The morphological and molecular variation of southern African Nannocharax (Characiformes: Distichodontidae), and its taxonomic implications
- Authors: Smith, Timothy
- Date: 2019
- Subjects: Nannocharax -- Africa, Southern , Distichodontidae -- Africa, Southern , Freshwater fishes -- Africa, Southern , Nannocharax -- Genetics -- Africa, Southern , Distichodontidae -- Genetics -- Africa, Southern , Freshwater fishes -- Genetics -- Africa, Southern
- Language: English
- Type: text , Thesis , Masters , MSc
- Identifier: http://hdl.handle.net/10962/69212 , vital:29446
- Description: Nannocharax is the most species rich genus in the family Distichodontidae, being currently represented by 41 species. The genus is widely distributed across much of sub-Saharan Africa, with a range extending from the Zambezi ichthyofaunal province in the south to the Nilo-Sudan ichthyofaunal province in the north. In southern Africa, the genus is currently represented by four species, Nannocharax dageti Jerep, Vari, & Vreven, 2014, N. machadoi (Poll, 1967), N. macropterus Pellegrin, 1926, and N. multifasciatus Boulenger, 1923. Each of these species exhibit considerable intraspecific pigmentation pattern variation across their respective distribution ranges, suggesting that the current taxonomy possibly underestimates the taxonomic diversity of Nannocharax species in southern Africa. Much pigmentation pattern variation within these southern African species has been observed by both collectors and scientists in the field, prompting an investigation into the extent of this morphological variation as well as what molecular variation may occur as well. The genus displays a high degree of morphological conservatisim, making it difficult to assign external morphological characters as diagnostic. To this end, this study was conducted to determine the extent of diversity of this genus in the region, employing an integrative approach with traditional morphological analysis techniques as well as sequencing the ‘barcoding gene’, cytochrome oxidase I, testing the hypothesis that there is a greater, hidden diversity of this genus in the region than currently recognised. This study aims to identify these potential lineages and accurately map their distributions. Phylogenetic analyses were performed using maximum parsimony, maximum likelihood, and Bayesian inference, using the mitochondrial cytochrome oxidase I gene region. Massive genetic divergence was detected between populations of taxa previously considered to be singular, widely distributed species. The three approaches of phylogenetic inference used in this study yielded trees of comparable overall topology, with the exception of the maximum parsimony tree which indicated additional lineages within the southern African N. multifasciatus group. These analyses revealed four deeply divergent (1.3 – 12.3%) lineages within southern African N. macropterus, as well as two deeply divergent (0.4-14.6%) populations from the Congo ichthyofaunal region, the lineages here named “N. macropterus Congo 1” and “N. macropterus Congo 2”. Within the southern African region, two deeply divergent (10.3%) lineages of N. macropterus were identified from the Okavango River system, identified as “N. macropterus Okavango 2” lineage restricted to the Cuito-Canavale tributary, and “N. macropterus Okavango 1” distributed throughout the remainder of the Okavango system. “N. macropterus Okavango 2” shares a closer relationship with the unique lineage from the Kwanza ichthyofaunal region, named N. macropterus “Kwanza”, which itself is deeply divergent from the N. macropterus “Okavango 1”, N. macropterus “Zambezi”, N. macropterus “Congo 1” and N. macropterus “Congo 2” lineages (3.1-14.4%). Principal component analyses (PCA) and discriminant function analyses (DFA) produced overlapping clusters for all identified lineages, with the exception of the N. macropterus “Kwanza” lineage, which in all analyses clustered away from the other lineages. Analysis of variance (ANOVA) and Kruskall-Wallis tests indicated significant differences in means between character traits between lineages, however, overlap in measurements and counts occurred in all instances except between the N. macropterus “Kwanza” and N. macropterus Congo lineages. However the N. macropterus “Kwanza” lineages could be distinguished from the other lineages by generally smaller fin lengths (dorsal fin 19.5%SL vs 20.0-22.1%SL in others; pectoral fin 16.5%SL vs 20.6-21.8%SL in others; pelvic fin 18.3%SL vs 21.3-22.4) and pigmentation pattern differences. The N. macropterus species group displayed extensive pigmentation pattern variation, to the extent that five pattern grades could be used to classify them. These pattern grades, while not specific to river systems, showed patterns similar to that which was seen in the molecular analyses and could be linked to lineages with only minor overlap between them. Three lineages of N. multifasciatus were identified, with two occurring in the southern African region, each corresponding to a river system, being the N. multifasciatus “Zambezi” and N. multifasciatus “Okavango” lineages. This species group displayed shallower divergence between lineages than did the N. macropterus group, at 2.5% genetic distance. Genetic