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The rock-dwelling cichlids from Lake Malaŵi, known as mbuna, comprise a diverse group of haplochromine fishes that are placed among 14 genera. Within the mbuna, Pseudotropheus is a polyphyletic genus, which has served as a catch-all for many of these fishes. Recently, many of the species-groups within Pseudotropheus have been elevated to separate genera. Herein, we describe an elongate form that was originally placed in the Pseudotropheus elongatus species group but is now described as a member of Metriaclima.

Relationship of L. chilingali to other taxa in the Lake Malawi radiation The present study has assumed that L. lethrinus is both the most likely sister taxon for L. chilingali and the species most likely to interbreed with it, should habitat barriers be broken down. The former proposition is based on their overall similar appearance, including very similar male breeding dress, and similar – although distinct- melanin patterns in the females and juveniles. They are the only two known Lethrinops species to share a largely horizontally-banded melanin pattern. Other Lake Malawi cichlids also share some of these features, notably species of Protomelas Eccles & Trewavas 1989 found in similar shallow weedy/muddy habitats, including Protomelas kirkii (Günther 1894), Protomelas similis (Regan 1922) and Protomelas labridens (Trewavas 1935) (Eccles & Trewavas 1989, Konings 2016, Turner 1996). These three species also have females/immatures with a sandy/countershaded appearance, with a strong horizontal dark band running along the flank. Males are also metallic blue-green, with a red and white dorsal fin margin. These species have shorter snouts and more upwardly-angled mouths than L. lethrinus, but so does L. chilingali, which is arguably morphologically intermediate between them. The genera Protomelas and Lethrinops can be distinguished by the shape of the lower jaw dental arcade, and it is presently assumed that this is a phylogenetically informative trait (Eccles & Trewavas 1989), although this requires confirmation from a phylogenetic analysis, ideally based on genome-scale sequence data. A published phylogenomic analysis places L. lethrinus in the middle of a clade of shallow water Lethrinops, Taeniolethrinops and Tramitichromis (Masonick et al. 2022), thus grouping these genera showing Lethrinops - type dentition (Eccles & Trewavas 1989). However, P. kirkii, P. similis and P. labridens were not included in that analysis (Masonick et al. 2022). Notably, however, an additional group of deep-water Lethrinops appears in a separate part of the phylogenetic tree, suggesting that the Lethrinops - type dentition is prone to parallelism. Thus, we conclude that available evidence does not conflict with L. chilingali being a sister species to L. lethrinus, but this requires detailed phylogenetic investigation for confirmation. If L. lethrinus shows relatively high levels of population structure, it could be paraphyletic (ancestral) with respect to L. chilingali. Distributions of L. chilingali and L. lethrinus Lethrinops chilingali has only been positively recorded from Lake Chilingali, but here we consider whether it may have a broader distribution in Lake Malawi, possibly extending to the central to northern part of the lake as an allopatric sister species to L. lethrinus. Although a lake-wide distribution has been claimed for L. lethrinus (Konings 2016), the great majority of records backed by preserved specimens or photographs come from the southern arms, Lake Malombe and the Shire River (Eccles & Lewis 1978, Turner 1996, Konings 2016). On the Global Biodiversity Information Facility website (GBIF 2023), there is a record of Lethrinops lethrinus from co-ordinates indicating a collection site off the Tanzanian shore near Ngkuyo Island, Mbamba Bay (11.334°S, 34.769°E), based on specimens at the South African Institute for Aquatic Biodiversity (SAIAB). An offshore location near a rocky headland seems an unlikely collecting site for Lethrinops lethrinus, which favours shallow sheltered vegetated habitats and the locality label is given as ‘Lifuwu’, which probably corresponds to the vicinity of Lifuwu village (13.69°S, 34.60°E) just north of Salima, suggesting that the co-ordinates have been recorded in error. The single small specimen shows no melanic markings (faded post-preservation?), but the head shape is consistent with Lethrinops lethrinus rather than L. chilingali. Another GBIF record from co-ordinates 13.35°S, 33.4°E would suggest specimens were collected from the Rusa River, a tributary of the Bua River, which joins Lake Malawi near Lake Chilingali. The site is far upstream, around 97km West of the Lake Malawi shore at Benga, and initially we thought this might suggest specimens of L. chilingali could be widespread in the river catchment. However, the collection label indicates the specimens were collected from Lake Malawi at Foo, which is a trawling station in the SE Arm of Lake Malawi (also sometimes written as Fowo), which is at approximately 14.14°S, 35.18°E, again suggesting an error in the co-ordinates. Photographs of the specimens show typical Lethrinops lethrinus, with long snouts and strong horizontal melanic markings. The catalogue of the Natural History Museum in London contains a single accession of three specimens labelled as L. lethrinus from Lupembe Sand Bar, collected by Cuthbert Christy in 1925 (BMNH 1935.6.14.2077-9; Figure 4). The electronic catalogue suggests that this location is in Tanzania, perhaps following Eccles & Trewavas (1989) who suggested it might represent a site at the mouth of the ‘Kivira River’. However, the town at the mouth of the Kiwira River (as presently named) is currently known as Itungi Port. It is more likely that the 1925 collection site is Lupembe on the Malawian lakeshore, just south of Karonga (10.055°S, 33.99°E). Notably, recent satellite images show a conspicuous sandbar (Google Earth). Examination of the unpublished diary of Cuthbert Christy held at the Natural History Museum shows a single