UWCScholar :: Browsing by Author "Maritz, Bryan"

Browsing by Author "Maritz, Bryan"

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An ambusher’s arsenal: chemical crypsis in the puff adder (Bitis arietans)
(The Royal Society, 2015) Miller, Ashadee Kay; Maritz, Bryan; McKay, Shannon; Glaudas, Xavier; Alexander, Graham J.
Ambush foragers use a hunting strategy that places them at risk of predation by both visual and olfaction-oriented predators. Resulting selective pressures have driven the evolution of impressive visual crypsis in many ambushing species, and may have led to the development of chemical crypsis. However, unlike for visual crypsis, few studies have attempted to demonstrate chemical crypsis. Field observations of puff adders (Bitis arietans) going undetected by several scent-orientated predator and prey species led us to investigate chemical crypsis in this ambushing species. We trained dogs (Canis familiaris) and meerkats (Suricata suricatta) to test whether a canid and a herpestid predator could detect B. arietans using olfaction. We also tested for chemical crypsis in five species of active foraging snakes, predicted to be easily detectable. Dogs and meerkats unambiguously indicated active foraging species, but failed to correctly indicate puff adder, confirming that B. arietans employs chemical crypsis. This is the first demonstration of chemical crypsis anti-predatory behaviour, though the phenomenon may be widespread among ambushers, especially those that experience high mortality rates owing to predation. Our study provides additional evidence for the existence of an ongoing chemically mediated arms race between predator and prey species.
Autecology of the rhombic egg-eater, Dasypeltis scabra
(University of the Western Cape, 2022) Barends, Jody Michael; Maritz, Bryan
Efforts to prevent global losses of biodiversity are hindered by incomplete knowledge of the functional importance of species. In part, this is due to a paucity of information on the basic biology of most species resulting in a poor understanding of their ecological roles. Snakes represent a speciose group of carnivorous predators that are likely important ecological components of a variety of environments given their roles in modulating the sizes of the populations of their prey. Unfortunately, the ecological importance of most species of snakes has yet to be empirically demonstrated, particularly in native systems.
A complete guide to the snakes of Southern Africa
(African Journal of Herpetology, 2023) Maritz, Bryan
Background: The new edition of A Complete Guide to the Snakes of Southern Africa represents the third iteration in a series of books that started three decades ago. The latest edition replaces the 2004 edition (Marais Citation2004)—an excellent book in its own right (Bauer Citation2004). In the nearly two decades since the last edition, there have been several important advances in the study of snakes in southern Africa. These include changes in systematics and the description of new species, a substantial formalisation of conservation assessment processes, and the remarkable growth of citizen/community science platforms that have improved the quantity and quality of geographic distribution data and natural history information
Conservation status and threats for African reptiles
(Elsevier, 2016) Tolley, Krystal A.; Alexander, Graham J.; Branch, William R.; Bowles, Philip; Maritz, Bryan
The assimilation of information on taxonomy, distribution, basic ecology and conservation status of Africa's reptiles lags far behind that for most other continents. Many regions of mainland Africa are rarely surveyed, resulting in severe knowledge gaps that currently limit effective conservation of African reptiles. Here, we provide a précis on the knowledge gaps and conservation status of mainland African reptiles, and quantify the main threats based on IUCN Red List of Threatened Species assessments using publicly available distribution data. Our results show that these data are insufficient to confidently identify areas of high biodiversity, with large gaps in knowledge in the Horn of Africa, central Africa and West Africa. There is a strong overall taxonomic bias in extinction risk with 45% of families more threatened than expected by chance. Furthermore, Amphisbaenidae, Chameleonidae, Gerrhosauridae, Testudinidae, Viperidae all have a high percentage of their constituent species at risk. Overall, land transformation for agriculture, particularly subsistence farming, constitutes the primary threat to African reptiles, and our derived Threat Index based on socio-economic traits of African countries show that risk is high in Burundi, Ethiopia, Liberia, Malawi, Rwanda and Sierra Leone. These findings highlight important challenges facing the conservation of African reptiles, and we suggest that conservation priorities in mainland Africa be focussed on areas where the potential for overall loss of biodiversity is high, particularly in regions where knowledge is inadequate.