analysis inferred a closer relationship between the N. multifasciatus “Zambezi” and N. multifasciatus “Congo” lineages than with the N. multifasciatus “Okavango” lineage. Morphological PCA and DFA analyses indicated morphological divergence of the N. multifasciatus “Congo” lineage, with generally larger proportional measurements than southern African specimens (body width 12.6%SL vs 9.5-9.7%SL; body depth 26.6%SL vs 21.6-21.9%SL; head width 12.0%SL vs 10.0-10.4%SL). PCA, DFA, and measurements show a near complete overlap between the N. multifasciatus “Okavango” and N. multifasciatus “Zambezi” lineages. Pigmentation pattern variation occurred within this group, but none that could be assigned to a particular lineage. The N. machadoi species group in southern Africa consists of five lineages: N. machadoi “Zambezi 1”, N. machadoi “Zambezi 2”, N. machadoi “Kafue 1”, N. machadoi “Kafue 2”, and N. machadoi “Okavango”. This group displayed shallower genetic divergence between lineages than the other southern African Nannocharax species groups (0.4-1.3%). This shallow genetic divergence is paralleled by near complete morphological overlap, with PCA and DFA producing overlapping clusters, and measurements, meristics, and pigmentation pattern metrics consisting of very similar values for the lineages. These results indicate that what is considered to be “N. macropterus” in southern Africa should not be named as such. The N. macropterus “Zambezi” and the N. macropterus “Okavango 1” lineages, are misidentifications of Nannocharax dageti. Other “N. macropterus” from the southern African region possesses fewer circumpeduncular scales than the true N. macropterus as described by Pellegrin (1926), and require taxonomic re-evaluation, each here being recognised as a unique lineage with species status, here named N. macropterus “Okavango 2” and N. macropterus “Kwanza”. In particular, N. macropterus “Kwanza” displays deep genetic divergence as well as morphological dissimilarity with the other southern African “N. macropterus” groups. Nannocharax fasciolaris and N. monardi are here placed as junior synonyms of N. multifasciatus, owing to vast overlaps in measurements and character counts of these species and N. multifasciatus, which is also known to occur within the same geographical distribution, as well as dubious arguments from the original publications in delineating these species from N. multifasciatus. Therefore, there is insufficient evidence indicating the presence of multiple species originating from the Okavango system, where it is here indicated that only a single lineage of banded, adipose fin-bearing Nannocharax occurs, namely N. multifasciatus “Okavango”.
- Full Text:
- Date Issued: 2019
- Authors: Smith, Timothy
- Date: 2019
- Subjects: Nannocharax -- Africa, Southern , Distichodontidae -- Africa, Southern , Freshwater fishes -- Africa, Southern , Nannocharax -- Genetics -- Africa, Southern , Distichodontidae -- Genetics -- Africa, Southern , Freshwater fishes -- Genetics -- Africa, Southern
- Language: English
- Type: text , Thesis , Masters , MSc
- Identifier: http://hdl.handle.net/10962/69212 , vital:29446
- Description: Nannocharax is the most species rich genus in the family Distichodontidae, being currently represented by 41 species. The genus is widely distributed across much of sub-Saharan Africa, with a range extending from the Zambezi ichthyofaunal province in the south to the Nilo-Sudan ichthyofaunal province in the north. In southern Africa, the genus is currently represented by four species, Nannocharax dageti Jerep, Vari, & Vreven, 2014, N. machadoi (Poll, 1967), N. macropterus Pellegrin, 1926, and N. multifasciatus Boulenger, 1923. Each of these species exhibit considerable intraspecific pigmentation pattern variation across their respective distribution ranges, suggesting that the current taxonomy possibly underestimates the taxonomic diversity of Nannocharax species in southern Africa. Much pigmentation pattern variation within these southern African species has been observed by both collectors and scientists in the field, prompting an investigation into the extent of this morphological variation as well as what molecular variation may occur as well. The genus displays a high degree of morphological conservatisim, making it difficult to assign external morphological characters as diagnostic. To this end, this study was conducted to determine the extent of diversity of this genus in the region, employing an integrative approach with traditional morphological analysis techniques as well as sequencing the ‘barcoding gene’, cytochrome oxidase I, testing the hypothesis that there is a greater, hidden diversity of this genus in the region than currently recognised. This study aims to identify these potential lineages and accurately map their distributions. Phylogenetic analyses were performed using maximum parsimony, maximum likelihood, and Bayesian inference, using the mitochondrial cytochrome oxidase I gene region. Massive genetic divergence was detected between populations of taxa previously considered to be singular, widely distributed species. The three approaches of phylogenetic inference used in this study yielded trees of comparable overall topology, with the exception of the