accession from Lupembe following an extensive collection of several hundred accessions from Vua / Deep Bay (Chilumba area) and immediately before another extensive collection from Mwaya in Tanzania, on the far north coast of the lake (itemising various river mouths visited). No other accessions were made from Lupembe. This suggests that the location was visited en-route from Chilumba to Tanzania, which would fit well with the location near Karonga. Unfortunately, the specimens (fig. 5a) are very small (44.8–50.9 mm SL) which makes morphological comparisons difficult with the larger specimens examined for this study, due to allometric effects. For example, they have notably relatively large eyes, making snout measurements relatively small. However, the low position of the mouth on the head and the largely continuous midlateral stripe, fit far better with L. lethrinus than with L. chilingali. Thus, available museum specimens strongly support the occurrence of typical Lethrinops lethrinus only in the southern arms of the lake, but tentatively indicate that they may also occur just north of Senga Bay and indeed almost at the northernmost extremity of the lake, but do not provide evidence for the occurrence of L. chilingali or any other similar form within Lake Malawi, Other published records are not backed by specimens available to examine or photographic evidence. Eccles & Lewis (1978) reported that they had found L. lethrinus at Nkhata Bay, which is well to the north of Lake Chilingali. However, they reported an unusual melanin pattern: “the dark line along the middle of the flank curves upwards anteriorly to merge with the lower of the two rows of spots and the spots themselves may run together posteriorly to form a stripe”. The occurrence of specimens with dramatically different stripe patterns at Nkhata Bay might lend credence to the idea that L. lethrinus represents a complex of allopatric taxa, which might increase the probability that L. chilingali might persist in the main Lake Malawi. Eccles & Lewis provided no illustration of this ‘ Nkhata Bay variant’. Their specimens were deposited in the collection of the Monkey Bay Fisheries Research Unit, Malawi and their present status is unknown. The pattern described is reminiscent of that seen on some of the type specimens of L. leptodon Regan 1922 (fig. 5b). In the same 1978 paper, Eccles & Lewis redescribed that species based on a single specimen collected from Chintheche in the NW of the lake, near Nkhata Bay, but their illustration of that specimen showed a clear midlateral blotch on the upper part of the flank. They reported examining, but not measuring, three of the type specimens of L. leptodon, which are held at the Natural History Museum in London (BMNH 1921.9.6.201- 207, collected by Wood from somewhere in ‘Lake Nyasa’). Thus, it seems unclear whether the reported Nkhata Bay populations represent L. lethrinus or L. leptodon, or indeed something else. In summary, the status of the northern populations of Lethrinops of this group is unclear but is consistent with the hypothesis that L. lethrinus is found in suitable habitats throughout Lake Malawi, and that L. chilingali is a satellite lake endemic extinct in the wild. Conservation status of Lethrinops chilingali Lake Chilingali is approximately 5km in length and a maximum of 1km in width, and is characterised by two deeper basins of approximately 5m depth separated by a shallower plateau (Turner et al. 2019). It has a single outflow, the Kaombe River, which meanders for approximately 12km before reaching the main body of Lake Malawi (Genner et al. 2007). The lake is a natural water body, and the two basins of the modern lake are represented on early European exploration maps, as two separate bodies of water, Lake Chikukutu to the south, and Lake Chilingali to the north (Turner et al. 2019). The lake level was raised when a dam was constructed across the single outflow for irrigation purposes, initially in the 1950s, before being modified in the early 1970s (Denys et al. 2013). The dam collapsed early in 2012 (Denys et al. 2013), and the single lake disappeared, reforming the two separate smaller shallow basins. In 2016 these basins were estimated to be only ~ 1m deep and fringed with macrophytes. The lake was refilled to approximately its pre-collapse-level in June-July 2019 following the construction of a new dam. During the period 2004 to 2011, before the collapse of the dam, L. chilingali was periodically and reliably sampled from the lake, alongside another apparently endemic haplochromine cichlid, the undescribed Rhamphochromis sp. “chilingali” (Genner et al. 2007; Turner et al. 2019). To our knowledge, the last sampling event where L. chilingali was recorded in the wild was on 25 June 2009 (by G. Turner), while representatives of R. sp. “chilingali” were last collected from an artisanal fishing catch on 12 January 2011 (by M. Genner). During sampling in February 2016, neither of the species was encountered in a survey of the main northern and southern basins of Lake Chilingali (Turner et al. 2019). A survey in April 2022 also failed to sample any either L. chilingali or R. sp “chilingali” but did find that Lake Malawi endemic Otopharynx tetrastigma (Günther 1894) was abundant (H. Svardal, pers comm). This species was absent between 2004 and 2016 and is likely to have been introduced during restocking after the lake was refilled in 2019 (H. Svardal, pers comm). Although further surveys of Lake Chilingali and the Kaombe river are warranted to determine if remnant populations of either L. chilingali or R. sp “chilingali” persist, on the basis of the current evidence, we consider it most likely that both species are no longer present in the natural environment. Breeding populations of L. chilingali or R. sp “chilingali” are, however, maintained in captivity, and may be candidates for reintroduction. On the basis of the evidence discussed above, we recommend that L. chilingali is attributed the status of Extinct in the Wild (EW) on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species.

