Corresponding author: Trond Andersen (
Academic editor: E. Rázuri-Gonzales
This paper is primarily based on collections in Tanzania and Ghana in 1990–1991 and 1991–1994, respectively. In all, 46 species of
Blahnik R, Andersen T (2022) New species of the genus
Currently, 115 species of
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Ghana, Guinea, Ivory Coast, Madagascar, Mali, Togo] |
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Ivory Coast, Mali, Togo] |
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*Described or redescribed in this article. **Species unassigned to subgroup are either based on females, inadequately illustrated, morphologically isolated from other species, known only from Madagascar or other islands, or some combination of above.
Most of the material on which this paper is based was collected during two projects by the Department of Natural History, University Museum of Bergen, Norway (former Museum of Zoology, University of Bergen). The field work during the first project targeted the fauna in the mountain rainforests in the West Usambara Mountains in northeastern Tanzania. The study area was located near the Mazumbai Forest Reserve, where caddisflies were collected in 1990 and 1991 along the Kaputu Stream (
Malaise trap across Kaputu Stream at 1535 m altitude in the Mazumbai Forest Reserve in the West Usambara Mountains, northeastern Tanzania (Photo: Trond Andersen).
The West Usambara Mountains belong to the Eastern Arc, a chain of mountains that stretch from the Taita Hills in Kenya in the north, south to the Udzungwa and Mahenge Mountains in Tanzania. They were formed at least one hundred million years ago along a fault lying to the east of the East African Rift, which is a more recent structure. Approximately thirty million years ago, all this area was covered by extensive rainforest. During a period, some ten million years ago, when the climate was cooler and drier, the lowland forests were converted to savannah, leaving the mountain ranges as “islands” where the tropical forests continued to flourish, fed by moisture from the Indian Ocean. This isolation of each mountain range has led to a great deal of endemism, and a very diverse flora and fauna (
During the second project,
Caddisflies were collected in several localities in the southern, forested part of Ghana. The Ankasa Conservation Area is situated in the Western Region in southwestern Ghana near the border to the Ivory Coast. The conservation area is ~ 500 square kilometers and incorporates the Nini Suhien National Park in the north and the Ankasa Forest Reserve in the south. The altitude varies from 35 m to 170 m and there are three larger rivers and many smaller streams in the area (Fig.
Malaise trap across a small tributary to Ankasa River in the wet evergreen forest in Ankasa Forest Reserve, southwestern Ghana (Photo: Jostein Kjærandsen).
The Kakum National Park is situated in the Central Region in southern Ghana. It was established as a reserve in 1931 and received the status as a national park in 1992. The Park covers 375 square kilometers and is generally flat with only a few undulating hills ranging 150–250 m above sea level. The forest is classified as moist semi-deciduous forest (
Most of the material treated here is from a study of the caddisfly community along a headwater stream in the Agumatsa Wildlife Sanctuary in the Volta Region, situated in the transition zone in the eastern part of Ghana (
The lower waterfall of Agumatsa-Nubui headwater stream in the Agumatsa Wildlife Sanctuary, eastern Ghana (Photo: Jostein Kjæranden).
Northern Ghana is covered with savannah and is crossed by several large rivers flowing southwards. Caddisflies were mainly collected at two of these rivers, the Black Volta and Oti rivers. The Black Volta originates in Burkina Faso and in Ghana it forms the border with Ivory Coast before it joins the White Volta. The Oti River has its headwaters in Benin and Burkina Faso and flows through Benin and Togo before it joins the Volta River in Ghana.
Because of the frequent use of species group names to refer to taxa in the subgenus
The fieldwork during the project in Tanzania targeted the fauna in the mountain rainforests in the West Usambara Mountains in northeastern Tanzania. The study area is located near the Mazumbai Forest Reserve, where caddisflies were collected along the Kaputu Stream (
During the second project,
During the project in Ghana, caddisflies were mainly collected in light traps, Malaise traps, and with sweep nets and preserved in 80% ethanol. The females were tentatively associated with the males based on their co-occurrence with males and relative similarity. Because of this uncertainty, and because multiple species were collected at many sites, females of most new species are not listed in the paratype series, but instead included as additional material.