The diet and feeding ecology of the brown house snake, Boaedon capensis
(University of the Western Cape, 2022) Bassier, Ielhaam; Maritz, Bryan
African brown house snakes (Boaedon capensis) are widely thought to be dietary specialists that predominantly consume rodents. Given their ubiquitous distribution, these snakes potentially play an important role in controlling rodent populations throughout their range. However, the full extent of the diet of this species remains poorly quantified, and the proportional importance of mammals to their diet is speculative. Moreover, little is known regarding intraspecific dietary variation of these snakes. In recent years, a dearth of reports of B. capensis feeding, particularly from novel information-sharing streams available through social media, suggests that the diet of these snakes may be broader than previously thought. B. capensis are not rodent specialists as amphibians (3%), birds (12%), and reptiles (38%) collectively comprise a significant proportion of their diet. There was no evidence for sexual size dimorphism or dietary variation between adult males and females. However, significant differences in morphology and prey utilisation between adults and juveniles indicate an ontogenetic shift in diet from small lizards to larger, and more diverse prey. Variation in climate and time of year did not affect diet. Importantly, the diet of B. capensis as presented by museum data vastly differed compared to literature and community science reports. In particular, the proportional use of reptiles differed across sources (museum data = 6%; community science data = 27%). Overall, my findings reiterate the importance of examining multiple sources of information when characterizing the diet of species, as these multiple sources provide different result outcomes.
Diet and feeding in the Cape Cobra, Naja nivea
(Taylor & Francis, 2017) Layloo, Inshaaf; Smith, Caitlin; Maritz, Bryan
It is widely reported that African cobras (genus Naja) are generalist predators, but examination of formal literature yields relatively little evidence of this for most species. Here, we review the diet of cape cobras (Naja nivea) based on examination of museum specimens, an extensive literature review, and social media reported natural history observations. We quantify dietary niche breadth and test the hypothesis that cape cobras are generalist predators. Additionally, we test the hypotheses that diet varies significantly (1) across biomes, and (2) across winter, summer and year-round rainfall regions. We gathered 101 feeding records for cape cobras and based on family-level prey frequencies calculated Levins’ measure of niche breadth as B = 6.57, which we standardised to BA = 0.29. We found no association between prey classes and biomes or rainfall region. Our data indicate that cape cobras consume a wide range of vertebrate prey from all four tetrapod classes, including a large number and diversity of snakes. Our findings suggest that cape cobras may be involved in several important ecological processes. Moreover, our study demonstrates the utility and value of collating natural history observations reported on social media platforms.
Foraging ecology of Naja nivea and Dispholidus typus
(University of the Western Cape, 2019) Greuel, Janine; Maritz, Bryan
It is widely reported that snakes can be major predators of avian nests, but the use of a single avian prey type by competing species has rarely been examined. This study aimed to investigate predation of a single food resource by the sympatric snakes Naja nivea and Dispholidus typus. Specifically, I aimed to 1) identify factors influencing snake presence in sociable weaver colonies and 2) quantify snake predation and potential differences in the consumption of prey by the two competing snakes. I used repeated visual surveys of sociable weavers to obtain presence-absence data of cape cobra and boomslang in sociable weaver colonies over an entire breeding season. I related the presence-absence data of the two snake species to spatially- and temporally-variable factors using principal component analyses (PCA) and multiple logistic regression analyses. The presence of snakes in sociable weaver colonies is primarily influenced by temporal factors, but spatial factors also play a role.
Functional importance of snakes in a strandveld ecosystem
(University of the Western Cape, 2018) Forgus, Juan-Jacques; Maritz, Bryan
Gaps in our knowledge of the functional roles of snakes within ecosystems limit our ability to predict the potentially cascading effects their removal from an ecosystem might create. Extirpation of snake species could potentially result in losses of ecosystem functionality if those taxa are ecologically unique. I used pitfall and funnel trap arrays, artificial cover object surveys, active searching, and passive camera trapping, as well as pre-existing faunal diversity data to identify terrestrial tetrapod species within the Koeberg Private Nature Reserve. This resulted in a list of 265 species, of which 13 were snakes. I then gathered data on dietary and four additional functional traits for each species from the literature. Next, using hierarchical and partitioning around medoids clustering, I identified ten broad dietary guilds and 54 functional guilds within the terrestrial tetrapod community. Of the dietary guilds Dasypeltis scabra was the only snake species that formed a unique single species guild and was one of four snake species (Pseudaspis cana, Homoroselaps lacteus and Lamprophis guttatus) to form four unique single species functional guilds. The remaining snakes clustered together within groups of other vertebrate predators. Functional diversity analysis was then used to simulate losing eight major taxonomic groups (birds, passerines, non-passerines, mammals, reptiles, snakes, non-snake reptiles and amphibians) and gauge the effects of those losses on overall community dietary and functional diversity. Functional diversity analysis revealed that the loss of certain snake species resulted in disproportionate losses of overall community dietary and functional diversity while losing others had negligible effects. These findings provide ambivalent support for the dietary and functional uniqueness of snakes suggesting that certain snake species are fulfilling unique functional roles within the ecosystem. Additionally, it is likely that losing those non-redundant species would result in significant losses of ecosystem functionality.