Turner, George F., Crampton, Denise A., Genner, Martin J. (2023): A new species of Lethrinops (Cichliformes: Cichlidae) from a Lake Malawi satellite lake, believed to be extinct in the wild. Zootaxa 5318 (4): 515-530, DOI: 10.11646/zootaxa.5318.4.5, URL: http://dx.doi.org/10.11646/zootaxa.5318.4.5

Lethrinops lethrinus (Günther, 1893) Holotype: Lethrinops lethrinus (Günther, 1893): BMNH 1893.11.15.15, 116.1 mm SL, coll. A. Whyte, Upper Shire River at Fort Johnston (Mangochi), March 1892

Crenicichla jupiaensis Britski & Luengo, 1968 ARGENTINA • 2 ex., 87.7–93.0 mm SL; Corrientes, Paraná River at Yahapé; MLP 11294., Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on page 63, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480

Crenicichla iguassuensis Haseman, 1911 BRAZIL • holotype, 137.0 mm SL; Iguaçu River, Porto Uniăo da Victoria; FMNH 54159., Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on page 63, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480

Crenicichla ama sp. nov. urn:lsid:zoobank.org:act: 9C8E18CF-8833-4AAA-9395-5B2D205F1997 Figs 1A, 2, 4A, 5–8; Table 1 Crenicichla sp. Piray Guazú – Piálek et al. 2012 (molecular phylogeny); 2019a (molecular phylogeny and genetic delimitation, photo of live specimen); 2019b (molecular phylogeny and genetic delimitation). — Říčan et al. 2021a (molecular phylogeny and genetic delimitation); 2021b (molecular phylogeny, genetic and morphological delimitation, photo of live specimens). Diagnosis Crenicichla ama sp. nov. is diagnosed from the here described sympatric C. aravera sp. nov. (and from C. yjhui, C. vittata and C. mandelburgeri) in the main colouration pattern, chiefly composed of midlateral blotches along the body side associated with variably developed vertical bars vs composed chiefly of a dominant continuous midlateral band. Crenicichla ama sp. nov. is distinguished from all other species of the C. mandelburgeri species complex, i.e., those with the main body colouration chiefly composed of midlateral blotches along the body side, by having small, black spots below the midlateral blotches on occasional scales in the series 0–E3 (i.e., generally between the lower lateral line and ventral border of the midlateral blotches) vs absence of such small, black spots on the flank in C. aravera sp. nov., C. hu, C. mandelburgeri, C. taikyra, C. yaha, C. yjhui, and C. ypo vs similar small dark spots however all over the flank in the Iguazú species C. iguassuensis, C. tesay, C. tuca and males of C. tapii (only on posterior part of body and caudal peduncle) vs much larger dark spots and blotches all over the flank, evidently originating from the fragmentation of the flank bars in C. gillmorlisi. Crenicichla ama is further distinguished from C. gillmorlisi in having the dots and blotches below the midlateral blotches small and sharply defined (as in the Iguazú species) vs much larger with softer borders, and in always lacking any traces of a continuous midlateral band (vs present in most juveniles and as a trace in many adults). Crenicichla ama sp. nov. is further distinguished from all these species (except C. aravera sp. nov., C. mandelburgeri, and C. gillmorlisi) by the colouration of the dorsal fin in breeding females which features a distinct dark white-margined blotch in the dorsal fin in C. ama (vs a long dark white-margined band in the rest). Crenicichla ama sp. nov. is additionally distinguished from the Iguazú species C. tapii and C. tuca by having a complete series of midlateral blotches (vs only the anterior one or two blotches), and from C. tapii, C. tuca, C. tesay, C. yaha and C. taikyra by a different head and mouth morphology (lower jaw prognathous with normal lips vs lower jaw hypognathous with thick lips in C. tuca) and by much less robust LPJ and LPJ teeth. Crenicichla ama is distinguished from the Iguazú species C. iguassuensis and C. tesay, and from C. gillmorlisi, all with the same complete series of midlateral blotches, by having the spots on body usually limited only to around the ventral borders of the midlateral blotches (vs found throughout the body including the dorsum and belly in same intensity; also in C. tuca). Crenicichla ama sp. nov. is most similar to C. ypo and C. hu who share the same main colouration pattern and the generalistic predatory ecomorphology (intermediate in oral jaw, LPJ jaws and teeth, and head and body proportions between all five ecomorphs of Crenicichla) vs C. aravera sp. nov., C. yjhui, C. iguassuensis and C. mandelburgeri which are specialized piscivorous species. Crenicichla ama is thus readily distinguished from C. aravera in lower jaw length (mean 15.7%, SD 0.63 vs 16.8% of SL, SD 0.71), upper jaw length (mean 11.7%, SD 0.38 vs 12.8% of SL, SD 0.86), head depth (mean 16.9%, SD 0.70 in C. ama vs 15.8% of SL, SD 1.39), more robust LPJ and LPJ teeth, body depth (mean 23.1%, SD 1.10 vs 19.8% of SL, SD 1.09, caudal peduncle depth (mean 12.8% of SL, SD 0.37 vs 11.8% of SL, SD 0.38), and pectoral fin length (mean 22.9% of SL, SD 0.87 vs 20.6% of SL, SD 0.86). In morphometric characters Crenicichla ama sp. nov. is distinguished from the two most similar species C. ypo and C. hu by tendency towards longer lower jaw (C. ama 15.7% in SL, SD 0.63 vs C. ypo 14.5%, SD 2.0, vs C. hu 14.3%, SD 4.55), longer pectoral fin (C. ama 22.8% in SL, SD 1.03 vs C. ypo 20.1%, SD 0.84, vs C. hu 19.5%, SD 5.84), and larger orbit (C. ama 7.7% in SL, SD 0.47 vs C. ypo 6.4%, SD 0.37 vs C. hu 6.4%, SD 1.93). Etymology The specific epithet ‘ ama ’ is a Guaraní