maximum parsimony tree which indicated additional lineages within the southern African N. multifasciatus group. These analyses revealed four deeply divergent (1.3 – 12.3%) lineages within southern African N. macropterus, as well as two deeply divergent (0.4-14.6%) populations from the Congo ichthyofaunal region, the lineages here named “N. macropterus Congo 1” and “N. macropterus Congo 2”. Within the southern African region, two deeply divergent (10.3%) lineages of N. macropterus were identified from the Okavango River system, identified as “N. macropterus Okavango 2” lineage restricted to the Cuito-Canavale tributary, and “N. macropterus Okavango 1” distributed throughout the remainder of the Okavango system. “N. macropterus Okavango 2” shares a closer relationship with the unique lineage from the Kwanza ichthyofaunal region, named N. macropterus “Kwanza”, which itself is deeply divergent from the N. macropterus “Okavango 1”, N. macropterus “Zambezi”, N. macropterus “Congo 1” and N. macropterus “Congo 2” lineages (3.1-14.4%). Principal component analyses (PCA) and discriminant function analyses (DFA) produced overlapping clusters for all identified lineages, with the exception of the N. macropterus “Kwanza” lineage, which in all analyses clustered away from the other lineages. Analysis of variance (ANOVA) and Kruskall-Wallis tests indicated significant differences in means between character traits between lineages, however, overlap in measurements and counts occurred in all instances except between the N. macropterus “Kwanza” and N. macropterus Congo lineages. However the N. macropterus “Kwanza” lineages could be distinguished from the other lineages by generally smaller fin lengths (dorsal fin 19.5%SL vs 20.0-22.1%SL in others; pectoral fin 16.5%SL vs 20.6-21.8%SL in others; pelvic fin 18.3%SL vs 21.3-22.4) and pigmentation pattern differences. The N. macropterus species group displayed extensive pigmentation pattern variation, to the extent that five pattern grades could be used to classify them. These pattern grades, while not specific to river systems, showed patterns similar to that which was seen in the molecular analyses and could be linked to lineages with only minor overlap between them. Three lineages of N. multifasciatus were identified, with two occurring in the southern African region, each corresponding to a river system, being the N. multifasciatus “Zambezi” and N. multifasciatus “Okavango” lineages. This species group displayed shallower divergence between lineages than did the N. macropterus group, at 2.5% genetic distance. Genetic analysis inferred a closer relationship between the N. multifasciatus “Zambezi” and N. multifasciatus “Congo” lineages than with the N. multifasciatus “Okavango” lineage. Morphological PCA and DFA analyses indicated morphological divergence of the N. multifasciatus “Congo” lineage, with generally larger proportional measurements than southern African specimens (body width 12.6%SL vs 9.5-9.7%SL; body depth 26.6%SL vs 21.6-21.9%SL; head width 12.0%SL vs 10.0-10.4%SL). PCA, DFA, and measurements show a near complete overlap between the N. multifasciatus “Okavango” and N. multifasciatus “Zambezi” lineages. Pigmentation pattern variation occurred within this group, but none that could be assigned to a particular lineage. The N. machadoi species group in southern Africa consists of five lineages: N. machadoi “Zambezi 1”, N. machadoi “Zambezi 2”, N. machadoi “Kafue 1”, N. machadoi “Kafue 2”, and N. machadoi “Okavango”. This group displayed shallower genetic divergence between lineages than the other southern African Nannocharax species groups (0.4-1.3%). This shallow genetic divergence is paralleled by near complete morphological overlap, with PCA and DFA producing overlapping clusters, and measurements, meristics, and pigmentation pattern metrics consisting of very similar values for the lineages. These results indicate that what is considered to be “N. macropterus” in southern Africa should not be named as such. The N. macropterus “Zambezi” and the N. macropterus “Okavango 1” lineages, are misidentifications of Nannocharax dageti. Other “N. macropterus” from the southern African region possesses fewer circumpeduncular scales than the true N. macropterus as described by Pellegrin (1926), and require taxonomic re-evaluation, each here being recognised as a unique lineage with species status, here named N. macropterus “Okavango 2” and N. macropterus “Kwanza”. In particular, N. macropterus “Kwanza” displays deep genetic divergence as well as morphological dissimilarity with the other southern African “N. macropterus” groups. Nannocharax fasciolaris and N. monardi are here placed as junior synonyms of N. multifasciatus, owing to vast overlaps in measurements and character counts of these species and N. multifasciatus, which is also known to occur within the same geographical distribution, as well as dubious arguments from the original publications in delineating these species from N. multifasciatus. Therefore, there is insufficient evidence indicating the presence of multiple species originating from the Okavango system, where it is here indicated that only a single lineage of banded, adipose fin-bearing Nannocharax occurs, namely N. multifasciatus “Okavango”.