Identifying global priorities for the conservation of vipers
(Elsevier, 2016) Maritz, Bryan; Penner, Johannes; Martins, Marcio
Vipers are among the most misunderstood and persecuted animals. They occupy most terrestrial ecosystems around the world, often at high population densities. However, certain aspects of their biology (e.g., low fecundity and slow growth) have resulted in vipers being disproportionately threatened by extinction. Despite increased extinction risk, relatively little is known about viper biology, severely limiting the development and implementation of conservation initiatives. Here, we review the conservation status of vipers globally, map species richness, and develop three indices (one reactive; one proactive; one combined index emphasising irreplaceable species facing severe threats) to identify species for which conservation action should be prioritised. Moreover, we map species richness weighted by each index to identify regions for conservation prioritisation. We ranked prioritisation scores for all species for which data were available. In doing so we identify species for which valuable data are missing and that should be prioritised for research. We additionally show that 17 species, currently listed as Not Assessed or Data Deficient by the IUCN, score sufficiently high on our Threat Index to be considered as Threatened in the future. We identify multiple regions for which viper conservation action should be prioritised. These areas broadly correlate with species richness patterns, suggesting that species richness may be an effective proxy for conservation planning. Finally, we discuss the major gaps in knowledge, as well as the major threats facing vipers.
Phylogenetic assessment and historical biogeography of the Psammophis leightoni complex
(University of the Western Cape) Taft, Jody M.; Maritz, Bryan; Tolley, Krystal A.
Stable and accurate taxonomy remains a primary component of conservation. By overlooking taxonomic disorder, conservation management strategies may ineffectively distribute resources. The Psammophis leightoni species complex is one such example where taxonomic confusion may have an influence on the conservation of threatened species. Psammophis leightoni, P. namibensis and P. trinasalis were all elevated to specific rank on the basis of ecological differences largely attributed to where these taxa occur. A molecular revision of Psammophiinae highlighted that P. leightoni and P. namibensis show levels of intraspecific divergence; however, this was based on single representatives per putative species. Psammophis leightoni is currently considered Threatened and is listed as Vulnerable [B1ab(iii)], but the taxonomic uncertainty surrounding the P. leightoni complex influences how these taxa should be regarded in a conservation context. To remedy this, I aim to validate the taxonomic status of members of the P. leightoni complex using phylogenetic analyses and species distribution modelling (SDM) techniques. Maximum likelihood and Bayesian inference approaches were applied to ascertain the genetic relationships within the P. leightoni complex. Uncorrected p-distances were generated to assess the level of divergence between taxa of the P. leightoni complex relative to its African congeners. Furthermore, a General Mixed Yule Coalescent model and its Bayesian implementation were used as tree-based methods for species delimitation. Species distribution models were carried out using a maximum entropy approach that estimated the climate suitability for these putative taxa defining their distributions during the last glacial maximum, mid-Holocene and under current climatic conditions. The phylogenetic analyses recovered all individuals of the P. leightoni complex in a monophyletic clade. Furthermore, the study taxa show intraspecific level divergences between taxa of the P. leightoni complex and are suggested to collectively represent a single taxon based on the species delimitation analyses. Additionally, the species distribution models showed no difference between these taxa's spatial distribution suggesting that taxa of the P. leightoni complex are not ecologically distinct. Assuming the P. leightoni complex represents a single species, meeting the prerequisites of the unified general species concept, a taxonomic revision is necessary to assign the appropriate taxonomic rank. Both P. namibensis Broadley, 1975 and P. trinasalis Werner, 1902 should be considered synonyms of P. leightoni Boulenger, 1902. As a result, P. leightoni should be considered widespread and in need of a conservation reassessment, potentially removing its current threat status.
