouts. Evolutionarily, this behavior is thought to have been selected because it allows animals to take in a large volume of plant material while feeding in areas where they are vulnerable to predation, then process their food in more secure areas.

The Infra-order Pecora includes species which have four-chambered stomachs. Most have boney protrusions, horns or antlers, from the frontal bones. These horns may be limited to one gender or occur in both and may be deciduous or permanent. Mountain reedbuck have been placed in the Family Bovidae (cows, sheep, goats, antelope, buffalo), Sub-family Hippotraginae (Old-World antelope with permanent, ridged horns that may curve but do not form "lyre" or spiral shapes [Sclater 1900]). Three species are recognized in the genus Redunca. Only males in this genus bear horns.

The southern or common reedbuck ( _R. arundinum_ ) (Fig. 2) is the largest member of the genus, with males weighing up to 180 pounds or ~80 kg (Kingdon 1982). It occupies grassland, savannah, and open woodland across southern Africa. The species is not well adapted to deserts, dense forest, or broken terrain. Bohor reedbuck ( _R. redunca_ ) (Fig. 3) occupy a band of savannah, grassland, and open woodland across East Africa, north into Ethiopia and west almost to the Atlantic coast. The range of Bohor reedbuck overlaps that of southern reedbuck in East Africa. Males of this species are reported to weigh up to 55 kg. The mountain reedbuck is the smallest species in the genus (males reported up to 84 pounds or 38 kg) and have the shortest horns. Mountain reedbuck, as the name suggests, tend to occupy open woodlands and grasslands associated with hills and mountains. The other Redunca species occupy more-or-less contiguous ranges while mountain reedbuck sub-species are found in disjunct ranges separated by hundreds of miles.

Figure 2. Southern or common Reedbuck (Redunca arundinum)

from iSimangaliso Wetland Park, KwaZulu-Natal, South Africa,

2019. Posted by Bernard Dupont on Wikimedia Commons

(https: //commons.wikimedia.org/wiki/).

Figure 3. Bohor reedbuck ram in the Serengeti National Park, 1971.

Three sub-species of mountain reedbuck have been identified (Fig. 4). Southern mountain reedbuck ( _R..f. fulvorufula_ ) are the largest and, at the time of European colonization, occupied broken terrain from the Cape of Good Hope north to southern Botswana (du Plessis 1969), generally east of the twenty-third longitude. Chanler's mountain reedbuck ( _R. f. chanleri_ ) are slightly smaller and, in the nineteenth century, occupied hills and mountains from Tanzania north of the Zambezi River through Kenya and Uganda and northward into Ethiopia and probably Sudan (Stevenson-Hamilton 1947, Bere 1958, Stewart and Stewart 1963, Dorst and Dandelot 1970). The third sub-species, Adamawa (also spelled Adamaoua) or western mountain reedbuck ( _R. f. adamauae_ ), is limited to the Adamawa Plateau in Cameroon. It is the smallest of the three sub-species and has the most restricted distribution. It was only proposed as a separate sub-species in the 1960s (Pfeffer 1962).

Figure 4. Range of 3 mountain reedbuck (Redunca fulvorufula) sub-species

reported in the IUCN Red list, 2017. The southern sub-species

(R. f. fulvorufula) is found in South Africa. Chanler's mountain reedbuck

(R. f. chanleri) occupy highlands in Kenya, Tanzania, and Ethiopia. The

western (also known as Adamawa or Adamaoua) sub-species (R. f. adamauae)

is restricted to northeastern Cameroons near the Nigerian border. The figure

was taken from the IUCN website ( <https://www.iucnredlist.org/species/>

19391/50193881).

This whole long-winded discussion of classification only serves to point out that mountain reedbuck have an unusual distribution pattern. The sub-species are separated by large areas that contain apparently suitable habitat. Gaps between mountain reedbuck sub-species are occupied by closely related species (Southern reedbuck in some areas. Bohor reedbuck in other areas), which suggests that sub-species of mountain reedbuck could have evolved from either southern reedbuck (for the southern mountain reedbuck) or Bohor reedbuck (for the Chanler's and western sub-species). The range overlap between each mountain reedbuck sub-species and another reedbuck species indicates _Redunca fulvorufula_ is a "good" species (i.e. can maintain itself with minimal gene mixing in the presence of related species). This implies that all of the sub-species evolved from a single ancestral mountain reedbuck then developed into sub-species due to historical geographic or ecologic isolation. When I went into the field, either hypothesis was viable.

In 1969, the literature on mountain reedbuck was sparse. Rough ideas of distribution were available. Physical descriptions (short, curved basally ringed horns on males, grayish bodies with reddish necks, prominent bare patch below ears, etc.). Imprecise information on social organization (small groups with a single adult male and several females and young) appeared in a few scientific articles and natural history books. Horn measurements were available from Rowland Ward's hunting records (https://sportsafield. com/rowland-ward-records-of-big-game-30th-edition/). Habitat preferences (broken terrain with some surface water, grassland to open woodland) were vaguely described. Food was thought to be predominantly grasses. The most frequently described characteristics were behavioral. When alarmed, mountain reedbuck issue a high-pitched whistle, raise their tails to expose a white ventral surface, and flee with a distinct "rocking-horse" gait.

At this time, the lack of attention to this species was understandable. Mountain reedbuck generally limited themselves to habitat that had little attraction to resident tribes or to colonizing Europeans. They did not provide spectacular trophies for hunters. They seldom, if ever, occurred in densities that threatened livestock forage or crops. They weren't known to carry virulent diseases that were transmissible to humans or livestock, other than the common pathogens and parasites that were common to all domestic and wild ungulates in Africa. They occupied areas that were difficult to hunt and presented a relatively small package of moderately palatable meat to hunters who did seek them. Scientists at the time were interested in studying showier animals, animals that presented threats to human interests, or animals that appeared to be in decline due to human activities. This is the familiar and understandable "squeaky wheel" approach to biology. There was no imminent threat to the species so, in the interest of improving relationships between the United States and the Republic of South Africa, no one objected to turning a skinny twenty-one-year-old foreigner loose in the wilds of Africa. From Texas A&M's standpoint, it was a relatively inexpensive project that more-or-less fit in with the interests of a large donor in Texas. From the University of Pretoria's standpoint, it was an opportunity to strengthen research bonds with academia in America. At the time, the South African policy of "apartheid" (apartness) was isolating South Africa from the scientific community in the West. From my standpoint, I got to play "Rama-of-the-Jungle," a long-time dream.

I was fortunate to receive funding from several sources. A stipend designed to cover basic field expenses for the first two years was provided by the Mammal Research Unit of the University of Pretoria. The Transvaal Nature Conservation Division funded living quarters, radio-telemetry equipment, and field crews for capture of animals. The staff of the Loskop Dam Nature Reserve furnished field technicians when needed and collected animals for my study. The Caesar Kleberg Research Program in Wildlife Ecology at Texas A&M University provided a stipend to support me in the field during the first three years of the study and equipment and field assistance in Kenya. The National Science Foundation covered stipend expenses after I returned to Texas A&M. Access and aid with data collection were also available from South African National Parks (now SANPark, a division of the Department of Environmental Affairs), the Natal Parks Board, and private land owners (L. van Rooyen, and A. Cole). Advice, direction, supplemental information, and advocacy for my project was provided by a number of individuals (J. Teer, W. Meester, L. Theron, S. Qvortrup, L. Blankenship, S. Smit, T. Smit, S. du Plessis, K. Bothma, J. Steencamp, P. Barnes, P. Gaymer, U. Pienaar, and many others). A one-man research project does not succeed without a lot of help.

I was in South Africa from July 1969 through August 1971 and again from June – August 1973. I spent September 1971 through March 1972 in East Africa. Due to archaic rules in effect at Texas A&M in the 1970s (if you skipped a master's degree, you had to take the equivalent of one in coursework), a stint in the U.S. Army to satisfy my military obligation, and my own slow pace as a writer, I didn't finish the Ph.D. until May 1976. I had access to scientific publications up to 1976 as I wrote my dissertation. Given the slow slog from fieldwork to analysis to publication, this means that when I wrote the dissertation the most recent information to which I had access was collected in the late sixties to early seventies. So I guess this memoir could be more accurately titled "a forty-five or so year retrospective on mountain reedbuck." Of course, at my normal writing pace, I might not finish this tome until fifty years after my dissertation was finished.

# Chapter 3 – Information Available in 1969

At the time I arrived in South Africa, the taxonomy of mountain reedbuck was as settled as it was likely to get in "pre-DNA" days. Most classification schemes for Artiodactyla basically followed Simpson (1945). The taxonomy of Bovidae (cow-like, antelope-like, sheep-like, etc. animals) was fiercely debated within taxonomic circles but unresolved. Depending on which system you supported, mountain reedbuck were placed in the sub-family with "horse-like" antelopes (Hippotraginae), not because they were closely related to horses or looked like horses, but species in this grouping were more horse-like than they were cow-like or gazelle-like or goat-like. Or they were classified as Reduncinae, a sub-family with only two genera. Whichever way you leaned, the mountain reedbuck was one of only three species in the genus Redunca. Southern reedbuck and Bohor reedbuck were known for their association with wet grasslands and marshlands (hence the name "reedbuck"). Mountain reedbuck were known to be associated with hilly or mountainous grassland or savannah (Roberts 1951, Astley-Maberly 1963, Dorst and Dandelot 1970).

Information collected prior to 1970, dealt primarily with physical descriptions and range. Dorst and Dandelot (1970) give a good, if brief, physical description in English measurements. To paraphrase: the mountain reedbuck is a small antelope, standing 24 to 30 inches (61-76 cm) at the shoulder. Its coat is "woolly" without distinct markings, gray on back and sides, white on the ventral side, and rufous on the neck and head. A distinct dark bare patch is visible below each ear. Tails are short and bushy with white undersides. Males have short horns (7-10 inches or 18-25 cm), ringed at the base with evenly-curved, un-ringed distal ends with no hooks at the top. Male necks may be more rufous than females, but except for the presence of horns in males, there is little sexual dimorphism.

Dorst and Dandelot (1970) also provide simple cues for distinguishing mountain reedbuck from other reedbucks and superficially similar antelope in other genera. Both the southern and Bohor reedbucks are larger, have similar-shaped but longer horns on males, and lack the dimorphic color scheme on the body. Both species tend towards a light brown coat. All species in Redunca have a bare dark patch below the ear. They record the southern reedbuck as standing 33-37 inches (85-94 cm) at the shoulder, weighing 120-170 pounds (55-77 kg), and horn length in males as 15-18 inches (38-46 cm). Bohor reedbuck are described as standing 27-35 inches (69-77 cm) at the shoulder, weighing 80-110 pounds (36-50 kg), and bearing horns on males that are 10-17 inches (25-43 cm).

Two other species, grey rhebok ( _Pelea capreolus_ ) and oribi ( _Ourebia ourebi_ ), overlap in range, occur in similar habitat to mountain reedbuck, and might be mistaken for them in brief sightings (Dorst and Dandelot 1970). Grey rhebok (Fig. 5) are roughly the same size as mountain reedbuck (30-inch or 76 cm shoulder height) but more slenderly-built (weight 50 pounds or 23 kg). They have a uniform grayish-brown coat color that may trend towards rufous on the neck. Males have basally-ringed horns with a slight forward curve (8-12 inches or 20-30 cm). Oribi (Fig. 6) are smaller (20-26 inches or 51- 66 cm at the shoulder and weighing 20-45 pounds or 9-20 kg) and more lightly-built than mountain reedbuck. Males have short (6-8 inch or 15-20 cm) straight horns. Oribi are uniformly tan, but they do raise and spread tails with white ventral surfaces when alarmed, as do mountain reedbuck. Oribi have bare glandular patches below the ear while grey rhebok do not.

Figure 5. Adult male Vaal (or grey) rhebok (Pelea

capreolus) in Karoo National Park, Western Cape,

South Africa, 2018. Posted by Bernard Dupont on Wikimedia

Commons (https://commons.wikimedia. org/wiki/).