word for ‘rain’ and is used as a noun in apposition. The name is given in allusion to the small diagnostic spots below midlateral blotches which appear as if rain (spots) is falling from clouds (midlateral blotches). Type material Holotype ARGENTINA • ♂, 105.8 mm; Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin, upper arroyo Piray Guazú on RP20; 26°26′34.1″ S, 54°08′29.4″ W; 18 Feb. 2012; Casciotta et al. leg.; MLP 11446 (Fig. 1A). Paratypes All from Argentina, Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin. ARGENTINA – Misiones • 16 ex., 83.6–111.2 mm; same collection data as for holotype; MLP 11182 (Fig. 2) • 11 ex., 67.3–103.9 mm; same collection data as for holotype; 30 Nov. 2007; Říčan et al. leg.; MLP 11180 (Fig. 2) • 2 ex., 75.6–84.0 mm; same collection data as for holotype; MLP 11447 • 8 ex., 77.5–150.8 mm; Paraná River Basin, Piray Guazú River Basin, tributary to upper arroyo Piray Guazú on RP16, 23 km from San Pedro; 26°35′45.90″ S, 54°16′59.96″ W; 26 Nov. 2016; Říčan leg.; MLP 11448 (Fig. 2). Description Body elongate, depth 4.0 to 4.8 times in SL (mean body depth 23.1% of SL, SD 1.10) (Figs 1A, 2). Head as deep as wide or slightly deeper (mean head depth16.9% of SL, SD 0.70). Snout bluntly pointed in lateral view, 2.8 to 3.1 times in head length (mean snout length 11.4% of SL, SD 0.46). Lower jaw slightly prognathous. Tip of maxilla reaching anterior margin of orbit in most specimens. Lower lip folds widely interrupted medially. Nostrils dorsolateral. Posterior margin of preopercle serrated (27 ex.) or smooth (3 ex.). Scales on flank strongly ctenoid. Head scales cycloid. Predorsal scales small. Interopercle naked. Cheek scaled, 7 to 9 scales below eye embedded in skin. Scales in E1 row 46 (2), 47 (3), 48 (4), 49 (4), 50 (3), 51 (5), 52 (5*), 53 (2), 55 (2). Scales in transverse row 9/11 (2), 9/12 (4*), 9/13 (2), 10/11 (1), 10/12 (13), 10/13 (7), 10/14 (1). Two to three scale rows between lateral lines. Upper lateral line scales 21 (1), 22 (4), 23 (10*), 24 (9), 25 (4), 26 (2). Lower lateral line scales 8 (2), 9 (8), 10 (16), 11 (3*), 13 (1). Dorsal, anal, pectoral and pelvic fins naked. Dorsal fin XXI,9 (1); XXI,11 (12); XXI,12 (4); XXII, 9 (1); XXII,10 (1); XXII,11 (9*); XXII,12 (1); XXIII,11 (1). Anal fin III,8 (11); III,9 (18*); III,10 (1). Pectoral fin 15 (7), 16 (16*), 17 (6). Caudal-fin squamation not reaching the middle of fin. Soft-dorsal fin rounded or pointed, extending beyond caudal-fin base. Tip of anal fin not reaching caudal-fin base. Caudal fin rounded. Pectoral fin rounded, not reaching tip of pelvic fin. Microbranchiospines present on second through fourth gill arches. Gill rakers externally on first gill arch: 3 on epibranchial, 1 on angle, and 8 on ceratobranchial. Two to five patches of unicuspid teeth on fourth ceratobranchial. Lower pharyngeal tooth plate with unicuspid recurved and bicuspid crenulated curved teeth, those of posterior and medial row larger than remaining ones (Fig. 4). Upper pharyngeal tooth plate with unicuspid and bicuspid teeth. Frayed zone bearing one concavity with small unicuspid teeth. Premaxillary ascending process longer than dentigerous process. Premaxilla with unicuspid teeth on outer row, larger than inner ones. Five teeth rows near symphysis. Dentary with unicuspid teeth on outer row. Premaxillary and dentary outer row teeth slightly movable, inner ones fully depressible. Colour in life Background colour of body grey to greenish-grey, darker on dorsum, lighter on venter. Grey preorbital stripe between anterior margin of orbit and snout tip. Postorbital stripe between posterior margin of orbit and preopercle or opercle distal margin grey. Flanks with 6 to 8 irregular black blotches between the upper and lower lateral lines, with fainter double-bar extensions above the upper lateral line up to the dorsal-fin base. Distinct black sharply defined spots below the midlateral blotches are diagnostic for the species. Suborbital stripe black, very narrow and pointed, not reaching ventral margin of cheek. Dorsal, anal, and caudal fins pale grey, males with numerous small dark scattered dots on dorsal, anal, and caudal fins, which are absent or rarely seen in females (Fig. 2). Dorsal fin of breeding females with a distinctive large black blotch margined with a white ring (Fig. 2). Reproductive females additionally with an orange band from pectoral-fin cleft to mid-body below the black midlateral markings. Caudal fin with a black subcircular spot, rarely bearing partial irregular pale ring, just above of midline of caudal fin. Colour in alcohol Similar to that of live specimens apart from lack of carotenoid pigments, most importantly concerning the orange area on flank of females. Main diagnostic characters, i.e., dark spots below midlateral blotches, and shape and configuration of suborbital stripe well visible (Fig. 1A) and as in live animals (Fig. 2). Distribution Crenicichla ama sp. nov. is endemic to the arroyo Piray Guazú Basin, Paraná River Basin, Misiones Province, Argentina (Fig. 5). Habitat Crenicichla ama sp. nov. is found throughout the Piray Guazú River Basin above the rapids separating it from the Paraná River (Fig. 5). The arroyo Piray Guazú is a moderately fast flowing river. It is a clear-water basin with a predominantly bouldery stream bed and running shallow water interspersed with pools (Figs 6–7). In the pools the bottom includes more silt accumulation and the pools bear macrophytes such as Echinodorus uruguayensis Arechav. and Potamogeton pseudopolygonus Hgstr.