Predicting reptile species distributions and biogeographic patterns within Kruger National Park
(University of the Western Cape, 2018) Barends, Jody Michael; Maritz, Bryan
Knowledge of global reptile ecology is limited and there remains much to understand in terms of detailed reptile species information, including that of their distributions. In South Africa, despite being one of SANParks best-studied reserves, surprisingly little is known about the distributions and spatial ecology of reptiles within Kruger National Park (KNP). Management within KNP follows a strategic adaptive management strategy which monitors the statuses of animals using species or group specific indicators. Indicators are given predetermined upper and lower ranges of acceptable fluctuation before actions are taken. These ranges are referred to as thresholds of potential concern (TPCs), and for reptiles these are based on changes to their distributions across the landscape of KNP. An apparent lack of high-quality reptile distribution data inhibits the effective monitoring of the statuses of these animals within KNP, which in turn limits management and conservation options. In this study, I use several methods to quantify available reptile occurrence data which formed the foundations for predicting the distributions of these species across KNP by means of species distribution modelling, with a view to gaining novel insight into reptile assemblage structure across the landscape of KNP.
Quantifying competition in two co-occurring southern African psammophiinae snakes: Psammophis crucifer and psammophylax r. Rhombeatus
(University of Western Cape, 2020) Tokota, Silindokuhle; Maritz, Bryan
Studies on snake competitive interactions have relatively been well documented globally, however, those examples tend to be dominated by non-African examples. Africa has a large and spectacular reptile diversity and yet robust and empirical studies on snake population ecology remain poorly understood or documented. Given the close phylogenetic relationship between the two species, as well as the remarkable similarities in overall appearance, morphology, reproductive biology, and most importantly geographic distribution, Psammophis crucifer and Psammophylax rhombeatus offered an ideal study system in which to ask questions related to interspecific competition and niche partitioning.
Reptile community responses to an avian ecosystem engineer
(University of the Western Cape, 2023) Buckley, Emma Elizabeth; Maritz, Bryan
In the Kalahari, sociable weavers (Philetairus socius) build huge, communal nests. Although Kalahari tree skinks (Trachylepis spilogaster) are known to preferentially use trees that host such colonies, the degree to which other reptile species use these trees is not well documented. Sociable weaver nests can provide benefits to different animals and can provide food and shelter for reptiles. I trapped reptiles at Tswalu Kalahari Reserve to assess the impact of sociable weaver colonies on selected reptile populations during weaver breeding and non-breeding seasons. I tested the hypothesis that the reptile community under trees with colonies would differ in diversity compared to those at nearby control trees without colonies, because the sociable weaver colonies provide more resources (food and shelter) to the reptiles than trees without colonies. I also tested whether the abundance of the Kalahari tree skink, Cape cobra (Naja nivea), and Cape thick-toed gecko (Pachydactylus capensis) under trees with colonies differed from those at nearby control trees without colonies, as these were the most abundant species found at the sociable weaver nests. Trapping for 11 days at 24 sites (12 colony trees, 12 control trees), on two occasions, once in March when weavers were breeding, and once in September before they were breeding, resulted in 665 total detections of 13 reptile species, including 148 total recaptures. Mean species richness was significantly higher at colony trees than at control trees during both surveys (2.5 times higher when the birds were breeding; 1.6 times higher when the birds were not breeding). Poisson N-mixture modelling revealed that Kalahari tree skink was significantly more abundant on colony trees during the breeding (3.2 times) and non-breeding seasons (3.7 times).