Figure 6. Adult male oribi (Ourebia montana) in Sudan, 2008.

Posted by Bernard Dupont on Wikimedia Commons

(<https://commons.wikimedia.org/w/index>.).

Descriptions of the distribution of sub-species prior to my study are relatively abundant. Naturalist/explorers and wildlife managers hired by various government organizations provided reasonable records of where they saw various species (Stevenson-Hamilton 1947, Roberts 1951, Rand 1955, Bere 1958, Bigalke 1958, Bigalke and Bateman 1962, Kettlitz 1962, Pfeffer 1962. Stewart and Stewart 1963, du Plessis 1969). During the middle of the 20th century, the three sub-species of mountain reedbuck had distributions assumed to be similar to those in the nineteenth century. There were concerns about many local populations (Rand 1955, Bere 1958, Bigalke 1958, Bigalke and Bateman 1962, Kettlitz 1962, van Ee 1962, Vincent 1962, Stewart and Stewart 1963, von Richter et al. 1972), and mountain reedbuck were regarded as "common game" only in the Cape Province of South Africa, but there was little concern for (or knowledge of) the population status for any sub-species as a whole.

Beyond a few natural history notes, very little was known in the scientific community about mountain reedbuck ecology in 1969. Mountain reedbuck were reported to live on mountain and hill slopes in small groups consisting of one adult male and several females and young (Sclater 1900, FitzSimons 1920, Hewitt 1931, Roberts 1937). Group sizes of 4 to 30 had been reported for the southern sub-species (Sclater 1900, FitzSimons 1920, Hewitt 1931, Roberts 1937, Stevenson-Hamilton 1947), up to 50 for the Chanler's sub-species (Simon 1962), and up to 12 for the western sub-species (Pfeffer 1962). Even though mountain reedbuck were reported to occupy broken terrain covered by a wide variety of vegetation types across a wide range of elevations, all sub-species were believed to feed primarily on grasses (Sclater 1900, FitzSimons 1920, Stevenson-Hamilton 1947, Roberts 1951).

Mentis (1972) summarized information available on the reproductive biology prior to 1970. His summary was largely based on anecdotal information and reports of explorers and zookeepers. In the Afrikaans tradition of referring to the young of large antelope as "calves" and small antelope as "lambs," he described the lambing season for southern mountain reedbuck as extending from October through December based on descriptions given by Sclater (1900), FitzSimons (1920), and Astley-Maberly (1963). Pienaar (1962) reported a broader season (September – February). Births in South African zoos (Brand 1963) have been reported in all months except May and June with a November through February peak. No multiple births are mentioned by any author. No information on age of reproductive maturity, gestation length, or reproductive potential was available in 1969 although a few authors speculated that the reproductive characteristics of mountain reedbuck might be similar to related species.

There were a few studies starting or underway in 1969 that provided me with insight into the biology of mountain reedbuck for my own study. Veterinarians from the Onderspoort Research Institute were engaged in a series of studies of antelope parasites. Baker and Boomker (1973) used specimens from my study to produce a detailed list of mountain reedbuck parasites. Hofmann (1972) described the sub-auricular gland (the dark bare patch below the ear) and its importance to mountain reedbuck as a scent radiator. Hofmann and Stewart (1972) did anatomical studies of mountain reedbuck alimentary tracts that explained their specializations for a graminoid (grass and sedge) diet. Drevemo et al. (1974) used blood collected from animals harvested in my study to produce the first blood chemistry results for the species. Information, published and personal communications prior to publication, collected by Jungius (1971), Mason (1977), Rowe-Rowe (1983), Theron (1973), and others on ecology of related species and vegetation of study areas where I worked were also useful for my output. As with most areas of science, you don't row the boat by yourself.

# Chapter 4 – Grand Design

When I arrived in South Africa, I thought I had spent two days on a plane only to wind up back in Amarillo. The "High Veld" of South Africa looked altogether too much like the high plains of Texas. I had expected wild country. I found a big city (Johannesburg), ugly mine tailings (the Witwatersrand), paved roads, plowed fields, and, at least in White suburbs, housing developments that could have been transplanted from Houston. I couldn't wait to get "out in the field." As things turned out, I was stuck at the University of Pretoria for nearly a month while final logistic details were worked out. It was not time wasted. I went through local libraries to find more material on mountain reedbuck (not much) and spent time talking with professors, government personnel, and graduate students (worth a lot).

South Africa was definitely more civilized than I had envisioned. I also had not envisioned the scope and implications of apartheid. Having been raised in the South, I knew about racism. But I had no idea what knots people could tie themselves into when racism was carried to extremes. I guess I took the coward's way out. Most white South Africans I met were friendly and helpful, basically good people like Southerners I had grown up with, except for that race thing. My study would have been impossible without their help, and I would have been had a lonely existence without the friends I made there. I didn't interact much with local tribes because it was discouraged by Whites, and I had little in common with tribal cultures. I tried to learn a couple of tribal languages but was never able to become fluent. Early in the game, I decided that I was in a country where I wasn't a citizen, didn't have a vote, and had best mind my own business. I was there to study wildlife not engage in politics.

By the time I was given the go-ahead to get to the field, I had a general game plan: I wanted to learn everything I could about the ecology and biology of mountain reedbuck. Executing this plan was more difficult than I had envisioned. My operational budget consisted of $200/month in a stipend from Texas A&M, $200/month for expenses (gas, equipment, a vehicle, etc.) from the University of Pretoria, a free room at a provincial nature reserve, and unspecified assistance promised for help with collecting data from the Transvaal Provincial Administration. I drove to Loskop Dam Nature Reserve in a GMC pick-up loaned to me by the University of Pretoria. I was disappointed that the whole trip was on paved roads through farm country, not pristine African wilderness. I was faintly proud of myself for staying on the left side of the road the whole ninety miles. In fact, the only time I screwed up was pulling into the entrance – I pulled right to the exit rather than left to the entrance.

I spent the first afternoon meeting reserve staff and settling into my housing, a thatched-roof rondavel. The "hut" was 50 yards from the swimming pool, tennis courts, and restaurant and came with daily maid service. This was quite posh compared to my lodging in East Africa, a tent on the back side of Arthur Cole's ranch. Although even in Kenya, I lived comfortably (I was only 300 yards from a volcanic spring where I could take warm baths and get fresh water, after it cooled).

Staff members gave me an orientation tour that covered all of the roads in the reserve, about forty miles. Loskop Dam Nature Reserve was a provincial game reserve formed from nine ranches bought purchased between 1942 and 1966 (Theron 1973) surrounding an irrigation reservoir on the Olifants River (Fig. 7). While I was there, the reserve included a little less than 32,000 acres (12,762 ha) surrounding a 4,200-acre (1,700 ha) reservoir. Basically, it was a bowl of rugged hills around a river valley with irrigated citrus and other crops grown downstream from the dam. Mountain reedbuck were in the reserve when it was founded and were one of the most numerous ungulates present in 1969. Beginning in the 1940s (du Plessis 1955), extirpated species were translocated from other areas so that by 1970, 18 ungulate species were present, ranging in size from steenbok ( _Raphicerus campestris_ ) to white rhinoceros ( _Ceratotherium simum_ ). Eleven of these species were gramivores (grass-eaters) that could potentially compete with mountain reedbuck for forage. Management strategies for smaller provincial reserves in South Africa at the time precluded re-introductions of hard to manage species such as elephants ( _Lexodonta africana_ ) and large predators. The reserve was enclosed with a 7-foot (2.1-meter) woven-wire fence that discouraged, if not completely prevented, egress by large animals such as buffalo ( _Syncerus caffer_ ), white rhinos, and hippopotami ( _Hippopotamus amphibious_ ). Several medium-sized predators, the species most likely to prey on mountain reedbuck anyhow, were present when the reserve was formed including leopards ( _Panthera pardus_ ), caracals ( _Felis caracal_ ), brown hyenas ( _Hyaena brunnea_ ), and black-backed jackals ( _Canis mesomelas_ ). Chacma baboons ( _Papio ursinus_ ), a potential predator on lambs, were abundant. Rock pythons ( _Python sebae_ ), another potential lamb predator, were present. There were a few crocodiles ( _Crocodylus niloticus_ ) large enough to eat any mountain reedbuck. The staff also pointed out that black mambas ( _Dendroaspis polylepis_ ), green mambas ( _D. angusticeps_ ), puff adders ( _Bitis arietans_ ), and spitting cobras ( _Naja nigricollis_ ) were abundant so walking around bare-footed at night was not advisable. Loskop was not a natural wilderness by any stretch of the imagination, but it met my expectations much better than Pretoria.

The second morning at Loskop, I was on my own. The first encounter with strange wildlife almost sent me scurrying back to my truck. Baboons make a hell of a lot of racket. When I decided that monkeys didn't present any imminent danger, I started searching for mountain reedbuck. I think I finally saw the first one a day later, after the reserve manager assigned a native worker to accompany me. He didn't speak any English. I didn't speak any Ndebele or Afrikaans. Using hand signs, he did guide me into areas where mountain reedbuck were common. I was off and running!

Figure 7. Overview of Loskop Dam Nature Reserve, South Africa, 1969.

After a few weeks, I began to wonder if the mountain reedbuck population at Loskop was typical. Many, if not most, ungulate species vary in population characteristics, habitat preferences, and even social organization in different areas of their range. I decided I needed to visit other areas to see what variation local populations might exhibit. I was eventually able to survey six areas in South Africa and three in East Africa (Fig. 8). Given limited time, low reedbuck densities, and my logistic constraints, most of these areas were not covered well. At suggestions from academic advisors and co-workers I also was able to obtain help in collecting animals (basic biology, condition, reproduction, parasites, food habits, etc.), capturing and marking animals (both visually identifiable collars and early radio-telemetry), testing census techniques, and raising captive animals (growth rate, reproductive maturation). In retrospect, I probably spread myself too thin, but it was fun.

Figure 8. Locations of mountain reedbuck study

areas in South and East Africa, 1969-1973

# Chapter 5 \- Behavior Patterns

Science tends to follow fads. In 1969, studies of behavior (ethology) were all the rage. Students all over the world were watching every conceivable variety of animal and dutifully recording activity patterns and behavioral interactions. I joined the pack. Sitting behind a telescope with a notepad and pencil fit my budget well. Unfortunately, I was ignorant of sampling protocols that reduce interdependence of individual sample points, and I discovered that mountain reedbuck were difficult to sample either randomly or consistently. I wound up oversampling early morning hours and under-sampling mid and late day hours. After active morning feeding, reedbuck frequently moved into brushy areas and didn't reappear until close to dark. I did use radio-telemetry and track counts to obtain crude data on nocturnal activity levels and movements. Most of my observations at Loskop (South Africa) and the Cole Ranch (Kenya) were made from specific observation points where I had a reasonable chance of observing one or more groups consistently. In Loskop, these sites included places where I had a chance to observe collared animals as well as uncollared animals. When reedbuck were visible, I noted numbers, age, gender, and activity at 5-minute intervals. I continued this approach for over two years at Loskop and six months on the Cole Ranch. I did not spend much time on observation of activity at the other study sites.

In October 1969, Transvaal Nature Conservation Department crews captured fifteen mountain reedbuck (6 adult males, 9 adult females) in the Loskop Dam study area using drive-nets (Fig. 9). These animals were used for behavioral observations and to test census techniques. All were fitted with individually recognizable plastic neckbands. In February and March 1971, five mountain reedbuck (3 adult females, 1 immature female, 1 immature male) were captured using drive-nets and fitted with radios manufactured by and tracked with a receiver provided by the South African Council for Scientific Research (Anderson and de Moor 1971). The antenna system included a monopole antenna and a double-yagi vehicle-mounted antenna (Anderson and Hitchins 1971). After pounding through the brush and never succeeding in seeing a collared animal without flushing it, I settled on two strategies for tracking radio-collared animals: 1) triangulating with fixed radio-beacons (a radio transmitter hung in a tree at a known location, a radio antenna on my vehicle parked at a known location, the azimuth to a radio on an animal); and 2) parking at a known location relative to a radio-beacon then periodically relocating animals detectable from that location to see if radio-collared animals moved up, down, or along a facing slope. I attempted to locate multiple animals (3-5 depending on how many radios were functioning) at 4-hour intervals over a 72-hour period each month. I also carried out four 12-hour night observations or two 24-hour observations on one or two animals each month to record "apparent activity" every thirty minutes. This allowed me to determine when animals moved towards surface water.