Keys to the species of Crenicichla from the Middle Rio Paraná Basin (the Crenicichla mandelburgeri species group plus C. vittata) Because of the taxonomical and morphological complexity of the C. mandelburgeri species complex we have chosen to provide two determination keys, each primarily based on a different main character. We believe this may facilitate determination of some species. The first key is more suited to the present study as it is based on the main colouration patterns, which also distinguish the two newly described species. The second key is more generally suited to the C. mandelburgeri species complex and the existing literature on parallel evolution of ecomorphologies in Crenicichla since it is based on oral jaw characteristics (the main charaters involved in the formation and parallel evolution of ecomorphs). For the keys we have chosen to use only readily visible external characters. Many of the characters used in the key will work only for adult specimens (at least 70 mm SL), especially the colour and colouration pattern characters. The main colouration pattern of the species is scored based on preserved specimens where there is less variation than in live specimens, while the colouration of the dorsal fin in females is best visible in live specimens. Key to the species of Crenicichla from the Middle Rio Paraná Basin (the Crenicichla mandelburgeri species group plus C. vittata) based on the main colouration pattern 1. Main colouration pattern of adults chiefly or most often composed of a dominant continuous midlateral band, in the latter case sometimes in combination with vertical bars............................. 2 – Main colouration pattern of adults chiefly composed of midlateral blotches along the body side without any continuous midlateral band........................................................................................... 5 – Main colouration pattern chiefly composed of paired vertical bars in young, and fragmented into dark blotches and spots all over the sides including side of belly in adults, midlateral band when present is continuous, endemic to the Acaray tributary, endemic to Paraguay............................................................................................................................... C. gillmorlisi Kullander & Lucena, 2013 2. The midlateral band is always present as a dominant colouration pattern in all growth stages, moods and under other conditions, scales in E1 row 50–64........................................................................ 3 – Depending on growth stage, mood and other factors the dominant body colouration can change from a midlateral band to vertical bars, or be a combination of the two, scales in E1 row either 42–57 or 78–85................................................................................................................................................ 4 3. Breeding females with a distinct dark white-margined blotch in the dorsal fin, scales in E1 row 50–55, endemic to the Piray Guazú tributary, endemic to Argentina................... C. aravera sp. nov. – Breeding females with a distinct dark white-margined long band in the dorsal fin, scales in E1 row 53–64, endemic to the Urugua-í tributary, endemic to Argentina................................................................................................................................... C. yjhui Piálek, Casciotta, Almirón & Říčan, 2019 4. Midlateral band when present always fully continuous, with barely discernible fluctuation in width, scales in E1 row 78–85, widespread in the Lower Paraná and entire Paraguay Basin, also present in Lower Uruguay and lower portions of the Middle Paraná in Paraguay, Brazil, Argentina and Uruguay......................................................................................................... C. vittata Heckel, 1840 – Midlateral band often semicontinuous or even distrupted (i.e., always of clearly fluctuating width), vertical bars when present always as clear double-bars, scales in E1 row 42–57, endemic to Middle Paraná main River and some non-isolated tributaries in Paraguay and Argentina................................................................................................................................... C. mandelburgeri Kullander, 2009 5. Breeding females with a distinct dark white-margined blotch in the dorsal fin, scattered spots on body below the midlateral band in both sexes, especially in males, endemic to the Piray Guazú tributary, endemic to Argentina................................................................................. C. ama sp. nov. – Breeding females with a distinct dark white-margined long band in the dorsal fin......................... 6 6. Scattered spots on body below the midlateral band in both sexes, especially in males, endemic to the Iguazú, Argentina and Brazil............................................................................................................ 7 – Without spots on body below the midlateral band in both sexes, endemic to Middle Paraná Basin apart from the Iguazú...................................................................................................................... 10 7. Oral jaws prognathous (i.e., lower jaw projecting in front of upper jaw), mouth large, corner of mouth reaching well below eye....................................................... C. iguassuensis Haseman, 1911 – Oral jaws isognathous or hypognathous (i.e., upper jaw projecting in front of lower jaw)............. 8 8. Mouth large, corner of mouth reaching below vertical from anterior margin of eye, lips very thick and bulbous.......................................... C. tuca Piálek, Dragová, Casciotta, Almirón & Říčan, 2015 – Mouth small, not reaching eye, lips normal..................................................................................... 9 9. Well-developed vertical double-bars (only species in the Iguazú with this character), anterior one or two midlateral blotches much more dominant over the remaining, which are usually only visible as connections between the double bars, a well-formed (not decomposed into spots) narrow suborbital stripe inclined significantly posteriad, near absence of dark scattered dots on flanks in males and complete absence in females............... C. tapii Piálek, Dragová, Casciotta, Almirón & Říčan, 2015 – Vertical bars and less developed double bars only visible above midlateral line, anterior four or more midlateral blotches well dominant, suborbital stripe decomposed into spots, wider, and less inclined posteriad, both sexes with dark scattered dots on flanks........... C. tesay Casciotta & Almirón, 2008 10. Oral jaws prognathous (i.e., lower jaw projecting in front of upper jaw), mouth rather large (i.e., corner of mouth reaching below vertical from anterior margin of eye)..........................................11 – Oral jaws isognathous or hypognathous (i.e., upper jaw projecting in front of lower jaw), mouth rather small (i.e., not reaching eyes)............................................................................................... 