Sampling bias in reptile occurrence data for the Kruger National Park
(AOSIS, 2020) Barends, Jody; Pietersen, Darren; Zambatis, Guinevere; Tye, Donovan; Maritz, Bryan
To effectively conserve and manage species, it is important to (1) understand how they are spatially distributed across the globe at both broad and fine spatial resolutions and (2) elucidate the determinants of these distributions. However, information pertaining to the distributions of many species remains poor as occurrence data are often scarce or collected with varying motivations, making the resulting patterns susceptible to sampling bias. Exacerbating an already limited quantity of occurrence data with an assortment of biases hinders their effectiveness for research, thus making it important to identify and understand the biases present within species occurrence data sets. We quantitatively assessed occurrence records of 126 reptile species occurring in the Kruger National Park (KNP), South Africa, to quantify the severity of sampling bias within this data set. We collated a data set of 7118 occurrence records from museum, literature and citizen science sources and analysed these at a biologically relevant spatial resolution of 1 km × 1 km. As a result of logistical challenges associated with sampling in KNP, approximately 92% of KNP is data deficient for reptile occurrences at the 1 km × 1 km resolution. Additionally, the spatial coverage of available occurrences varied at species and family levels, and the majority of occurrence records were strongly associated with publicly accessible human infrastructure. Furthermore, we found that sampled areas within KNP were not necessarily ecologically representative of KNP as a whole, suggesting that areas of unique environmental space remain to be sampled. Our findings highlight the need for substantially greater sampling effort for reptiles across KNP and emphasise the need to carefully consider the sampling biases within existing data should these be used for conservation management decision-making. Modelling species distributions could potentially serve as a short-term solution, but a concomitant increase in surveys across the park is needed. Conservation implications: The sampling biases present within KNP reptile occurrence data inhibit the inference of fine-scale species distributions within and across the park, which limits the usage of these data towards meaningfully informing conservation management decisions as applicable to reptile species in KNP
Scale-dependent orientation in movement paths: a case study of an African viper
(Wiley, 2016) Maritz, Bryan; Alexander, Graham J.
Decisions relating to the orientation of movement by animals and how this translates into movement patterns can occur at multiple spatial scales simultaneously, but this interaction is poorly understood for many groups of animals. Using the tracks left by moving snakes in their sandy habitat, we studied the movement paths of the African snake Bitis schneideri (Namaqua dwarf adder) for evidence of broad-scale directional persistence and short-range avoidance of exposure. Although snakes clearly displayed directional persistence, they preferentially moved to nearby shrubs, thereby minimizing exposure to solar and thermal radiation and/or predation. Thus, snakes made decisions relating to orientation at a minimum of two scales, the interaction of which resulted in snakes moving ≈17% (mean straightness index = 0.85) further than the simple broad-scale straight-line distance. We assert that the actual path chosen by moving snakes represents a trade-off of various costs and risks that include risk of predation, exposure to the elements, time and energy expenditure. Our study highlights the need for cognizance of the possibility of the scale dependence of orientation and movement in studies of snake movement, and adds to a growing literature demonstrating previously unrecognized behavioural complexity in snakes.
Sharing for science: High-resolution trophic interactions revealed rapidly by social media
(PeerJ, 2020) Maritz, Robin A.; Maritz, Bryan
Discrete, ephemeral natural phenomena with low spatial or temporal predictability are incredibly challenging to study systematically. In ecology, species interactions, which constitute the functional backbone of ecological communities, can be notoriously difficult to characterise especially when taxa are inconspicuous and the interactions of interest (e.g., trophic events) occur infrequently, rapidly, or variably in space and time. Overcoming such issues has historically required significant time and resource investment to collect sufficient data, precluding the answering of many ecological and evolutionary questions. Here, we show the utility of social media for rapidly collecting observations of ephemeral ecological phenomena with low spatial and temporal predictability by using a Facebook group dedicated to collecting predation events involving reptiles and amphibians in sub-Saharan Africa. We collected over 1900 independent feeding observations using Facebook from 2015 to 2019 involving 83 families of predators and 129 families of prey. Feeding events by snakes were particularly well-represented with close to 1,100 feeding observations recorded.