Figure 9. Capturing mountain reedbuck in the Loskop Dam Nature Reserve, South Africa, using drivers and lines of collapsible nets.

Track counts at Loskop were unreliable, given the wide variety of ungulates and the rocky terrain, but I was able to take advantage of daily drawdowns of the reservoir along one stretch of the shoreline to get an estimate of how often mountain reedbuck drank surface water during the dry season of 1970. I knew how many groups occupied a slope above a 1.8-mile (2.9-km) stretch of silty shoreline on the south shore of Loskop Reservoir. I also knew there were few other antelope species of similar size to mountain reedbuck in this area. Drawdown rates were relatively constant (1 to 6 ft or 0.3. to 2.0 m per day) (Fig. 10). I cleared tracks along the shoreline on 19 August and walked the stretch nine times between 20 August and 1 September, clearing tracks each time. I followed each track set back to the vegetation line to minimize recounts of animals taking more than one drink. I had to abandon the counts after September 1st because of the quagmire created by my tracks and those of an ostrich (Struthio camelus) group that persisted in following me.

Figure 10. Loskop Reservoir in dry season showing draw-down lines, August 1970.

Over the course of the study in South Africa, I recorded 7,714 visual activity records (Irby 1981). In Kenya, I recorded 2,819 activity notes (Irby 1982). In summary, mountain reedbuck are not any more active than they have to be. Roughly 80% of their activity involved grazing. Peaks occurred during the first and last hours of daylight (Fig. 11), appeared to be similar in males and females (young lambs generally were inactive most of the time), and apparently increased, at least during daylight hours, during the dry winter season in South Africa. This may have been a response to more moderate mid-day temperatures or an attempt to process more forage as quality decreased through the dry season. Drops in temperature associated with approaching rain often stimulated the animals to begin grazing. Intense rainfall led them to turn their tails towards the blowing rain, lay down, or move into cover provided by terrain or vegetation.

Figure 11. Observed frequency of activity for mountain reedbuck in the Loskop Dam Nature Reserve during daylight hours, 1969-1971. Percentages are based on 7,714 observations.

Radio-telemetry data indicated they were also active at night (Fig. 12). I was able to use variation in radio signals associated with head movements to separate animals moving their heads from those not moving their heads, but I could not tell the difference between animals feeding or just grooming or shaking heads to dislodge insects. If head movement was correlated with feeding activity, reedbuck in fall and early winter were less active during night hours than early morning or late afternoon but more active than in mid-day hours. There appeared to be a midnight peak in feeding during late winter but that may have been an interaction with moonlight.

Figure 12. Results of radio-telemetry observations of apparent diurnal and nocturnal activity for mountain reedbuck in the Loskop Dam Nature Reserve, March – August 1971

Other behavioral patterns (grooming, defecation, urination) occupied a small portion of their time, except head shaking in seasons with high fly densities. I never observed reactions to predators in South or East Africa. I presume the reedbuck would whistle and bound away into rough terrain as they did when I startled groups. Mountain reedbuck males were territorial in every population I observed. They would occasionally tolerate males from adjacent territories in Loskop. Adjacent territorial males were reasonably tolerant of one another in the Cole Ranch, and I saw a few instances of adult males passing through territories of other males at Loskop. The trespassers appeared to adopt submissive postures. Encounters between neighbors were ritualized (whistling, stiff postures, _stotting)_. I saw no obvious marking of vegetation or use of fecal piles or urine to delineate territorial boundaries. If Hofmann's (1972) idea that the sub-auricular bare patches on mountain reedbuck heads served as scent radiators, they could have marked territories using only scent.

Adult male tolerance of male lambs decreased starting at about six months of age. Territorial males would pursue older lambs, sometimes relentlessly. Male lambs persisted in trying to return to their mothers; territorial males persisted in chasing them. Tolerance of immature males

seemed to increase with population density. I only observed 5 of 153 territorial group sightings with yearling males at Loskop versus 53 of 349 in the Cole Ranch. Pairs or groups of males were also more common in the Cole Ranch than at Loskop (16 versus 4). I seldom saw adult males fighting, but it undoubtedly occurred. In one group in Kenya, a male had escaped from a wire snare, breaking the wire near its attachment point on a fence. I saw him interact with another male, using the wire as a whip to gain an advantage. The snare fell off before I saw if this odd use of a "tool" had any reproductive advantages.

I seldom saw mountain reedbuck drinking (<1% of activity notes in all seasons), but there was a higher incidence (0.8% vs. 0.1% of observations) in early morning drinking during the late dry season than in the wet or early dry season at Loskop. Surface water was relatively widely distributed in the Loskop study area except in the late dry season. Radio-telemetry at Loskop suggested animals might move a half-mile or more (0.3 – 1.5 km) to surface water during the dry season and drink on alternate days. Track counts during the late dry season of 1970 at Loskop indicated that resident mountain reedbuck that had to move less than a half-mile (<0.8 km) drank 0.4 times/ 24 hours (cloudy, cool conditions) to 1.5 times/24 hours (hot, dry days). All of these estimates are based on small sample sizes, and, for the track counts, duiker ( _Sylvicapra grimmia_ ) tracks could have been confused with mountain reedbuck tracks. The only mountain reedbuck I observed as far as 3 miles (5 km) from surface water were on the Akira Ranch in Kenya. This observation occurred during the short rains so pools or livestock watering tanks that I missed may have been available. I have no idea if this group would have remained so far from water during the dry season.

Observed reproductive behavior was more frequent than water use. I saw six full copulatory sequences and 50 partial sequences. The behavior patterns were similar to those described for other African antelope. Males investigated female receptivity by sniffing the vaginal region and using the flehmen (lip curl) to test urine. If the female stood for the male, he would begin kicking between her hindlegs with a foreleg. If she was receptive, she would stand for copulation. If not, she either lay down or ran away. Sometimes males were persistent and followed. Other times they left the female alone. Copulation was brief, although I saw one male attempt 43 mounts over 5 minutes before he ejaculated. Males seemed more relaxed than females afterwards, but both genders usually resumed grazing within a minute or two. Male sexual activity was spread through the year with an indistinct peak in the autumn (March – April) in South Africa. Lowest levels occurred in the winter dry season and again in what appeared to be the peak of the parturition season, mid-summer. On the Cole Ranch in Kenya, males showed interest in females through the whole period I was in the field (September 1971 – March 1972). I observed young lambs in Loskop throughout the year with an indistinct summer–autumn peak. Observations of other populations in South Africa suggested a narrower summer peak. My estimates of lamb age for mountain reedbuck observed in the Cole Ranch in Kenya throughout the September – March study period indicated a possible peak during the March – May rainy season. Lambs were hidden at birth in all populations and gradually became more active until their activity patterns resembled those of adults at >2 months of age. Play was much more common in lambs up to one year than in adults.

Copulation of an unmarked male with a collared female gave me my best estimate of the gestation period for the species. Prior to my study, gestation length was unknown but assumed to be similar to common reedbuck, a much larger species. On March 9, 1970, I observed a complete copulatory sequence (including ejaculation) involving a collared female. I re-sighted her one or more times without a lamb in every month except April 1970. The last sighting without a lamb occurred on October 31, 1970. I saw her nursing a young lamb (less than 2 weeks-old) on November 16, 1970. Assuming the copulation I observed was successful, this indicated a gestion period of 236 – 251 days (7.75 – 8.25 months) which seemed long for a relatively small antelope.

# Chapter 6 \- Social Organization

Literature available when I began this project indicated mountain reedbuck lived in small groups (Fig. 13). Groups were reported to be comprised of an adult male, one to several adult females, and immature animals. I found that social organization could be more accurately described as a three-class system: territorial males (alone or with female groups); non-territorial males (usually immature and alone but occasionally in small groups in high density populations); and female-young groups. I only had access to 12 individually recognizable animals that could be observed reasonably frequently, and those were only in the Loskop population (7 adult males, 5 with recognizable physical features and 2 with collars, and 5 adult females, all collared), so conclusions I reached on social organization must be regarded as tentative.

My observations in Loskop indicated male territories were maintained throughout the year and were large enough to provide forage for the male and apparently for females and young that shared the territory. Minimum sizes for territories occupied by five distinguishable males in the Loskop Reserve varied from 37 to 119 acres (15 – 48 ha). Four of them were adjacent to year-round surface water. Males remained in territories unless forced out by disturbance (human activities, fire, predators, etc.), but I observed a few incidences where males left territories to drink or interact with other reedbuck.

Figure 16. Chanler's mountain reedbuck group in a lava field on the Cole Ranch, Kenya, 1971.

Based on my small sample of distinguishable individual females, it appeared that females varied in the time spent with and fidelity to individual territorial males. One collared female was observed with the same collared male from October 1969 through July 1971 then again with the same male in July 1973. She was never observed with a different male. Another was observed for almost a year with the same territorial male. Other females were more fickle. Three observed during 1969 – 1971 were observed with three or four different males, spending anywhere from a single day to several months with specific males and sometimes switching back and forth between adjacent male territories. When marked females moved from one territory to another, they appeared to move independently of other adult females. Minimum home ranges for five collared females averaged 91 acres (37 ha). I only observed males attempting to keep females in a territory two times. Males occasionally left a group of females untended to pursue a single female (presumably a female in estrus), to drink, or to interact with neighboring males.

Social groupings I observed in the nine populations varied from 1 to >20 (although the larger aggregations may represent more than one group). Reports of groups with up to 30 southern mountain reedbuck (Stevenson-Hamilton 1947) and 50 Chanler's mountain reedbuck (Simon 1962) were likely due to human disturbance. Mountain reedbuck censuses through the 1970s were frequently undertaken by lines of 10 to 50 beaters moving across the census area. Undoubtedly, this level of disturbance would force individual groups to coalesce as they fled from humans. The percentage of sightings composed of a single individual varied from 10% to 41% across the nine populations I observed. The highest percentage was recorded at Loskop, which was not only the most intensively studied population but the most heavily wooded. In many cases I only had a glance of flushing animals so I probably over-estimated the percentage of one-animal groups (if the concept of a "one-animal group" makes sense) and under-estimated average group size. For mountain reedbuck in groups, averages across the nine populations I surveyed varied only from three to five. Populations exposed to a full complement of predators (Kruger National Park, Umfolozi Game Reserve, Akira Ranch, and Ngorongoro Crater Conservation Area) had similar average group sizes as protected populations with reduced top-tier predators (Loskop, Ohrigstad, Giant's Castle). Two ranches which made efforts to eliminate large and some medium-sized predators (van Rooyen Ranch, Cole Ranch) had group sizes similar to protected areas.

Group sizes in the two populations in South Africa I was able to survey through a full year or more showed some variation through the year with smallest group sizes in the late dry season and largest in the early dry season (Fig. 14). This would make sense for a species that produced lambs that remained secluded when young and only joined mothers with groups as they matured. If, as I suspect, mortality peaks in the late dry season, changes in group size might well reflect these losses.

Figure 14. Average size of southern mountain reedbuck groups observed in six populations in South Africa, 1971-198-73.