12 11. Anterior two to three midlateral blotches dominant, background colouration light grey, midlateral blotches distinctly H-like with dorsally continuing double bars, females with a wide wide orange to red stripe along the entire length of the basal portion of the dorsal fin above which is found a shorter black band margined in white, suborbital stripe of normal length and width, composed of spots, which do not form horizontal lines, endemic to the Urugua-í tributary, endemic to Argentina........................................................................... C. ypo Casciotta, Almirón, Piálek, Gómez & Říčan, 2010 – All midlateral blotches well developed, background colouration very dark grey to blackish, midlateral blotches square-like with dorsally continuing single wide bars, females only with the shorter black band margined in white in the dorsal fin without the orange to red band, suborbital stripe short and wide, ventrally widest, composed of spots which do form horizontal lines, endemic to the Piray Miní tributary, endemic to Argentina.............................. C. hu Piálek, Říčan, Casciotta & Almirón, 2010 12. Anterior two to three midlateral blotches usually more dominant, midlateral blotches distinctly H-like with dorsally continuing double bars, suborbital stripe shorter and wider, composed of spots which tend to form horizontal lines, scales in E1 row 48–51 (mean 50), endemic to the Urugua-í tributary, endemic to Argentina.................................... C. yaha Casciotta, Almirón & Gómez, 2006 – All midlateral blotches equally developed, midlateral blotches square-like with dorsally continuing single wide bars, suborbital stripe of normal length and width, composed of spots which tend to form horizontal lines, scales in E1 row 54–60 (mean 56–57), endemic to the lower portion of the Middle Paraná and some non-isolated tributaries in Paraguay and Argentina............................................................................................. C. taikyra Casciotta, Almirón, Aichino, Gómez, Piálek & Říčan, 2013 Key to the species of Crenicichla from the Middle Rio Paraná Basin (the Crenicichla mandelburgeri species group plus C. vittata) based on oral jaw characteristics 1. Oral jaws isognathous or hypognathous (i.e., upper jaw projecting in front of lower jaw), mouth generally rather small to small (i.e., not reaching eyes), except in one species where quite large... 2 – Oral jaws prognathous (i.e., lower jaw projecting in front of upper jaw), mouth rather large to large (i.e., corner of mouth reaching below vertical from anterior margin of eye)................................... 6 2. Mouth large, corner of mouth reaching below vertical from anterior margin of eye, lips very thick and bulbous.......................................... C. tuca Piálek, Dragová, Casciotta, Almirón & Říčan, 2015 – Mouth small, not reaching eye, lips normal..................................................................................... 3 3. Scattered spots on body below the midlateral band in both sexes, especially in males (except for one species where weakly developed), endemic to the Iguazú, Argentina and Brazil............................ 4 – Without spots on body below the midlateral band in both sexes, endemic to Middle Paraná Basin apart from the Iguazú........................................................................................................................ 5 4. Well-developed vertical double-bars (only species in the Iguazú with this character), anterior one or two midlateral blotches much more dominant over the remaining, which are usually only visible as connections between the double bars, a well-formed (not decomposed into spots) narrow suborbital stripe inclined significantly posteriad, near absence of dark scattered dots on flanks in males and complete absence in females............... C. tapii Piálek, Dragová, Casciotta, Almirón & Říčan, 2015 – Vertical bars and less developed double bars only visible above midlateral line, anterior four or more midlateral blotches well dominant, suborbital stripe decomposed into spots, wider, and less inclined posteriad, both sexes with dark scattered dots on flanks........... C. tesay Casciotta & Almirón, 2008 5. Anterior two to three midlateral blotches usually more dominant, midlateral blotches distinctly H-like with dorsally continuing double bars, suborbital stripe shorter and wider, composed of spots which tend to form horizontal lines, scales in E1 row 48–51 (mean 50), endemic to the Urugua-í tributary, endemic to Argentina.................................... C. yaha Casciotta, Almirón & Gómez, 2006 – All midlateral blotches equally developed, midlateral blotches square-like with dorsally continuing single wide bars, suborbital stripe of normal length and width, composed of spots which tend to form horizontal lines, scales in E1 row 54–60 (mean 56–57), endemic to the lower portion of the Middle Paraná and some non-isolated tributaries in Paraguay and Argentina............................................................................................. C. taikyra Casciotta, Almirón, Aichino, Gómez, Piálek & Říčan, 2013 6. Main colouration pattern of adults chiefly or most often composed of a dominant continuous midlateral band, in the latter case sometimes in combination with vertical bars............................. 7 – Main colouration pattern of adults chiefly composed of midlateral blotches along the body side without any continuous midlateral band..........................................................................................11 – Main colouration pattern chiefly composed of paired vertical bars in young, and fragmented into dark blotches and spots all over the sides including side of belly in adults, midlateral band when present is continuous, endemic to the Acaray tributary, endemic to Paraguay............................................................................................................................... C. gillmorlisi Kullander & Lucena, 2013 7. The midlateral band is always present as a dominant colouration pattern in all growth stages, moods and under other conditions, scales in E1 row 50–64........................................................................ 8 – Depending on growth stage, mood and other factors the dominant body colouration can change from a midlateral band to vertical bars, or be a combination of the two, scales in E1 row either 42–57 or 78–85................................................................................................................................................ 