Illustrations were made using an ocular grid and inked in Adobe Illustrator. All figures are drawn of the left side or appendage, unless otherwise noted. Setation is generally omitted from the right side. Terminology used follows
Type material is deposited in the collections of the
The species described here can be placed into two well-defined species groups, the
Most of the mainland African species of the
The
In addition to new species described in this paper, other species from Africa and Madagascar were also assessed for their relationships. A list of the species and their assignment to subgroups within the major species groups is found in Table
The first characterization of evolutionary relationships in the genus
The first, and most useful, character defining the
A character usually used to define the subgenus
The presence of an inflected Rs vein is often accompanied by other character state changes and can be useful for assessing relationships within the subgenus when the characters are considered in combination. Characters frequently occurring in the
Described species of the
Members of this subgroup have the same generalized features that characterize the
In the molecular analysis of species in the genus
Ghana –
The two species included in this subgroup are very similar. The differences, as evident from the original illustrations, lie mostly in the shape and length of the lateral lobes of tergum X, the relative shape of the apices of these lobes, as well as in the more bulbous bases of the phallic spines in
Ghana, Ivory Coast.
The
Species of the
The subgroup is probably most closely related to the
In the study by
Ghana –
Phallobase with ventral apex greatly produced and strongly bent, apex rounded; phallic spines both rather short; inferior appendage tapered, bent, acute apically, cusps of ventromesal margin not evident in lateral view.
Angola, Democratic Republic of the Congo, Ghana.
Cape Verde ● 1♂; Brava, Fajâ d’ Agua; 100 m a.s.l.; 17 Feb. 2007; E Aistleitner leg.;
Phallobase with ventral margin greatly produced and strongly bent, apex enlarged and subtruncate, dorsal margin slightly upturned; phallic spines both moderately elongate and narrow; inferior appendage with dorsal projection abruptly narrowed, nearly uniform in width and distinctly bent, apex acute, ventromesal cusps of inferior appendage often both evident in lateral view (character possibly variable or inconsistent).
Our illustration closely matches that presented by
Burkina Faso, Cape Verde, Democratic Republic of the Congo, Ghana, Guinea, Ivory Coast, Madagascar, Mali, Togo.
Democratic Republic Of The Congo ● 3♂♂; South Kivu, CRSN Lwiro, Kabindi, Guest House, Site 3;
Democratic Republic of the Congo, Kenya.
Ghana –
Inferior appendage short and rounded apically; phallobase with ventral apex short and only weakly projecting, not enlarged apically; phallic spines both relatively elongate and narrow, differing in length; periphallic processes fused mesally, comparatively narrow and weakly developed; posteroventral margin of sternum VIII distinctly projecting.
Within the
Within the group of taxa assigned to the
Cameroon, Democratic Republic of the Congo, Ghana, Madagascar, Sao Tomé.
Phallobase with ventral apex produced and strongly bent, extreme apex enlarged and bent downward; both phallic spines narrow and elongate; inferior appendage with apex strongly narrowed and only weakly bent, cusps of ventromesal margin not evident in lateral view.
Although similar in overall morphology to other species in the
Ghana –
Ventral process of segment IX incredibly elongate; tergum X without spine-like dorsal projection, ventral part completely divided laterally, but not strongly deflexed; inferior appendage mounted at approximately midheight on segment, far above ventral process, dorsal apex strongly narrowed, acute apically, and also very strongly posteriorly bent, ventral cusps visible in lateral view; phallobase with ventral apex extended and sclerotized, but not bent or excessively produced; phallic spines both moderately elongate and narrow; sternum VIII with posteroventral margin weakly produced.
Tanzania –
Phallobase with ventral apex greatly produced, but only weakly bent, apex slightly enlarged and more or less rounded apically; phallic spines both relatively short; inferior appendage relatively narrow overall, with dorsal projection narrow and tapering, not or only scarcely bent, cusps of ventromesal surface not evident in lateral view.
We are somewhat unsure of our attribution of the specimen illustrated (Fig.
Democratic Republic of the Congo, Tanzania.
Ghana –
Ghana, Togo.
General features of subgroup: tergum X with lateral lobes entire (or sometimes cleft apically), with digitate process near dorsal margin bearing two sensilla, process sometimes short. Inferior appendage with distinct basal inflection and mesal curvature, variable in length, but generally relatively narrow, with variably modified apex; mesal cusps absent. Ventral process of segment IX short, usually posteriorly projecting; the shape of the ventral process and genital capsule is variable among species and often usefully diagnostic. Phallus often with a pair of symmetrically positioned spines.
The number of known species assigned to this subgroup is large. Although there is considerable variation in the shape of the genital capsule, ventral process, phallic armature, etc., most species have a general similarity that readily allows them to be placed in the subgroup. Distinguishing differences between some species are relatively minor. A revision of the subgroup would be a useful contribution.
Ghana –
Characters, in combination, that confirm the identification and can be used to distinguish
The form illustrated here (Fig.