# Chapter 7 – Carcass Collections

Wildlife managers have used a variety of invasive and non-invasive techniques (Silvy 2012) to determine the basic biology of wild animals and ecological factors that affect populations. The most bang for the buck (i.e. the most information that can be collected per individual) comes from live animals killed for necropsy. The Transvaal Nature Conservation Division and the Kenyan Wildlife Service (through a project funded by the Caesar Kleberg Research Program in Wildlife Ecology) authorized collection of animals for this project. Forty-one mountain reedbuck were shot in the Loskop population between January 1970 and May 1971. Seven additional fresh carcasses from predator kills or accidents were also necropsied. The sample (24 adult females, 16 adult males, 3 immature females. And 5 immature males) included animals taken in all seasons. Animal carcasses were moved to a warehouse I used as a laboratory for necropsy. In the Cole Ranch in Kenya, 21 individuals were shot and one animal was necropsied after it was killed by a car (10 adult females, 2 adult males, 7 immature females, and 3 immature males). This sample includes 2 killed in June 1972 and necropsied by Lytle Blankenship. Specimens from the Cole Ranch were examined at the kill site. We attempted to recover as much information as possible from each carcass, which was somewhat limited given the primitive lab facilities available. Information ranged from body measurements to condition indices to food habits to reproductive condition to age indices to parasite loads. Where possible, I preserved biological specimens for later examination (rumen contents, parasites, ovary and testicle samples, blood samples, mandibles for aging, etc.). Three carcasses and four complete digestive tracts were examined by personnel from the Veterinary Research Institute at Onderstepoort (then in the Transvaal Province of South Africa).

In addition, I collected remains of animals that were not fresh enough for necropsy. From August 1969 through August 1971, 48 mountain reedbuck mortalities were located in the Loskop Reserve. I found remains of 14 mountain reedbuck that were too decomposed for necropsy on the Cole Ranch between September 1971 and March 1972. Cause of death, probable season of death, age, and gender were determined for all of these finds where possible. My finds at Loskop were supplemented by records maintained by reserve staff. Neither I nor staff members made any systematic searches for carcasses, but 268 dead mountain reedbuck were reported from 1961 – 1973, including those I located and those picked up by reserve staff.

Body weights of mountain reedbuck from Loskop showed a distinct seasonal pattern (Table 1), as did condition indices such as the kidney fat index (mass of kidney divided into mass of fat surrounding kidneys). Lightest weights and lowest kidney fat indices occurred in the late dry season (July - October) for adult males and females. Body weights and kidney fat indices in adult males were highest during the early dry season (March-June). Female kidney fat indices peaked in the early dry season, but body weights for adult females were similar during the wet season (November-February) and early dry season (March -June). Adult Chanler's mountain reedbuck were taken before, during, and after the "short rains." Vegetation remained green and surface water was widely distributed during the whole September through March collection period. Body weights were similar to southern mountain reedbuck collected during the late dry season.

Immature animal weights varied with estimated age. I was unable to distinguish distinct annuli when I sectioned molars, but the irruption pattern of molars indicated full adult dentition was present at about two years of age. Immature animals collected indicated males and females reached the range of adult body weights at between 15 and 20 months of age in Loskop and the Cole Ranch. Females evidently reached reproductive maturity around the age of one year. Three immature females collected in Loskop (estimated age range 4 – 12 months) did not have mature follicles. Ovaries of one lamb (estimated age 4 months) collected in the Cole Ranch showed little ovarian activity. Four of the six yearlings collected there contained mature follicles or corpus lutea. Two were pregnant. A small sample of males <12 months of age (four from Loskop and one from the Cole Ranch) had much smaller testes than adult males. Three males (two from Loskop and one from the Cole Ranch) estimated to be yearlings from horn length and shape and dental characteristics had testes within or approaching the adult male range. One had sperm in the epididymis

Table 1. Body mass and kidney fat indices for adult (estimated 2 years of age or older) mountain reedbuck collected in the Loskop Dam Nature Research and Cole Ranch, 1970-1972. Means are live weights minus blood loss. KFI = fat around kidney/kidney weight and is an indicator of physical condition (higher = better condition).

I could not determine the ages of adults, but I was able to divide them into classes based on tooth wear. Mandibles examined from Loskop were clustered in heavier wear classes than those from the Cole Ranch. This suggests that the age structure of the Loskop population was more skewed towards older animals than the Cole Ranch population or that tooth wear patterns were different in the two populations. I should note that soil type and physical characteristics of cells in the plant species eaten by herbivores can definitely influence tooth wear.

Reproductive status and estimated conception dates for fetuses in yearling and adult females collected in Loskop suggested an indistinct peak in conception in the early dry season (autumn) and a peak in lambing in the wet season (summer). Weights of testes in males were consistent with this pattern. Two of the three females collected in Loskop with fetuses <100 g were lactating. Fetuses were not found in five lactating females from Loskop, but ovarian activity indicated they were already primed for the next estrus (developing follicles or corpora lutea). Reproductive status of females collected in the Cole Ranch indicated a high incidence of lamb production but no strong seasonal peaks. Only one adult female (from the Cole Ranch) showed no indication of recent reproductive activity.

Rumen samples from animals collected in Loskop and the Cole Ranch verified that mountain reedbuck feed heavily on graminoids. In Loskop, grasses and other monocots made up more than 90% of diets in all seasons. A comparison of frequency of plant fragments in rumens collected from mountain reedbuck with vegetation measurements made by Theron (1973) indicated grasses were eaten in greater than expected proportions in all seasons. Vegetation measurements indicated that use of graminoids was greater than their abundance in the ground stratum. Figure 15 illustrates this pattern, although it involves "apple versus orange" comparisons (frequency of monocots and dicots in rumen samples versus relative importance of the two plant groups in vegetation transects measured by Theron [1973]). A small-scale test of protein content of seven common grass species sampled during the late dry season at Loskop were all below the 7% minimum designated as maintenance level for livestock. Mountain reedbuck could mitigate for this deficiency somewhat by selective grazing, but the decline in condition and carcass weights in the late dry season indicated that the Loskop population could not completely overcome quality deficiencies.

Figure 15. Relative frequency of monocots (grasses and grass-like

plants) and dicots (forbs and low-growing shrubs) in rumen samples

versus relative frequency of monocots and dicots in the ground

stratum measured in vegetation transects in areas where mountain

reedbuck were collected in the Loskop Dam Nature Reserve, 1969-1971.

Vegetation measurements were made by Theron (1973).

Monocots made up over 90 percent of the diets of Chanler's mountain reedbuck collected in the Cole Ranch. I was unable to do protein analyses for grasses in Kenya, but I did calculate leaf to stem ratios for rumen samples. Over 90% of the grass fragments in my samples were from leaves or leaf sheaths. Less than 8% of the fragments were grass stems. Stems are far inferior in digestibility to leaves.

For samples taken in the Cole Ranch, I had access to a key to epidermal characteristics of grass genera prepared by Svend Qvortrup (unpublished) and data from vegetation transects measured by Lytle Blankeship and Svend Qvortrup. (unpublished and Blankenship and Qvortrup 1974). Using this key, I could identify rumen fragments from 11 genera/species with reasonable confidence (Table 2). A comparison of percent frequency in rumen samples with relative importance of grass availability from sward measurements (another "apples versus oranges" comparison) indicated that Chanler's mountain reedbuck not only selected for the most nutritious parts of grasses, but they were able to selectively choose among grass taxa. (Table 2).

Table 2. Plant categories found in rumen samples from Chanler's mountain reedbuck collected on the Cole Ranch during December 1971 through June 1972. Scientific name for each taxon, common names where known, relative importance of these taxa in the ground stratum, and percent occurrence of fragments from these taxa in rumen samples are given in the table.

Staff from the Onderstepoort Veterinary Institute examined animals we collected at Loskop and found 11 new host records for helminths and one new species of tick. Overall, they identified 18 species of internal parasites, four species of ticks, two species of lice, and sub-cutaneous maggots from parasitic flies ( _Strobiloestrus_ spp.). None of these parasites were regarded as serious threats for epidemics in mountain reedbuck or livestock, and loads were deemed low to moderate in most cases. _Strobiloestrous_ maggots, lice, and ticks were exceptions. Numbers of ticks and lice peaked in the late dry season. Peaks in subcutaneous maggots occurred during the wet season. Parasites from animals collected in the Cole Ranch were generally lower in number than in Loskop and were not identified to species.

Carcasses located in Loskop between 1961 and 1973 peaked in the late dry season (Fig. 16). This may represent a peak in deaths or simply greater visibility associated with lower vegetation cover. Over the whole sample, recoveries of males and females were similar (45% versus 42%). Lamb recoveries were much lower (8%) probably due to smaller size and greater probability that a predator would consume the whole carcass. Probable or likely causes of death were available for only 33 of the carcasses reported. Predators (leopard, caracal, jackal, hyena, feral dogs, and an unidentified snake) were responsible for 23 (70%) of the deaths, accidents for 4 (12%), poachers for 2 (6%), and malnutrition for 2 (6%). I located 14 carcasses on the Cole Ranch in 7 months (September 1971 – March 1972). Cause of death could only be determined for five. Four were likely predator kills and one was a collision with a vehicle.

Figure 16. Sex/Age (top figure) of recovered mountain reedbuck carcasses and

the season (bottom figure) of recovery from the Loskop Dam Nature Reserve,

1961 – 1973. Bars represent percentages of total sample. Numbers of individuals

appear at the top of bars.

# Chapter 8 \- Captive Animals

Very little was known about growth or maturation in mountain reedbuck when I began my study. I obtained permission to capture lambs and hold them in captivity. Obtaining permission was much easier than capturing lambs. Attempts during 1969 yielded one female lamb estimated to be <2 weeks of age. Searches during the rainy season of 1970 yielded one female <2 weeks of age, and one male ~1month of age. The three were held in a 10 x 20-meter pen and provided milk (as long as they would take it), water, and livestock pellets. The male never became tame enough to handle and died after two months in captivity. The two females tolerated my close presence and would allow me to measure them as they were bottle-fed. Measurements were continued on one female for 12 months and the other for 9 (Fig. 17). Hindfoot length reached the adult range at 8 months, body length at 9-10 months, and shoulder height at 8-9 months. I was able to weigh one female to the age of 9 months, and she reached the low end of the adult female weight range by then.

Figure 17. Growth of captive mountain reedbuck lambs from the Loskop Dam Nature Reserve compared to adult females (mean, SD, and range) collected in the reserve.

The female captured in December 1969 no longer tolerated my measurements when she was a year old, but I did keep her in captivity for feeding studies. When she was ~18-months of age, I tranquilized her to see if the third permanent molar had irrupted. I never found out the status of the molar, but I did determine that muscles used in chewing were not affected by the tranquilizer. I still bear scars on my right index finger.

This female also provided information on female reproductive maturity. When she was a year old, an adult male started hanging around the pen. She managed to escape from the pen a week or so later, presumably related to estrus, but returned after a few days. I guess the affair didn't work out, or she preferred living in a predator-free environment where food and water were readily available. I should have sacrificed both of these animals before I left South Africa, but they were basically pets by August 1971. For readers interested in happy endings, they were shipped to another provincial reserve after I left the country where they lived out their lives as pests, eating gardens and landscaping around the reserve manager's house.

I sampled grasses and herbaceous dicots (point-centered quarter technique) in the pen during March 1970, August 1970, November 1970, and December 1970 (after clearing annual dicots from the pen). The pen included 9 taxa of grasses when the first female was released in the pen. Thirteen months later, only 5 taxa remained. Herbaceous dicots increased from 15% of ground coverage in March 1970 to 60% of ground coverage in November 1970.

The captive animals were tolerant of my close presence in the pen so I tried to see how forage over-use would affect dietary choices. Neither female required food from the plants in the pen, but both would begin grazing when I entered. From March 1970 – April 1971, I spent 1-4 hours per month observing the female captured in December 1969 grazing. I classified over 17,000 bites during the 13-month sampling period. It was difficult to record much more than "monocot or dicot" for most species, but two common grass species had growth forms and distributions that allowed reasonably accurate species-specific counts. Herbaceous forbs and woody plants within reach of the female made up <1% of bites recorded through May 1970 then increased to a peak of 30% in August 1970 before declining to 3% in March 1970 (Fig. 18). The peak of forb use coincided with the height of the dry season when no green grass shoots were available. It was obvious that the female selected grass leaves over stems and was selective about which species of graminoids she ate, when choices were available. For the two species I was able to identify as the female fed, Chloris virgate was used in greater proportions than availability in the early dry season and used less than expected most of the rest of the year. Panicum maximum was selected in the rainy season and early dry season but used less than expected in the late dry season (Chi-square tests, P <0.05).