- Full Text:
- Date Issued: 2019
Atlas of Southern African freshwater fishes
- Scott, L E P, Skelton, P H, Booth, A J, Verheust, L, Dooley, J, Harris, R
- Authors: Scott, L E P , Skelton, P H , Booth, A J , Verheust, L , Dooley, J , Harris, R
- Date: 2006
- Subjects: Freshwater fishes -- South Africa , Freshwater fishes -- Africa, Southern , Freshwater fishes -- Africa, Southern -- Identification
- Language: English
- Type: text , book
- Identifier: http://hdl.handle.net/10962/199097 , vital:46068
- Description: Worldwide concern over the loss of biological diversity during the past decade has provided the impetus for the development of biodiversity databases and biological atlas projects in order to make information accessible. This Geographic Information Systems-based Atlas of Southern African Freshwater Fishes is the first of its kind for fishes in Africa and, as such, can be seen as contributing towards this global trend of consolidation and refining of biodiversity information. The technological advances made in computing systems over the past decade have facilitated the extraction and utilisation of much of the untapped information in museum collections. The size of most natural history collections had tended to prohibit any comprehensive applied use of the specimens on a large scale, but the computerisation of these collections has proved to be an invaluable tool to systematists, taxonomists and ecologists, by allowing faster and more efficient access to specimen record data of interest. With the automation of many biological databases, public attention has been brought to their potential contribution to biodiversity studies and conservation efforts around the world. The application of Geographic Information Systems (GIS) in the field of southern African ichthyology and freshwater fisheries management has been limited, until now, by the lack of good-quality, comprehensive data sets, adequate budgets and GIS skills. Sophisticated investigations of fish distributions for conservation and management purposes need a powerful computing platform. This is realised in the Atlas of Southern African Freshwater Fishes, which has been tested and used for biogeographic analysis in the southern African region. The atlas was established using specimen records from nineteen museum collections, and currently contains 35145 geo-referenced specimen records, collected between 1885 and 2000 in eleven countries of southern Africa, which represent 254 fish species from 37 families. The Atlas of Southern African Freshwater Fishes was developed with the objective of being a dynamic, powerful information system. It has been constructed to allow the import of additional information at any stage, and it is designed to be as accessible and as user-friendly as current technology allows. This Atlas is the product of the successful integration of fish distributional information with other spatial data layers to provide a new tool for the study of freshwater fish in southern Africa.
- Full Text:
- Date Issued: 2006
- Authors: Scott, L E P , Skelton, P H , Booth, A J , Verheust, L , Dooley, J , Harris, R
- Date: 2006
- Subjects: Freshwater fishes -- South Africa , Freshwater fishes -- Africa, Southern , Freshwater fishes -- Africa, Southern -- Identification
- Language: English
- Type: text , book
- Identifier: http://hdl.handle.net/10962/199097 , vital:46068
- Description: Worldwide concern over the loss of biological diversity during the past decade has provided the impetus for the development of biodiversity databases and biological atlas projects in order to make information accessible. This Geographic Information Systems-based Atlas of Southern African Freshwater Fishes is the first of its kind for fishes in Africa and, as such, can be seen as contributing towards this global trend of consolidation and refining of biodiversity information. The technological advances made in computing systems over the past decade have facilitated the extraction and utilisation of much of the untapped information in museum collections. The size of most natural history collections had tended to prohibit any comprehensive applied use of the specimens on a large scale, but the computerisation of these collections has proved to be an invaluable tool to systematists, taxonomists and ecologists, by allowing faster and more efficient access to specimen record data of interest. With the automation of many biological databases, public attention has been brought to their potential contribution to biodiversity studies and conservation efforts around the world. The application of Geographic Information Systems (GIS) in the field of southern African ichthyology and freshwater fisheries management has been limited, until now, by the lack of good-quality, comprehensive data sets, adequate budgets and GIS skills. Sophisticated investigations of fish distributions for conservation and management purposes need a powerful computing platform. This is realised in the Atlas of Southern African Freshwater Fishes, which has been tested and used for biogeographic analysis in the southern African region. The atlas was established using specimen records from nineteen museum collections, and currently contains 35145 geo-referenced specimen records, collected between 1885 and 2000 in eleven countries of southern Africa, which represent 254 fish species from 37 families. The Atlas of Southern African Freshwater Fishes was developed with the objective of being a dynamic, powerful information system. It has been constructed to allow the import of additional information at any stage, and it is designed to be as accessible and as user-friendly as current technology allows. This Atlas is the product of the successful integration of fish distributional information with other spatial data layers to provide a new tool for the study of freshwater fish in southern Africa.