Diagnostic features of
Ghana, Ivory Coast.
Tanzania –
The most diagnostic aspects of
Republic of South Africa, Tanzania.
Ghana –
Inferior appendage relatively short, uniform in width in lateral view, apex subacute in ventral view; ventral apex of phallobase expanded, laterally compressed, and ventrally deflexed; phallus with pair of small, curved, symmetrically placed spines; apex of lateral lobe of tergum X subacute.
Tanzania –
Tanzania.
Ghana –
Characters, in combination, that confirm the identification and can be used to distinguish
Ghana, Ivory Coast.
Ghana –
Characters, in combination, that confirm the identification and can be used to distinguish
Both
Ghana, Ivory Coast.
Characters tentatively used to define the group include an elongate, narrow tergum X, with preanal appendages flattened and fused basally, sensilla of tergum X on a rounded mesally directed process, nearly linear arrangement of the
Ghana –
Three species are assigned to this new group, the two from Tanzania very evidently closely related and the third from Ghana more speculatively associated. All of the species have scabrous lobes associated to terga IX or X, and relatively short phalli with a prominent sclerotized ventral apex. The endotheca is relatively simple, short and untextured (without small spines). A single short spine is present in the endotheca, as well as a short phallotremal sclerite complex, composed of a short rod and ring structure. The ventral process of segment IX is also very short. Superficially, the species look very much like species in the
Ghana –
Superficial similarities would suggest a relationship to species in the
The
Ghana –
Ghana, Guinea, Mali, Sierra-Leone.
Ghana –
Ghana, Guinea, Ivory Coast, Sierra-Leone.
Ghana –
Benin, Burkino Faso, Cameroon, Ghana, Guinea, Ivory Coast, Mali, Togo.
Ghana –
Cameroon, Ghana, Guinea, Ivory Coast, Mali, Togo.
This subgroup is probably closely related to the
The differences characterizing the
Ghana –
Because of having multiple sensilla on tergum X, rather than just two, as is typical of species in the
This group was designated by
The two subgroups of the
This subgroup has the R1 vein of the hind wing either completely obsolete, or present basally, but obsolete or fused to the subcosta apically. Notably, only one vein intersects the wing margin. The venation of the hind wing is otherwise complete for
To date, only a single species of this subgroup has been reported from Asia (
Ghana–
Ghana –
Ghana –
Ghana –
Ghana –
Ghana –
Ghana –
This subgroup is distinguished from the
Ghana –
This species is probably most closely related to
We take pleasure in naming this species
Ghana –
This species is undoubtedly closely related to
Ghana –
Ghana –
Ghana –
General diagnostic characters of the species include the general shape of segment IX, which is relatively elongate and lacks anterodorsal apodemes, the ventrally projecting ventral process of the same segment, the simple lateral lobes of tergum X, and the short curved inferior appendages. Additionally, the numerous small spines in the phallus, its relative length, absence of a projecting ventral apex on the phallobase, as well as the relatively desclerotized posteromesal margin of segment VIII are also all useful diagnostic characters, not found in any other species of the
We are pleased to name this species for Jostein Kjærandsen, who participated in the collecting expedition that generated much of the material that the current study is based on, in addition to doing an initial sorting of the material and initiating the study.
The project in Tanzania was a joint project between the Faculty of Forestry, Sokoine University of Agriculture (SUA) and the University Museum of Bergen. The second author is indebted to Jørn Bjørndalen, Kjell Arne Johanson, Geir E.E. Søli and the other members of the team for help and company during the field work. Financial support was given by the Norwegian Research Council (NFR), the Norwegian Agency for Development Cooperation (Norad) and the University of Bergen. The project in Ghana was a joint project between the Department of Zoology, University of Ghana; Institute of Aquatic Entomology, CSIR, Accra, Ghana, and the University Museum of Bergen. We are indebted Joseph S. Amakye, Institute of Aquatic Entomology, and the other members of the team. Financial support was granted by the Norwegian Universities’ Committee for Research and Education (NUFU) and by the Norwegian Research Council (NFR). Special thanks are extended to Jostein Kjærandsen who collected much of the material and did much of the initial sorting.
RB acknowledges support by the University of Minnesota through its Agricultural Experimental Station project AES-017-094, and also extends much gratitude to Ralph Holzenthal, without whose continued support, in terms of space, equipment, and materials, this research and much previous work would never have been possible. TA is also much indebted to Ralph Holzenthal for his hospitality and support during sabbaticals at the Department of Entomology, University of Minnesota.