Figure 18. Percentages of grasses in the diet of captive mountain reedbuck and of grasses in the ground stratum of the pen holding the mountain reedbuck, 1970-1971.

# Chapter 9 \- Habitat Use

I was fortunate to have vegetation surveys available for Loskop Dam Nature Reserve (Theron 1973) which allowed me to investigate habitat preferences. With a limited statistical background, I didn't design the study with "modern" statistical approaches, but with help from the Statistics Department at Texas A&M, I did cobble together a multiple regression approach that incorporated census (an estimate of the influence of population and seasonal changes that occurred over the four periods when I sampled large areas of the reserve for census purposes), vegetation type, sample unit size, sub-area within the reserve, an estimate of the impacts of shrub/tree density on visibility, slope class, and time since the last burn. I only used transects I walked for census attempts to limit over-sampling of sites near groups I used in behavioral observation. At the time, the models were "state of the art" even if the sampling design was deficient.

After burning a lot of shoe leather, picking up more ticks than I care to remember, and stepping over more poisonous snakes than I thought healthy (my personal best was two spitting cobras and two puff adders in one afternoon), I had built a model that could explain <10% of the variation in sightings! Both models I ran (one with actual numbers of reedbuck sighted and one with a square-root transformation of the number of groups sighted) were significantly (P<0.01) related to mountain reedbuck habitat use which indicated I had chosen variables that were important to mountain reedbuck, but there were not enough sightings to support elaborate statistical modeling.

For what it is worth, the longer the transect I sampled and the better the visibility (as long as some trees were present), the more reedbuck I saw. Also, numbers varied between sub-units of the reserve and among slope classes (hills were favored over flats). The significance of sub-unit as a variable was not surprising. The seven sub-units I defined had different topographical configurations, were bought in different years, and had different land-use histories. All sub-units had been heavily grazed for up to 100 years prior to purchase and most had been burned annually to encourage growth of new grass. An analysis of variance on terrain character measured from maps indicated that the sub-unit with the highest percentage of valley bottom flats, the greatest elevation change, and the least complex geography had lower densities of mountain reedbuck than sections with lover overall elevation change but more complex hill and canyon systems. The most complex areas were also the first sub-units fenced and stocked with introduced species.

A less ambitious chi-square analysis of sightings in Loskop (a 5-variable classification including: 1] sub-unit within the reserve; 2] dominant vegetation species; 3] density of woody vegetation; 4] slope characteristics; and 5] burn status) yielded more interpretable results. The pattern of distribution was not tight, but mountain reedbuck were most frequently seen on dry slopes (north and east-faces) with moderate tree/shrub density. Fresh burns were uncommon during the period when I observed mountain reedbuck, but I did see a trend towards preference in sub-units that had been burned within a year of my observations. Dense woodland was not common on Loskop and was apparently avoided, although part of the apparent avoidance may have been associated with observer bias (reedbuck were less visible to observers), reduced forage availability (herbaceous vegetation was less abundant under dense tree cover) and/or predator avoidance strategies used by mountain reedbuck (reedbuck rely on vision as well as sound and odor to avoid predators). The dominant tree species did not seem to be important in that mountain reedbuck tended to favor moderately open stands of whatever tree species grew on dry slopes.

I did not attempt to analyze habitat use in the eight supplementary populations I visited, but my overall impression was that mountain reedbuck could occupy habitats ranging from high mountain grassland (Giant's Castle) to volcanic crater/lava fields (Cole Ranch, Akira Ranch, and Ngorongoro Crater), to breaks leading from mid-elevation plateaus to mid-elevation bushveld (Ohrigstad Dam Game Reserve) to low elevation mountain and hill systems (Kruger National Park, Umfolozi Game Reserve, and the von Rooyen Ranch). The important characteristics appeared to be broken terrain, a good grass cover, some tree cover, and surface water during the dry season.

# Chapter 10 - Census Attempts

Counting wild animals has always been a problem for wildlife managers. Counting inconspicuous animals in non-migratory populations that routinely exist at low densities in complex terrain and wooded environments is a major headache. It is difficult to manage any species if you don't know how many critters there are, and you generally need some idea of numbers to determine population trends. Mountain reedbuck at Loskop (and the other populations I visited) met all of the criteria for census difficulty.

In October – December 1969, I attempted to cover the whole reserve (with the exception of a newly fenced plateau grassland with little suitable mountain reedbuck habitat) using belt transects. I walked another randomly selected set of belt transects in September – October 1970 and a replicated set of belt transects in June – August 1973. I also tested an "area-count" technique (sitting at an observation point for a specific length of time, 1 hour, and counting all mountain reedbuck observed in a defined area). Finally, I flew with a team of New Zealand pilots to do a full-coverage helicopter survey of the reserve in 1971.

None of the techniques I employed worked well. The helicopter census was a total bust, yielding <100 sightings. The belt-censuses and area-counts in 1969-1971 produced similar overall population estimates, but the confidence intervals were extremely wide (Fig. 19). Helicopters made too much noise. Mountain reedbuck could move under tree cover well in advance of the machine, and many of the drainages at Loskop did not accommodate the helicopter rotor with reasonable safety margins. Belt-censuses were hampered by investigator bias. I was unable to move through the rough terrain that characterizes mountain reedbuck habitat without making excessive noise, and I sometimes had to deviate from planned routes due to cliffs or encounters with buffalo and rhino. The area-counts produced a high proportion of "zeroes" and were excruciatingly boring. Sitting on a hot slope for an hour, staring across a drainage at another slope while swatting sweat bees is not an entertaining exercise.

Figure 19. Census attempts (population estimates with 95% confidence intervals) on the Loskop Dam Nature Reserve, 1969-1973.

The most exciting incident in all my census attempts did occur when I was using the area-count technique. One afternoon I took a boat (a 17-foot wooden rowboat with a mighty 1.5-horsepower Seagull motor) to a remote section of the reserve. I was sitting on a hill sweating, as usual, when I heard a noise behind me. I looked over my shoulder and saw a black mamba sliding down the rocks ten feet above me. The black mamba is known as a "one-stepper." It is a big snake with abundant highly potent poison. In theory, you die after taking a single step after being bitten. This is likely an exaggeration, but I did not want to test it. I was over an hour from the Loskop headquarters where they might or might not have anti-venom. I picked up a stick, hoping to throw it and use it to distract the snake, but I did not want to move enough to focus its attention on me. In accordance with Murphy's Law (whatever can go wrong, will), I tossed the stick up, shielding my arm movement with my body. The stick hit a branch over my head and fell back less than a foot from where I was sitting. I glanced over my shoulder. The snake had raised its head. Mambas are reputed to be able to run down a galloping horse then kill the horse and the rider, and a green mamba had killed a giraffe on the reserve the year before I arrived. I don't know if I outpaced a horse or not, but I picked up my spotting scope and ran like hell to the boat. I have no idea if the snake tried to follow or just stopped and snake-laughed at the poor fool it had just scared.

Despite their limitations, the point estimates from ground surveys in 1969-1971 compared to the 1973 strip-census results indicated a decline between 1971 and 1973 of 10-15%. I believe the decline was real and was likely a function of harvesting animals for my study. The Loskop population was evidently stable in 1969-1971 but had little resilience to added mortality.

Before I arrived in South Africa, managers at Loskop attempted to survey ungulate populations using game drives. Ten or more employees would form a line and move across the landscape noting species and numbers of animals they flushed. Observers were stationed ahead of the line to count animals that flushed before drivers reached them. Then, as each area covered in the drive was finished, observers and drivers attempted to reconcile counts. Drives were held every 1-3 years and covered different sets of sub-units (generally the area covered included only those fenced and stocked with introduced species). The first report of a census in Loskop Dam Nature Reserve was for 1960 (Riney and Kettlitz 1964) when 150 mountain reedbuck were counted in the north-central units, the only fenced areas in the reserve at the time. Counts from drives in the same units from 1961 when compared to belt-transects I walked in 1969-1973 indicated an increase in 1961-65, a decline between 1965 and 1969 (possibly due to drought), an increase in 1969-70, and a decline by 1973. Counts from monthly reports for Sections 2-6 obtained from drive counts (1968, 1969, 1970) roughly agreed with estimates from my strip censuses in 1969-1971.

How Loskop compared to other populations was difficult to determine. Table 3 gives best population guesses for nine populations during the period I was in the field. The estimates (with the exception of Loskop) are based on short surveys or estimates from biologists and managers familiar with the areas. I have converted these estimates to densities for comparison. I suspect that the confidence intervals on estimates would likely be as wide as those for Loskop if they were available. Visits to these supplemental populations did provide me with some of my best tales of adventure in Africa. I had a few encounters with lions and other large beasts that made me uneasy, but these run-ins have given me material for bar conversations for more than forty years. The worst experience by far was stomping through the van Rooyen Ranch during the tick season. I had no facilities for washing them off so I suffered through tick bites for weeks after I left the ranch.

Table 3. Habitat available or surveyed, estimated mountain reedbuck density, estimated total population in survey area, and sources of estimates for 9 populations, 1969-1973.

I did have more run-ins with more conventionally "dangerous" animals while surveying mountain reedbuck populations. Two encounters with lions in supplemental populations stand out in my mind. The first encounter of a leonine kind occurred in Kruger National Park as I was walking through the Lebombo Mountains, the best mountain reedbuck habitat in the park, trying to get an idea of mountain reedbuck density. The park staff had assigned a game guard, a Tsonga named Isaac, to accompany me. Isaac didn't speak English or Afrikaans, which I was reasonably fluent in by this time. Given the density of lions in the area, the Lee-Enfield rifle he carried was comforting. After a full day of walking and sweating (it was the winter season, but I am sure the temperature was in the mid-nineties), we were less than a mile from my Land Rover when lion heads started popping out of the grass ten yards in front of us. You would think a bunch of 200 to 400-pound cats would be pretty obvious at that distance. Think again. Figure 20 is a photo of six lions taken at that range in Botswana (from a Land Rover). I think I counted six in the pride we surprised in Kruger, but I was more interested in what they would do than in accurate counting. I stood with one foot in the air, my only defensive weapon a 40-power spotting scope, for several seconds. Then I looked over my shoulder. Isaac was slowly retreating down the hill. I stood one-footed for a couple of more seconds and decided it might be a good idea to emulate him. Before I could turn, lions broke out of the grass going in all directions. One passed within ten feet of me. When I put my foot down, the lions were long gone, and Isaac was smiling at me with a sheepish expression on his face. I mentally reassessed my disappointment in having a primary study area that did not have a lion population.

Figure 24.(new Fig 20) Lions in tall grass in the Okavango Delta, Botswana, 1970.

A year later, I was sitting on top of my Land Rover in the Ngorongoro Crater using a spotting scope to scan for mountain reedbuck when a pride of lions broke out of tall reeds a few yards away. They were after a Bohor reedbuck grazing twenty or thirty yards from my vehicle. The reedbuck escaped, and the lions ignored me. I gave the incident considerable thought the next day, however, as I walked out of Ngorongoro. Shortly after seeing the lions, I managed to stick my Land Rover in deep mud while crossing a donga on the crater floor. I dug all afternoon, slept in the vehicle that night, and dug all the next morning without freeing my "go-anywhere" four-wheel drive vehicle. In one of the most famous tourist sites in East Africa, I did not see another car, truck, or tourist van either day. I finally decided to walk the ten miles to the reserve headquarters on the crater rim. No tourist cars appeared during the whole walk. On the other hand, I didn't run into any lions, either.