Snake body size frequency distributions are robust to the description of novel species
(Ecological Society of America, 2016) Maritz, Bryan; Kgaditse, Mimmie; Alexander, Graham John
The accurate interpretation of body size frequency distributions (BSFDs) has important implications for understanding large-scale ecological and evolutionary patterns because they are shaped by ecological and evolutionary processes. However, incomplete species sampling, resulting either from a paucity of reliable body size data or because those species have not yet been detected/described, also has the potential to influence BSFD shape and skew when missing species are disproportionately large or small. Missing species may thus influence the skew of a BSFD, misleading subsequent interpretations. In contrast to many vertebrate taxa, the BSFDs of snakes appears to be log-normal, with no significant skew. However, the influence of imperfect species sampling on these distributions has however not been previously evaluated. We collated body size data and date of description for 300 species of elapid, and 264 species of viperid snakes and assessed the shape and skew of the BSFD of each clade. Next we tested the hypothesis that date of description was related to body size in each clade. We show that the BSFDs of elapid and viperid snakes are both log-normal, with no significant skew. Moreover we demonstrate size-related differences in the probability of detection (as measured by year of description) among elapids and vipers: in both lineages, species with larger body size tend to have been described earlier. However, simulations testing the effect of the addition of novel, small-bodied species revealed that the distributions of elapids and vipers are not highly vulnerable to becoming significantly skewed in the future. Our results support current interpretations of BSFDs in snakes that suggest a lack of size-associated biases in the speciation and extinction rates of this clade.
Snake community structure in a heterogeneous fynbos ecosystem
(University of the Western Cape, 2022) Cloete, Bryan Nathan; Maritz, Bryan
Snakes occur in most terrestrial ecosystems, forming an integral part of many ecological communities. A diverse community of snakes in an ecosystem can indicate a complex habitat structure capable of supporting a diverse assemblage of faunal species. I utilised multiple non-invasive and non-lethal trapping methods to quantify diversity metrics for the snake community occurring in a heterogeneous fynbos ecosystem within the Koeberg Private Nature Reserve (KPNR) over one-year period. Additionally, I sampled the species richness and abundance of snake predators and prey within the reserve and tested the hypothesis that the snake community was impacted by the abundance and diversity of snake predators and prey. I performed several statistical analyses to identify the factors that influence the snake community structure observed in KPNR.
Snake predators of bird eggs: A review and bibliography
(Wiley, 2022) Barends, Jody M.; Maritz, Bryan
Snakes are frequent predators of bird nests and therefore potentially have an important impact on bird population dynamics. However, while many species are known to consume nestlings and chicks, few species have been recorded consuming bird eggs. To effectively quantify the effects of bird egg predation by snakes on bird demographics, a key first step is to identify which snake species consume bird eggs. Unfortunately, detailed information on the dietary habits of most snakes is scarce and feeding records are poorly cataloged, making it difficult to ascertain which species do and do not eat bird eggs. We reviewed the literature and online community science reports to compile a global list of confirmed snake predators of bird eggs. In total, we gathered 471 feeding records of 123 snake taxa consuming the eggs of at least 210 bird species from 238 individual data sources. Geographical locations of records disproportionately represented well-sampled regions, and we infer that many snake species not included on our list also consume bird eggs.
Specialized morphology, not relatively large head size, facilitates competition between a small-bodied specialist and large-bodied generalist competitors
(Journal of Zoology, 2021) Barends, Jody, M; Maritz, Bryan
Interspecific competition for limited resources should theoretically occur between species that are morphologically similar to each other. Consequently, species that reduce competition by adapting to specialize on a specific resource should be morphologically disparate to sympatric contemporaries and show evidence of phenotypic specialization. However, few studies have compared the morphologies of specialist and generalist competitors. In this context, we compare the feeding morphology and diet of an obligate, specialist, bird-egg-eating snake to three sympatric generalists that only facultatively consume bird eggs. We measured and compared body and head morphology of preserved museum specimens of each of four, syntopic snake species from southern Africa: the obligate bird-egg-eating rhombic egg-eater (Dasypeltis scabra), and the facultative bird-egg-eating boomslang (Dispholidus typus), cape cobra (Naja nivea) and mole snake (Pseudaspis cana). Given the physical challenges of consuming bird eggs in snakes, we predicted that consumption of bird eggs would be facilitated by the evolution of relatively larger heads in the smaller bodied Dasypeltis. We found that head size was not phylogenetically conserved in the clades of these taxa and that contrary to our expectations, the specialist egg-eaters evolved to possess significantly smaller heads relative to body size than their competitors. We found a positive relationship between dietary niche breadth and head size within these species and their close relatives. Thus, relatively large-headed species have evolved diverse diets that overlap with the restricted diets of the small-headed specialist thereby producing this atypical competitive interaction. Our findings suggest that specialized adaptations can decouple typical body-size-constrained competition dynamics between sympatric snake species and highlight the complexity of the origins of dietary specialization.

Browsing by Author "Maritz, Bryan"

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