- Full Text:
- Date Issued: 2006
The development of a geographic information systems based atlas of southern African freshwater fish, and its application to biogeographic analysis
- Scott, Lucy Elizabeth Powell
- Authors: Scott, Lucy Elizabeth Powell
- Date: 2000
- Subjects: Freshwater fishes -- Africa, Southern , Freshwater fishes -- Geographical distribution -- Atlases , Freshwater fishes -- Africa, Southern -- Geographical distribution -- Atlases , Geographic information systems
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5256 , http://hdl.handle.net/10962/d1005099 , Freshwater fishes -- Africa, Southern , Freshwater fishes -- Geographical distribution -- Atlases , Freshwater fishes -- Africa, Southern -- Geographical distribution -- Atlases , Geographic information systems
- Description: A Geographic Information Systems (GIS) atlas of southern African freshwater fish was developed for the SADC countries from natural history collection specimens, hydrological, topographical and climatological data. The primary purpose of the development of the atlas of freshwater fish was the construction of a practical framework to transform vast amounts of existing biological data for use in research and management of aquatic resources. The database of freshwater fish collection specimens that was incorporated into the atlas, was developed in association with ALCOM (Aquatic Resources Management for Local Community Development Programme). The development of advanced computing and GIS technology has increased the scope of biological atlas projects by facilitating the integration of large amounts of spatial data to produce derived databases for specific applications. The atlas of freshwater fish was constructed using TNTmips GIS software as the most practical system available for managing and analysing biological data with a spatial component. The atlas contains 35 180 comprehensive distribution records of 735 species of fish. It has many applications as an inventory of ichthyofaunal spatial biodiversity, including those of conservation planning, environmental assessment and biogeographic research. Biogeographic studies have traditionally been subjective due to the logistical problems of working with large amounts of distribution data, although some small-scale quantitative research has been carried out in the past. The content of the atlas of freshwater fish is tested with respect to these previous studies, on known patterns of freshwater fish distributions, and the analytical capability of the atlas is tested and demonstrated with some new preliminary approaches to the analysis of freshwater fish distributions in southern Africa.
- Full Text:
- Date Issued: 2000
- Authors: Scott, Lucy Elizabeth Powell
- Date: 2000
- Subjects: Freshwater fishes -- Africa, Southern , Freshwater fishes -- Geographical distribution -- Atlases , Freshwater fishes -- Africa, Southern -- Geographical distribution -- Atlases , Geographic information systems
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5256 , http://hdl.handle.net/10962/d1005099 , Freshwater fishes -- Africa, Southern , Freshwater fishes -- Geographical distribution -- Atlases , Freshwater fishes -- Africa, Southern -- Geographical distribution -- Atlases , Geographic information systems
- Description: A Geographic Information Systems (GIS) atlas of southern African freshwater fish was developed for the SADC countries from natural history collection specimens, hydrological, topographical and climatological data. The primary purpose of the development of the atlas of freshwater fish was the construction of a practical framework to transform vast amounts of existing biological data for use in research and management of aquatic resources. The database of freshwater fish collection specimens that was incorporated into the atlas, was developed in association with ALCOM (Aquatic Resources Management for Local Community Development Programme). The development of advanced computing and GIS technology has increased the scope of biological atlas projects by facilitating the integration of large amounts of spatial data to produce derived databases for specific applications. The atlas of freshwater fish was constructed using TNTmips GIS software as the most practical system available for managing and analysing biological data with a spatial component. The atlas contains 35 180 comprehensive distribution records of 735 species of fish. It has many applications as an inventory of ichthyofaunal spatial biodiversity, including those of conservation planning, environmental assessment and biogeographic research. Biogeographic studies have traditionally been subjective due to the logistical problems of working with large amounts of distribution data, although some small-scale quantitative research has been carried out in the past. The content of the atlas of freshwater fish is tested with respect to these previous studies, on known patterns of freshwater fish distributions, and the analytical capability of the atlas is tested and demonstrated with some new preliminary approaches to the analysis of freshwater fish distributions in southern Africa.
- Full Text:
- Date Issued: 2000
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