I encountered lions, elephants, rhinos, buffalo, and other potentially dangerous species a number of other times, but these encounters were not in the line of work. Two such incidents stand out in my mind after all these years. In 1970, I was "safari-ing" with friends in the Okavango Delta of Botswana. We drove in, set up camp on Chief's Island and prepared to go to sleep. There were lions roaring somewhere off to our south. One of my friends had a tape recorder and decided to record the roars. Another friend had the bright idea of playing the recorded roars back on loudspeakers. The lions were not pleased with intruders. We heard the roaring come closer and closer and wound up spending the night in our vehicles. The next night, we refrained from playing back recorded lions. We heard no distant roars so we ate, engaged in philosophical conversations, and finally retreated to our tents for a night's sleep. At some time during the night, someone woke up and saw a pride of lions walking through our camp. The other campers watched nervously as the lions wandered around for a half-hour. Nobody woke me up so I missed the whole episode. The party included several folks who had distinguished careers as biologists in South Africa so I won't embarrass them by mentioning names, but they watched as lions went up to my tent, sniffed around, and eventually left. It seemed to me that my friends had used me as bait to observe lion behavior. Or maybe they just didn't want to yell to draw attention of the lions to themselves. Anyhow, the lions left, and I slept peacefully through the night. I sometimes wonder what would have happened today. This was in the pre-I-phone pre-Facebook era. Nobody had a handy video device so there is no physical evidence of the episode.

The second incident occurred as we were driving back from the Okavango along the Chobe River that formed the border between Botswana and Zambia. We were all looking forward to visiting the Vic Falls Hotel and, after two weeks of warm beer, getting a cold one at Victoria Falls. We saw a bunch of elephants grazing along the Chobe so I stopped on the road, set the emergency brake, and prepared to take pictures. There were a dozen cows, calves, and sub-adults in the little herd (Fig. 21). They seemed amenable to letting us sit and take pictures. I tried to focus the long lens I was using but couldn't. When I looked up, an adolescent male was charging! I popped the clutch and immediately killed the engine (I forgot about the emergency brake). The elephant veered and passed by the Land Rover. I can swear I heard the little bastard chuckling as he thundered by.

Figure 21. Elephants feeding along the Chobe River innorthern Botswana, 1970.

One of the best things about a generalized study with few defined objectives is that it allows you to wander. I was lucky to be doing field work in southern and eastern Africa during a quiet political time. South Africa had yet to shed apartheid. Rhodesia (now Zimbabwe) and Mozambique were still under colonial control with no big wars going on. Botswana had achieved independence 10 years before I arrived and was blessed with a stable government. Kenya and Tanzania were independent, but most folks in both countries were unsure exactly how to treat lone white men (Were they oppressors or bwanas?) and were beginning to realize that white tourists could be a cash cow.

# Chapter 11 - Limiting Factors

In my dissertation, I considered a variety of factors that could limit mountain reedbuck population numbers in the Loskop Dam Nature Reserve. Large predators, with the exception of leopards, could not have been a factor in Loskop because lions, cheetahs, spotted hyenas, hunting dogs and all but a few leopards had been eliminated from the area in the 19th century. Given their hunting strategies (see Dorst and Dandelot 1970, Schaller 1972, Kruuk 1972, or any number of other fine books completed on African carnivores in the past half-century), I doubted that lions, spotted hyenas, cheetahs, and hunting dogs would have done more than take mountain reedbuck incidentally. Lions prefer animals larger than mountain reedbuck. Cheetahs, wild dogs, and spotted hyenas generally hunt in flat terrain. Medium-sized predators did prey on mountain reedbuck, and their numbers had increased since the formation of the Loskop reserve. Prior to formation of the reserve, ranchers made efforts to eliminate all mammalian predators so medium-sized predators were likely rare prior to 1948. Leopards, caracals, and black-backed jackals were capable of killing adult mountain reedbuck. Brown hyenas and baboons could take lambs. It seems unlikely that these species had much influence on mountain reedbuck during my time at Loskop (1969-1971, 1973). Mountain reedbuck in the Loskop Dam Nature Reserve both increased and decreased during the period for which I had population estimates (1961 – 1973) while medium-sized predator numbers were thought to have increased steadily over the period. The low lamb to ewe ratios at Loskop could have indicated the predation effect, if significant, was likely due to predation on juveniles.

Disease and parasites, unless they increased mortality in very young lambs, were not players at Loskop when I was in the field. The terrain and vegetation communities in the Loskop reserve appeared to be well within the range of conditions at supplemental study sites. Based on what I observed in the Orhigstad Dam Nature Reserve and the Cole Ranch (Table 8), Loskop could have supported a denser population.

Dry season forage quality, or lack thereof, in the grass sward was a prime suspect in population regulation. The condition of mountain reedbuck collected during the late dry season was noticeably lower than during the early dry season (Table 1). External and some internal parasite loads followed a similar pattern. Soils in Loskop were generally shallow, rocky, and low to moderate in nutritive status (Theron 1973). Protein levels for common grass species in the late dry season were all below the 7% level recommended as a minimum for livestock. Mountain reedbuck could compensate for some nutritional deficiencies by selectively feeding on the "best of the bad" species and selecting the most nutritious plant parts of all species (newer leaves), but selectivity can only accomplish so much when all of the plants that you can reach are desiccated. Mountain reedbuck were very reluctant to switch from graminoids to dicots even when grasses were poor in quality. There was no indication that amount of graminoids was limiting.

Nutritional stress is a common limiting factor in ruminants (see Meredith, D. 1955 and Chapman and Feldhamer 1982). In Africa, this has been shown to result in delayed puberty, increased calving intervals, and decreased calf survival in several antelope species (Skinner and van Zyl 1969, Bigalke 1970, Liversidge 1973, Penzhorn 1973). Observations of higher lamb to ewe ratios and a younger age structure in animals collected from the Cole Ranch, where grass protein levels stayed above 7% throughout my study (Blankenship and Qvortrup 1974) support the idea of nutritional stress as a limiting factor at Loskop. While surface water was suspected to be a major factor limiting distribution of mountain reedbuck (Dorst and Dandelot 1970), it was unlikely an issue in Loskop where no areas were more than two miles (3.2 km) from dry season water sources.

I concluded that the Loskop population was likely regulated by dry season forage quality and could not effectively respond to moderate harvest pressure because lower densities would not result in added resources to surviving animals. If maximizing mountain reedbuck density was a primary goal for the reserve, predator control might help. A better strategy would involve supplemental feeding with high quality forage, fertilizing and irrigating native grasses, or planting and irrigating cover crops such as oats. Developing supplemental water sources (dams, wells) might increase density in the areas most distant from dry season surface water. The management of Loskop Dam Nature Reserve, however, was geared towards maintaining healthy populations of a diverse set of indigenous species for tourism so I maintained that intensive management for mountain reedbuck was unnecessary.

For the species as a whole, I concluded that many areas with grass, broken terrain, and year-round surface water should support mountain reedbuck. Mountain reedbuck probably didn't reach many suitable areas simply because they were sedentary, and unoccupied areas were distant from occupied areas. Established populations seemed to be at only low to moderate risk from impacts by indigenous predators and human land use. Because many populations were restricted to isolated areas of broken terrain, these populations would be at risk from localized events (drought, pandemics, mine development, over-grazing by livestock. excessive predation, etc.), and recolonization of extirpated populations could be difficult given the sedentary nature of the beasts. When I finished my dissertation, I believed that the future of the species as a whole was as secure as that of any antelope species in Africa. It turned out I was wrong.

# Chapter 12 - New Information 1976-2019

Over the last 50 years, relatively few studies of mountain reedbuck were completed, and most of them were done on the southern sub-species. I had hoped that modern genetics techniques would have resolved my questions about sub-speciation, but I was unable to find papers that addressed any taxonomic level lower than the species (Birungi and P. Arctander 2001, Rubes et al. 2008, Heywood 2010, Pagacova et al. 2011, Cabelova et al. 2012).

Recent studies (or at least more recent than mine) on the biology of the species generally confirmed my findings. Body weights of adults were within the range I found at Loskop and the Cole Ranch (Skinner 1980, Anderson and Koen 1983, Taylor et al. 2005). Norton and Fairall (1991) were able to determine ages of adults using dental cemental lines, which I had been unable to do. Their paper did indicate that my estimates of the timing of tooth eruptions in younger animals were reasonably accurate. Based on irruption of molars and molar annuli, Norton and Fairall (1991) showed that males and females reached full adult body weight at 2 to 3 years of age. Male horn growth leveled off during the same period. I had estimated full adult size at 2 years of age. Reproductive studies confirmed that mountain reedbuck females reached reproductive maturity at or slightly less than the age of one year and produced their first lambs as yearlings (Skinner 1980, Els 1991, Norton and Fairall 1991, Anderson and Koen 1993).

Males reached adult body mass range at two years of age. They achieved adult horn shape by the age of three but maximum length and mass at around five years of age (Norton and Fairall 1991). All of the studies I found indicated some females were bred as early as one year of age, produced single lambs, and produced a lamb a year with a "relaxed" seasonal pattern (i.e. they tended to give birth during the period with highest availability of green grass but could produce lambs outside this period). Norton and Fairall (1991) believed that females were reproductively active at least through their tenth year of life.

Matschei (2004) reported the reproductive histories of 73 mountain reedbuck born in a Berlin zoo (derived from eight mountain reedbuck captured in South Africa) from 1984–2004. Females produced their first lambs at 11–25 months. Males reached reproductive maturity at 17–25 months. Gestation periods were recorded as 240-253 days with intervals between lambs of 11-12 months. Males were actively breeding at up to 11 years of age, and some females produced their last lambs at the age of 14. My observations of age of reproductive maturity (~1year for females and 1-2 years for males) and gestation (236-251 days) were evidently reasonable, especially for the limited observations I based them on.

I observed some breeding seasonality in South Africa and Kenya. Peaks in lamb births seemed to correspond to wet seasons. Mason (1977) observed young lambs during November – April, the rainy and early dry season in his Transvaal study area. Oliver et al. (1978) observed young lambs in eight months of the year suggesting a prolonged lambing season with a rainy season peak in their study area in the Drakensberg Mountains. Skinner (1980) found the highest concentration of births in the Mountain Zebra National Park in the Cape Province in October – December (the rainy season there) with scattered births in other months. Skinner et al. (2002) found similar rainy season peaks in mountain reedbuck born in the Pretoria zoo.

My descriptions of social organization and behavior were confirmed by later authors (Mason 1977, Oliver et al. 1978, Wirtz and Lorscher 1983, Roberts and Dunbar 1991, Dunbar and Roberts 1992, Taylor and Skinner 2006, Taylor et al. 2006). Mountain reedbuck males seem to be territorial year-round in all populations studied. No authors mentioned territorial behavior in females. Reported group sizes were similar to those I saw (average of ~3 with a range of 1-10), but a few authors mentioned anecdotal reports of larger groups (which I still think are the result of driving adjacent groups together when they were disturbed by humans).

Habitat selection reported in other studies (Mason 1977, Oliver et al. 1978, Skinner 1980, Rowe-Rowe 1983, Beardall et al. 1984, Pienaar et al. 1993, Newmark 1996, Taylor et al. 2007, Taylor et al. 2016a, Kahana et al. 2013) appeared to be similar to what I saw. Mountain reedbuck occupy a specific niche which allows them to "fit in" among a large suite of other herbivores. They select rough terrain, usually with shrub or tree cover of some sort, but open enough to support a dense ground cover of grasses. This allows them to make use of slopes to escape predators and to feed on the most abundant vegetation available at ground level, graminoids. The problem with a grass diet is that graminoids tend to be low in quality (most frequently low in protein and high in difficult to digest cellulose) in dry or cold conditions. Mountain reedbuck are among the smallest African antelopes to subsist on a diet dominated by graminoids. Their rumen structure is similar to that of larger species in the genus Redunca, and they have morphological adaptations in their digestive tract (Hofmann and Stewart 1972, Pérez-Barbería et al. 2001) typical of grazers. Most ruminant species that rely on a grass diet are either very large (so they can support a larger fermentation chamber) or small enough to select grass parts that are above average in digestibility (new leaves, succulent seed heads, etc.). Mountain reedbuck seem to use both strategies. Their muzzles are small enough to be somewhat selective in what they bite, and they have a comparatively large rumen.