9 8. Breeding females with a distinct dark white-margined blotch in the dorsal fin, scales in E1 row 50–55, endemic to the Piray Guazú tributary, endemic to Argentina................... C. aravera sp. nov. – Breeding females with a distinct dark white-margined long band in the dorsal fin, scales in E1 row 53–64, endemic to the Urugua-í tributary, endemic to Argentina................................................................................................................................... C. yjhui Piálek, Casciotta, Almirón & Říčan, 2019 9. Midlateral band when present always fully continuous, with barely discernible fluctuation in width, scales in E1 row 78–85, widespread in the Lower Paraná and entire Paraguay Basin, also present in Lower Uruguay and lower portions of the Middle Paraná in Paraguay, Brazil, Argentina and Uruguay......................................................................................................... C. vittata Heckel, 1840 – Midlateral band most often semicontinuous or even distrupted (i.e., always of clearly fluctuating width), vertical bars when present always as clear double-bars, scales in E1 row 42–57, endemic to Middle Paraná main River and some non-isolated tributaries in Paraguay and Argentina...................................................................................................................... C. mandelburgeri Kullander, 2009 10. Breeding females with a distinct dark white-margined blotch in the dorsal fin, a few scattered spots on body below the midlateral band in both sexes, especially in males, endemic to the Piray Guazú tributary, endemic to Argentina................................................................................. C. ama sp. nov. – Breeding females with a distinct dark white-margined long band in the dorsal fin........................11 11. Scattered spots on body below the midlateral band in both sexes, especially in males, endemic to the Iguazú, Argentina and Brazil........................................................... C. iguassuensis Haseman, 1911 – Without spots on body below the midlateral band in both sexes, endemic to Middle Paraná Basin apart from the Iguazú, endemic to Argentina................................................................................. 12 12. Anterior two to three midlateral blotches dominant, background colouration light grey, midlateral blotches distinctly H-like with dorsally continuing double bars, females with a wide wide orange to red stripe along the entire length of the basal portion of the dorsal fin above which is found a shorter black band margined in white, suborbital stripe of normal length and width, composed of spots, which do not form horizontal lines, endemic to the Urugua-í tributary, endemic to Argentina........................................................................... C. ypo Casciotta, Almirón, Piálek, Gómez & Říčan, 2010 – All midlateral blotches well developed, backgroundcolouration very dark grey to blackish, midlateral blotches square-like with dorsally continuing single wide bars, females only with the shorter black band margined in white in the dorsal fin without the orange to red band, suborbital stripe short and wide, ventrally widest, composed of spots which do form horizontal lines, endemic to the Piray Miní tributary, endemic to Argentina.............................. C. hu Piálek, Říčan, Casciotta & Almirón, 2010, Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on pages 53-57, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480, {"references":["Kullander S. O. & Lucena C. A. S. 2013. Crenicichla gillmforlisi, a new species of cichlid fish (Teleostei: Cichlidae) from the Parana river drainage in Paraguay. Zootaxa 3641 (2): 149 - 164. https: // doi. org / 10.11646 / zootaxa. 3641.2.3","Pialek L., Casciotta J., Almiron A. & Rican O. 2019 b. A new pelagic predatory pike cichlid (Teleostei: Cichlidae: Crenicichla) from the C. mandelburgeri species complex with parallel and reticulate evolution. Hydrobiologia 832: 377 - 395. https: // doi. org / 10.1007 / s 10750 - 018 - 3754 - 1","Kullander S. O. 2009. Crenicichla mandelburgeri, a new species of cichlid fish (Teleostei: Cichlidae) from the Parana river drainage in Paraguay. Zootaxa 2006 (1): 41 - 50. https: // doi. org / 10.11646 / zootaxa. 2006.1.3","Haseman J. D. 1911. An annotated catalog of the cichlid fishes collected by the expedition of the Carnegie Museum to central South America, 1907 - 10. Annals of the Carnegie Museum 7: 329 - 373. Available from https: // www. biodiversitylibrary. org / page / 9956994 [accessed 2 Jun. 2023].","Pialek L., Dragova K., Casciotta J., Almiron A. & Rican O. 2015. Description of two new species of Crenicichla (Teleostei: Cichlidae) from the Lower Iguazu River with a taxonomic reappraisal of C. iguassuensis, C. tesay and C. yaha. Historia Natural (Tercera Serie) 5: 5 - 27.","Casciotta J. & Almiron A. 2008. Crenicichla tesay, a new species of cichlid (Perciformes: Labroidei) from the rio Iguazu basin in Argentina. Revue suisse de Zoologie 115: 651 - 659. https: // doi. org / 10.5962 / bhl. part. 80451 Casciotta J. R. & Arratia G. 1993. Jaws and teeth of American Cichlids (Pisces: Labroidei). Journal of Morphology 217: 1 - 36. https: // doi. org / 10.1002 / jmor. 1052170102","Casciotta J., Almiron A., Pialek L., Gomez S. & Rican O. 2010. Crenicichla ypo (Teleostei: Cichlidae), a new species from the middle Parana basin in Misiones, Argentina. Neotropical Ichthyology 8: 643 - 648. https: // doi. org / 10.1590 / S 1679 - 62252010000300009","Casciotta J., Almiron A. E. & Gomez S. E. 2006. Crenicichla yaha sp. n. (Perciformes: Labroidei: Cichlidae), a new species from the rio Iguazu and arroyo Urugua-i basins, northeastern Argentina. Zoologische Abhandlungen 56: 107 - 112.","Casciotta J., Almiron A., Aichino D., Gomez S., Pialek L. & Rican O. 2013. Crenicichla taikyra (Teleostei: Cichlidae), a new species of pike cichlid from the middle rio Parana, Argentina. Zootaxa 3721 (4): 379 - 386. https: // doi. org / 10.11646 / zootaxa. 3721.4.5"]}

Crenicichla lepidota (Heckel, 1840) ARGENTINA • 1 ex., 151.6 mm SL; Corrientes, Paraná River Basin, Isla Apipé Grande, Ituzaingó; MACN-ict 7275 • 1 ex., 98.4 mm SL; Formosa, Paraguay River Basin, Riacho de Oro; MACN-ict 4091 • 2 ex., 116.0– 165.7 mm SL; Misiones, Represa Estación Experimental Cerro Azul; MACN-ict 3656., Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on page 63, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480

Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana, Casciotta, Jorge (2023): Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex. European Journal of Taxonomy 879: 38-63, DOI: https://doi.org/10.5852/ejt.2023.879.2159, URL: http://dx.doi.org/10.5852/ejt.2023.879.2159

Crenicichla jupiaensis Britski & Luengo, 1968 ARGENTINA • 2 ex., 87.7–93.0 mm SL; Corrientes, Paraná River at Yahapé; MLP 11294.