The topographic placement of suitable habitat seems less important than the presence of rough terrain and a grass sward. They occur in the foothills and lower slopes of high mountains, at all elevations in lower mountain ranges, along escarpments between shrubland and plateau grasslands, and in rugged lava flows below volcanos. In the Drakensberg Mountains of South Africa and the massive volcanos of East Africa, mountain reedbuck are reported to occupy the zone between montane forests and moorland (Rowe-Rowe 1983, Newmark 1996, Kahana et al. 2013).

Studies completed after I finished my fieldwork (Mason 1977, Oliver et al. 1978, Skinner 1980, Rowe-Rowe 1983, Pienaar et al. 1993, Newmark 1996, Taylor et al. 2007, Taylor et al. 2016a) note that mountain reedbuck seem to be found only in areas where surface water is available during the dry season. This dependence on surface water is likely inherited from ancestral reedbucks and limits mountain reedbuck distribution within environments that have plenty of rugged terrain but limited availability of dry-season surface water. Territorial behavior in males and restricted home ranges in females ensure that they know the structure of the area they live in well, thus allowing them to escape predators. This high fidelity to specific areas, however, makes them poor colonizers. When a population is extirpated, it may be difficult for animals from neighboring populations to locate suitable empty habitat.

As with most African antelope, mountain reedbuck have adapted to co-exist with other antelope species. The greatest geographic overlap between mountain reedbuck and other species would be with grey rhebok, oribi, and other Redunca species (Oliver et al. 1978, Rowe-Rowe 1983). The ranges of southern reedbuck ( _R. arundinum_ ) and southern mountain reedbuck ( _R. fulvorufula fulvorufula_ ) and those of Bohor reedbuck ( _R. redunca_ ) and Chanler's mountain reedbuck ( _R. fulvorufula chanleri_ ) do overlap in some areas, but the larger reedbuck species prefer tall grass areas in flood plains or other mesic environments and seldom move into complex terrain with shorter grasses (Jungius 1971, Kingdon 1982). Oribi feed primarily on grass but tend to occupy open grasslands and plateaus as well as savannah. They are not limited to areas with rugged terrain (Kingdon 1982). They are smaller than mountain reedbuck and presumably able to feed more selectively than mountain reedbuck. They are dependent on surface water during dry seasons. Grey rheboks prefer areas with low vegetation with or without steep slopes (Taylor et al. 2016b). Their diet separates them from mountain reedbuck. Grey rhebok feed primarily on herbaceous dicots (forbs) and low woody shrubs and are largely independent of surface water. They use steep slopes but are not limited to them.

Until recently, southern mountain reedbuck and Chanler's mountain reedbuck were regarded as relatively secure compared to many other antelope species. They were found over large geographic areas, occupied marginal lands unlikely to be attractive to settlers, were found on both government and private lands, did not possess large horns that would be attractive to trophy hunters, were inconspicuous and relatively small, and lived in small groups that would not attract the ire of farmers. The status of Adamawa (also known as Adamaoua or western) mountain reedbuck was of much greater concern, primarily because of their limited range. During the late 20th century and early 21st century, southern mountain reedbuck were being promoted as a game animal that could provide high quality food to humans, challenging prey for hunters, and supplementary income to farmers in South Africa (Skinner 1980, Taylor et al. 2005, von Brandeis and Reilly 2007, von Brandeis and Reilly 2008, Cousins et al. 2010).

Studies identifying possible limits to exploitation began to appear as early as the 1970s (Irby 1976, O'Connor and Kruger 2003), but serious widespread declines were not expected. These scattered cautionary reports were outnumbered by advertisements for mountain reedbuck hunts as ranchers began to increase emphasis on game ranching. As I was writing this book, I did a Google search on " _Redunca fulvorufula_." I got 20,200 hits. When I searched for "hunting mountain reedbuck," I got 56,900 hits. Most of the " _Redunca fulvorufula_ " sites were photos or natural history blurbs, but 7-8% were hunting sites. I can't fathom Google algorithms, but it looks like interest is high in hunting mountain reedbuck even as a review of population trends in over 400 populations in southern Africa, including protected areas (40) and herds on private lands (395) (Taylor et al. 2016a) indicated a decline from 33,000 individuals in 1999 to 15,000 in 2013. The population estimates in most of the surveyed populations were based more on educated guesses than formal censuses, but more than half the few protected areas with routine census procedures reported substantial declines.

Poaching (snares, guns, and dogs) and encroachment by humans on habitat were the most likely causes of the decline (Taylor et al. 2016a), but some researchers have reported population declines driven by native predators. Increases in jackal, leopard, and caracal populations are common where ranchers reduce predator control efforts as a result of switching from sheep to cattle or to changing from livestock operations to hunting and game viewing. I was surprised to find a report of re-introduced cheetah and Cape hunting dogs implicated in a decline in Chanler's mountain reedbuck in Kenya (de Jong 2015, http//lolldaiga.com/mountain-reedbuck). I believed those species tended to hunt in habitat much different than that occupied by mountain reedbuck. Local populations may also be threatened by drought and unusually cold weather (Taylor et al. 2005). As climate changes, surface water availability in dry seasons is likely to decrease and productivity in the grass sward may decline as rainfall patterns change. Even if the two most likely threats, human population growth and poaching, are controlled, recovery of mountain reedbuck would likely be slow. As territorial animals, they are disinclined to travel far to find empty habitat.

# Chapter 13 – Status of Study Areas 2019

I attempted to determine the status of the nine populations with which I worked without much success. The internet is a font of information, but it can never replace fieldwork and conversations with biologists familiar with local conditions. I did uncover information on five of the six South African populations in which I collected data and general information on the three populations I observed in East Africa. All reported extant populations of mountain reedbuck, even if they did not give details on numbers or population trends. I was surprised by some of the changes that had occurred since I last visited in 1973. I was not surprised that the agencies managing public herds in South Africa had been re-organized following the end of apartheid, but I was surprised by changes in the status and/or sizes of some of the areas in both South and East Africa where I observed the species.

The **Loskop Dam Nature Reserve** is now in Mpumalanga Province and managed by the Mpumalanga Tourism and Parks Agency. The dam was raised in the 1970s with the full pool area increasing from 1,700 ha to 2,300 ha to support expansion of irrigated agriculture below the dam. The reserve was expanded by nearly 11,000 ha (85%) to compensate for land loss due to the raised dam and to keep waterfront areas in public ownership. This increased the reserve (land and water) to 23,000 ha (57,000 acres). Habitat lost along the river valley was replaced with land purchased on uplands. This likely benefited mountain reedbuck while harming species limited to riparian habitats. The website for the reserve (http://www. mpumalanga.com /our-provincial-parks/loskop-dam-nature-reserve) lists mountain reedbuck as "common." Poaching is an issue in the reserve with two poachers killed by police in a shootout in 2018 (https://www. news24.com/SouthAfrica/News/two-suspected-poachers-killed-in-loskop-nature-reserve-shootout-20180716).

**Orhigstad Dam Nature Reserve** , the southern African area I spent the second greatest amount of time in, is now in the Mpumalanga Province and managed by the same organization as Loskop. It has not changed in size since 1973 (2,600 ha – 6400 acres - around an 85-ha – 210 acre - irrigation reservoir), and the antelope species present in 1973 are still reported to occupy the reserve today. The reserve website (http://www.mpumalanga.com/our-provincial-parks/ohrigstad-dam-nature-reserve) gives no information on numbers or population trends in mountain reedbuck but mentions that tourists can expect to see mountain reedbuck.

I spent only a few weeks in **Giant's Castle Game Reserve** (Fig. 22). The game reserve is now a part of the Ukhahlamba Drakensberg Park, which is a World Heritage Site. The park is managed by the Ezemvelo KZN Wildlife, a division of the KwaZulu-Natal Provincial government, covers 2,400 km2 (940 mi2), and is managed in co-ordination with the adjoining Sehlabathebe National Park in the Kingdom of Lesotho. Ukhahlamba Drakensberg Park includes the 34,638-ha Giant's Castle Game Reserve as well as several other parks and reserves. The Drakensberg Mountains are the highest mountain range in Southern Africa. According to the official website (http://www.kznwildlife.com/form.php): "Game populations in the Ukhahlamba Drakensberg Park are kept at a level that can be sustained by winter grazing because with neighboring farms and fences, the wildlife is no longer able to migrate into lower more plentiful lands. As part of the grassland management program, large areas of the park are burnt annually to improve and maintain quality grasslands. Eland antelope and chacma baboons are commonly seen. Mountain Reedbuck and the rare caracal are occasionally spotted." This implies that mountain reedbuck numbers may have declined since the 1970s.

Figure 22. Mountain grassland and savannah in Giant' Castle Game Reserve, Natal, 1970.

I spent 1-2 weeks in 1970-71 surveying mountain reedbuck in the **Hluhluwe-Umfolozi Game Reserve** (now Hluhluwe-Imfolozi Provincial Park) located in what is now the KwaZulu-Natal Province. Mountain reedbuck at that time occupied only the hillier areas of the reserve, and their numbers were low. The park is now managed by the Ezemvelo KZN Wildlife Authority and encompasses 237,000 acres (96,000 ha). From park information on the web, it seems that the mountain reedbuck population is still spatially restricted and persists at low densities.

The **van Rooyen Ranch** , owned by a faculty member at the University of Pretoria in 1970, was located in the Natal Province and would be in KwaZulu-Natal today. I was unable to locate any information on the current status of the population on this property. Taylor et al. (2013) reported mountain reedbuck status on seven private properties (ownership not given) in KwaZulu-Natal that indicated relative stability. Van Rooyen is a common name in South Africa, but I did find a newspaper report of a Billy van Rooyen killed by robbers on April 20, 2016 (Vryheid Herald 20 Apr 16 - https://vryheidherald.co.za/ 41480/breaking-news-two-killed-in-farm-attack/) on his farm by robbers in the Vryheid District of KwaZulu-Natal. His father's first name was Lood (the same initial as the faculty member). I have no idea if this L. van Rooyen has any association with the ranch where I spent a week surveying mountain reedbuck in 1970.

**Kruger National Park** is one of the largest protected areas in Africa with an area of 7,500 mi2 (19,500 km2) in the provinces of Limpopo and Mpumalanga in northeastern South Africa. When I was in South Africa, Kruger was managed by the South African Parks Board. Today, it is still managed as a national park. Under the new constitution, it falls under SANPark (South African National Parks), a division of the Department of Environmental Affairs. Kruger also forms part of the Great Limpopo Transfrontier Park which includes protected areas in Mozambique and Zimbabwe. Most mountain reedbuck are found in the Lebombo Mountains in the southern portion of the park. I spent two weeks surveying mountain reedbuck in this area in 1970-71. At the time, densities were low. A report in 2009 (https://www.sanparks. org/parks/ kruger/conservation/scientific/ff/ biodiversity_statistics.php) includes an estimate of 150 mountain reedbuck for the entire park. My guess is this is a ballpark guestimation. Mountain reedbuck are notoriously difficult to census, and the park has no systematic program for counting this species. The estimate does indicate that the population may be stable.

The **Cole Ranch** was located in the Rift Valley 60 miles south of the equator in Kenya. With elevations ranging from 6,000 – 7,600 feet (1,800 – 2,300 m), the climate was sub-tropical. Approximately 20,000 acres (8,000 ha) of the 45,000-acre (18,200 ha) ranch was suitable mountain reedbuck habitat in 1971-72 (Fig. 23). Arthur Cole, the owner, sold the ranch to a community cooperative in 1977. It was divided into small farms soon after. Roberts and Dunbar (1991) completed a behavioral study of Chanler's mountain reedbuck on ranchland near the former Cole Ranch in 1991, but I would be surprised if any native antelope survive on areas converted to farmland.

Figure 32. (new Fig 23)Horseshoe Crater in the Cole Ranch, 1971. The savannah/woodland on the walls of this crater supported the densest mountain reedbuck population I encountered during my study.

The **Akira Ranch** is still in operation, but the portion of the ranch that supported mountain reedbuck, the slopes of the volcano Longonot, is under development for geothermal power, with the first commercial generating facility commissioned in the early 1980s. This geothermal development eventually required displacement of ~3,000 Masai pastoralists (Schade and Faist 2017), and the case was still being argued in Kenyan courts as of 2019. Environmental impact statements written in the 1990s and early 2000s included surveys of antelope at geothermal sites. Mountain reedbuck were not mentioned, but they were rare compared to several other species in the 1970s and likely still a minor player on the ranch.

The **Ngorongoro Conservation Area** (Fig. 24), protected as a World Heritage Site, is located in the Crater Highlands area of Tanzania and is now administered by the Ngorongoro Conservation Area Authority (http://www.ngorongorocrater.org/index.html). In 2009, the Ngorongoro Wildlife Conservation Act placed new restrictions on human settlement and subsistence farming in the Crater, displacing Maasai pastoralists, most of whom had been relocated to Ngorongoro from their ancestral lands to the north when the British colonial government established Serengeti National Park in 1959. Mountain reedbuck are restricted to woodland and mountain savannah on the crater walls. I spent several days surveying ~ 5,000 acres (2,000 ha) on the dry southern and western interior walls in 1971 and sighted all of 28 individuals. At the time, biologists estimated that more than 20,000 ungulates used the crater floor and inner walls. EasyTravel, a comprehensive African travel site (https://www.easytravel.co.tz/ngorongoro-crater/, updated 2019), mentioned that 30,000 ungulates and predators use the crater today. Mountain reedbuck were noted as "rare" and seldom seen by tourists.

Figure 24. Interior slopes of Ngorongoro Crater, 1971. Mountain

reedbuck habitat was limited to dry woodland and savannah on

the crater walls.

# Chapter 14 - Future Prospects

It seems that the mountain reedbuck is an example of a specialized species whose specializations do not match changes in the world in which they live. Territoriality, gramivory, dependence on radiated scent to communicate with con-specifics, warning whistles to alert other members of a social group to danger, a bouncing escape gait, and preference for rugged topography near surface water worked well for hundreds of thousands of years, but the pace of change in Africa has turned these specializations into disadvantages.

Territoriality allows an animal to learn its home range intimately, but it discourages range expansion and colonization to re-establish populations that have been extirpated. Mountain reedbuck eventually could return to empty habitat, but it is a slow process that works at the margins of occupied range. When patches of suitable habitat are separated by long distances, discovery of new habitat or reoccupation of lost habitat could take centuries.

Dependence on grasses as a primary food source and the associated dentary, digestive, and behavioral specializations that allow mountain reedbuck to exploit the most abundant ground-level plant species in many environments worked well until the modern world changed the game. With over-grazing by domestic livestock and global climate change turning more and more landscapes into desert, dependence on grasses for food is no longer as advantageous strategy as it used to be.

Use of radiated odor to communicate is advantageous to identify con-specifics as individuals and as individuals of specific genders, ages, and reproductive status. It has been hypothesized that the Redunca line may have developed olfactory radiation because ancestral species lived in seasonally changing riverine environments with tall graminoids (reeds, rushes, etc.) (Kingdon 1982). Use of stationary territorial markers such as fecal piles or glandular secretions deposited on plants would be constantly lost as rivers ebbed and flowed. Mountain reedbuck do not change their territories seasonally, but odor radiation may be a carryover from an ancestral species. In extant mountain reedbuck, strong odors radiated from their bodies could alert predators to the exact location of mountain reedbuck. It is possible that using scent markers (glandular secretions, feces, etc.) in territories as many other territorial African antelope do, may send intra-specific information without identifying exact physical locations to predators. This is more of a hypothesis than an established fact, but it is worth investigating.

Vocal signals (whistles in mountain reedbuck) are used by many species to alert con-specifics to the presence of predators. The exact location of an animal that uses a shrill whistle to warn of danger may be difficult for a predator to determine. At least it works that way for many passerine bird species that use similar alarm calls to alert nearby birds of several species to the presence of raptors (Magrath et al. 2007, Hetrick and Sieving 2012). This strategy is less successful against human predators. Mountain reedbuck are inconspicuous to human hunters until they announce themselves with whistles. This species also uses whistles in territorial displays which may lead con-specifics to ignore whistles directed towards predators. I couldn't tell the difference between the warning whistles and territorial whistles. I could differentiate between whistles issued by common reedbuck and mountain reedbuck. Maybe mountain reedbuck can differentiate between territorial and alarm whistles in their species.

The "rocking-horse" gait and raised tail (with its conspicuous white underside) provide cues to alert con-specifics of the presence of predators and the direction which may be safest to flee. I couldn't see any difference between the use of these visual signals in territorial displays and in situations where the animals perceived danger. Maybe reedbuck can. As with whistles, visual signals do alert human hunters to the locations of mountain reedbuck. If the animals stood still, they might be overlooked.

Habitat specializations are probably the most detrimental hinderances to mountain reedbuck success in today's world. Suitable areas of rough terrain are reasonably abundant in Africa. Suitable terrain with open to moderate tree cover, abundant grass, and access to surface water in the dry season is much more restricted and geographically scattered. When I worked on mountain reedbuck, most of the areas they occupied were regarded as marginal for agriculture. Over-grazing by livestock was sometimes a problem, but many ranchers provided surface water for their stock and tried to control the medium-sized predators most likely to kill mountain reedbuck. As the human population in Africa has exploded, areas which were once "marginal" are attracting farmers. Herders are also trying to graze their livestock on poorer pastures. Either situation puts mountain reedbuck in greater contact with poor people who are struggling to feed themselves.

Territorial behavior may be an acceptable strategy to minimize predation by indigenous predators, but it doesn't work well against poor rural agriculturists. Even poor people have access to wire for snares. They use it to control their livestock and can easily steal it from fences built by more prosperous neighbors. Areas occupied by mountain reedbuck can be easily identified and the best sites to place snares (fencelines on approaches to surface water, for example) located. Most countries within mountain reedbuck range have strict gun control laws, but local residents have greater access to guns than in the colonial era thanks to the interminable wars on the continent. Poor rural people also have dogs. A pack of dogs chasing mountain reedbuck will eventually run the animals, who are attempting to remain within a fixed territory or home range, past areas where hunters with guns can pick them off. Or dogs may kill or constrain mountain reedbuck while hunters follow the dogs to reclaim their prey before the dogs can eat them. All countries that have mountain reedbuck have game laws that regulate or prohibit take. Few countries have hunting traditions that limit take to sustainable levels or funds to effectively enforce laws. As in the United States, hunting is a popular rural tradition in Africa. There are also millions of people that have limited access to protein so "bushmeat" is avidly sought.

Changes in climate exacerbate these problems. As surface water becomes scarcer and forage productivity declines, mountain reedbuck will have fewer and fewer places where they can survive. The poor pioneering ability inherent in mountain reedbuck will inhibit their ability to find what empty suitable habitat there is. Ironically, as the continent warms and dries, some high elevation areas, such as the Kenyan highlands, Mount Kilimanjaro, and the Drakensberg Mountains, may become suitable habitat for mountain reedbuck – if they can find it. Overall, there is likely to be more habitat loss than gain since high elevation sites are scarcer than rough terrain at lower elevations.

Grey rhebok and oribi, species that are frequently sympatric with mountain reedbuck, seem to be less vulnerable to conditions in the modern world than mountain reedbuck. Grey rhebok are forb and low-growing shrub specialists, plant forms likely to increase as lands become drier. They are less dependent on surface water than mountain reedbuck and are less strongly tied to complex topography. Oribi are specialized gramivores and may be seasonally dependent on surface water, but they are a smaller meat package than mountain reedbuck, live in smaller (thus harder to locate) groups, do not announce their presence with whistles, and do not require rough terrain.

Soulé (1991) proposed a scenario for extinction where populations decline and eventually become fragmented. Once fragmentation occurs, population remnants begin to "blink out." Each disappearance may be due to different proximal causes. Some populations may disappear due to habitat loss. Others fall prey to hunting or disease. Still others may be eliminated due to weather extremes. At some point, low population density and/or restrictions on gene flow may lead to in-breeding, loss of fitness, and finally extinction. Kingdon (1982) believed that mountain reedbuck had evolved into an evolutionary cul-de-sac in that they have restricted habitat preferences and depend on proximity to surface water. This, according to his theory, made them more susceptible to extinction than many other African antelope.

I have painted a bleak picture of the future for mountain reedbuck, but there are potential solutions to the problems they are having. The distribution of the species is fragmented, but fragmentation can be beneficial. I spent over 20 years studying bighorn sheep ( _Ovis canadensis_ ) in the United States. Bighorns are also restricted to rugged terrain. They are not territorial but, in many areas, have to move long distances between seasonal ranges, and they are poor pioneers. If they cannot see suitable habitat (steep slopes, open vegetation, escape terrain in the form of cliffs, etc.), they tend to expand slowly even in areas where forests provide the only barrier between suitable patches of mountain habitat. But they do eventually manage to colonize empty habitat.

The disinclination of bighorns to colonize new areas has been a problem, but disease and competition with domestic livestock are bigger problems. Bighorn sheep are notoriously susceptible to diseases carried by domestic sheep. Separation, either by exclusion of domestic sheep or by habitat fragmentation that keeps individual bighorn sheep populations from interacting, may help control the spread of disease. When a strong tradition of sustainable harvest is developed by local hunters and funding for law enforcement and wildlife management are present, extant populations can be maintained, and populations that go extinct can be replaced by translocation. Mountain reedbuck do not seem to be particularly susceptible to epizootics common in Africa, but populations lost to poaching or over-grazing by domestic livestock can be restored, given public will, adequate law enforcement, and application of modern wildlife and range management techniques.

Climate change is normal for the earth. Human-induced climate change is a threat to most species which will not be halted until humans make a major commitment to control their behavior. In the meantime, there are some mitigation actions that might aid mountain reedbuck. Bighorn populations in desert areas of North America have been maintained by providing surface water in the desert. This can be achieved by drilling wells or passive storage systems for rain that does fall. Increasing surface water will not benefit wildlife unless provisions are made to regulate use of new sources by pastoralists. Providing boreholes in some areas of Africa has only allowed traditional migratory grazing to be expanded to year-round grazing which is detrimental to wildlife and the land.

Mountain reedbuck do have a few things going for them. In East Africa, several populations of Chanler's mountain reedbuck live in well protected areas with strong support from international conservation organizations. In South Africa, southern mountain reedbuck in many national and provincial parks enjoy substantial protection from poaching and habitat destruction. This sub-species also occupies several thousand private ranches where ranch owners have monetary incentives to maintain them as a huntable species. The Adamawa mountain reedbuck in Cameroon is highly vulnerable to extirpation. It has a very limited range in a poor country with little money to spend on wildlife protection. The other two sub-species have a good chance of surviving for the next century.

# Chapter 15 – Calling It a Day

As you have noticed, this is a highly personal view of the current status of an obscure species. My perspective on the current status of mountain reedbuck is based on information I could find in the public record (i.e. the internet and the Montana State University library). I live in Montana and have neither the funds nor the physical stamina to do on-site investigations in Africa. After working on this project for a year, I can tell you that long-distance biological assessment is less than ideal. Obtaining recent agency reports was difficult. Busy biologists and administrators have little time and interest in responding to requests from someone half a world away who has not been involved in mountain reedbuck research in half a century. I wish I did have the resources to revisit all of my old study areas, but even on-site work where you can talk to other biologists and obtain obscure reports from local sources has limits. You may wind up studying atypical populations or typical populations under atypical conditions. You will always face limits on funding and feasible techniques. And no biologist can follow a population over an unlimited time span.

This assessment sounds pessimistic, but limits on studies are a fact of life in science. Every study is limited by the state of current knowledge. Science can only base the search for new information on what is already known. New hypotheses are formulated on what has been discovered by past researchers. Few researchers are able to pursue one set of questions throughout their careers, and careers seldom last more than a few decades. Questions regarded as important at one point in time may seem trivial at another time. Techniques considered "state of the art" are replaced by new approaches. How do you integrate old information with new information? Which population questions are worth centuries of monitoring? How do you allocate monitoring effort among individual populations? I wish I had answers.

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