Crenicichla aravera sp. nov. urn:lsid:zoobank.org:act: 1B66C894-3DE8-4439-8431-7F6D9EE41790 Figs 1B, 3, 4B, 5–8; Table 2 Crenicichla sp. Piray Guazú line – Piálek et al. 2012 (molecular phylogeny); 2019a (molecular phylogeny and genetic delimitation, photo of live specimen); 2019b (molecular phylogeny and genetic delimitation). — Říčan et al. 2021a (molecular phylogeny and genetic delimitation); 2021b (molecular phylogeny, genetic and morphological delimitation, photo of live specimens). Diagnosis Crenicichla aravera sp. nov. is diagnosed by a rare main colouration pattern (shared only with C. yjhui and to some extent with C. mandelburgeri within the C. mandelburgeri species complex, plus with C. vittata) which features a dominant continuous midlateral band vs the most common main colouration pattern composed of midlateral blotches along the body, present in all other species of the complex including C. ama sp. nov. Crenicichla aravera sp. nov. is readily distinguished from C. vittata by 50–55 scales in E1 row (vs 78– 85) and also by colouration, where the midlateral band in C. vittata (and in C. mandelburgeri) is not always present, e.g., in breeding or stressed individuals (vs always present and the dominant colouration pattern in C. aravera). Crenicichla aravera sp. nov. can also be distinguished from the most similar C. yjhui in having less scales in E1 row (50–55 vs 53–64) and in transverse row (9/12–12/10 vs 10/13–11/16). Crenicichla aravera is further distinguished from C. yjhui in one colouration-pattern character in adult breeding females, which have a distinct dark white-margined blotch in the dorsal fin (shared with C. ama sp. nov. vs a long dark white-margined band in C. yjhui). Crenicichla aravera can further be marginally distinguished from C. yjhui in the better degree of development of all of the piscivory-associated characters, i.e., in statistically longer snout (12.1% of SL, SD 1.04 vs 11.4%, SD 0.43), longer head (33.1%, SD 0.90 vs 31.9%, SD 0.78), longer lower (16.8%, SD 0.71 vs 15.9%, SD 0.50) and upper jaws (12.8%, SD 0.86 vs 12.5%, SD 0.61), longer caudal peduncle (15.4%, SD 0.69 vs 13.6%, SD 0.76), and also in several other characters, i.e., in statistically deeper head (15.8%, SD 1.39 vs 15.1%, SD 0.64), longer pectoral fin (20.6%, SD 0.85 vs 19.5%, SD 0.98), larger eye (7.2%, SD 0.49 vs 6.8%, SD 0.47). Crenicichla aravera sp. nov. is further diagnosed by its specialized piscivorous ecomorphology characterized by long oral jaws (mean lower jaw length 16.8% of SL, SD 0.71; mean upper jaw length 12.8% of SL, SD 0.86), shallow head (mean head depth 15.8% of SL, SD 1.39), delicate LPJs and LPJ teeth, streamlined bodies (mean body depth 19.8% of SL, SD 1.09) which it shares with C. yjhui, C. iguassuensis, C. mandelburgeri and with C. tuca (in oral jaws and head length) within the C. mandelburgeri species complex plus with C. vittata vs species with other ecomorphologies (C. ama sp. nov., C. gillmorlisi, C. ypo, C. hu, C. taikyra, C. tesay, C. yaha, C. tapii and C. tuca [in head depth and body depth]) with shorter oral jaws (mean lower jaw length 11.0–15.7% of SL; mean upper jaw length 9.1–11.7% of SL), deeper heads (mean head depth 16.0–18.2% of SL), more robust LPJs and LPJ teeth, and less streamlined and deeper bodies (mean body depth 22.0–24.9% of SL). Crenicichla aravera is further diagnosed from C. ama by caudal peduncle shape (mean depth 11.8% of SL, SD 0.38 vs 12.8% of SL, SD 0.37) and by pectoral fin length (mean 20.6% of SL, SD 0.86 vs 22.9% of SL, SD 0.87). Etymology The specific epithet ‘ aravera ’ is a Guaraní word for ‘flash’ (as associated with stormy and rainy weather) and is used as a noun in apposition. The name is given in association with its sister-species C. ama sp. nov. (meaning rain) in allusion to the long body, pointed head, and a dominant colouration marking of a black midlateral band and rapid hunting strategy, all alluding to a flash. Type material Holotype ARGENTINA • ♂, 102.5 mm; Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin, upper arroyo Piray Guazú on RP20; 26°26′34.1″ S, 54°08′29.4″ W; 30 Nov. 2007; Říčan et al. leg.; MLP 11450 (Figs 1B, 3). Paratypes All from Argentina, Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin, same locality as holotype. ARGENTINA – Misiones • 2 ex., 81.2–83.3 mm; same collection data as for holotype; MLP 11451 (Fig. 3) • 4 ex., 2 (C&S), 90.1–135.3 mm; Paraná River Basin, arroyo Piray Guazú Basin, upper arroyo Piray Guazú on RP20; 26°26′34.1″ S, 54°08′29.4″ W; 18 Feb. 2012; Casciotta et al. leg.; MLP 11452 (Fig. 3). Non-type material ARGENTINA • 12 ex., 2 (C&S), 79.9–101 mm; Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin, lower arroyo Piray Guazú at Balneario Piray Guazú; 26°29′1.66″ S, 54°35′11.98″ W; 23 Feb. 2012; Casciotta et al. leg.; MLP 11453. Description Body elongate, depth 4.6 to 5.3 times in SL (mean body depth 19.8% of SL, SD 1.09) (Figs 1B, 3). Head as deep as wide or slightly deeper (mean head depth 15.8% of SL, SD 1.39). Snout bluntly pointed in lateral view, 2.5 to 3.0 times in head length (mean snout length 12.1% of SL, SD 1.04). Lower jaw slightly prognathous. Tip of maxilla reaching anterior margin of orbit in specimens over 80.0 mm SL. Lower lip folds widely separated along symphysis. Nostrils dorsolateral. Posterior margin of preopercle serrated. Scales on flank strongly ctenoid. Head scales cycloid. Predorsal and prepelvic scales small. Interopercle naked. Cheek scaled, 8 to 11 scales below eye embedded in skin. Scales in E1 row 50 (1), 51 (1), 52 (1), 53 (1), 55 (3). Scales in transverse row 9/12 (2), 9/13 (1), 10/13 (1), 10/14 (1*), 12/10 (1). Two to three scale rows between lateral lines. Upper lateral line scales 21 (1), 23 (1), 25 (3), 26 (2*). Lower lateral line scales 8 (1), 10 (3*), 11 (2), 12 (1). Dorsal, anal, pectoral and pelvic fins naked. Dorsal fin XX,12 (1); XXI,11 (2); XXI,12 (2*); XXII,11 (1); XXIII,11 (1). Anal fin III,8 (4); III,9 (3*). Pectoral fin 16 (4); 17 (3*). Caudal-fin squamation not reaching the middle of fin. Soft-dorsal fin rounded or pointed, surpassing caudal-fin base. Tip of anal fin reaching or not caudal-fin base. Caudal fin rounded. Pectoral fin rounded, not reaching tip of pelvic fin. Microbranchiospines present on second through fourth gill arches. Gill rakers externally on first gill arch: 3 on epibranchial, 1 on angle, and 8 on ceratobranchial. Three to five patches of unicuspid teeth on fourth ceratobranchial. Lower pharyngeal tooth plate with unicuspid recurved and bicuspid crenulated curved anteriorly teeth, those of posterior and medial row larger than remaining ones (Fig. 4). Upper pharyngeal tooth plate with unicuspid and bicuspid teeth. Frayed zone bearing one concavity with small unicuspid teeth. Premaxillary ascending process longer than dentigerous one. Premaxilla with unicuspid teeth on outer row, larger than inner ones. Five teeth rows near symphysis. Dentary with unicuspid teeth on outer row. Premaxillary and dentary outer row teeth slightly movable, inner ones fully depressible. Colour in life Background colour of body grey to yellowish-grey, darker on dorsum, lighter on venter, where almost white. Deep grey preorbital stripe between anterior margin of orbit and snout tip. Postorbital stripe between posterior margin of orbit and preopercle or opercle distal margin deep grey to black. Deep grey to black midlateral band along entire body as dominant colouration patter and main diagnostic character on body. Light grey hints of vertical bars only visible on dorsum above midlateral band. Suborbital stripe black, rather wide, not distinctly pointed, and short, not reaching ventral margin of cheek. Dorsal, anal, and caudal fins pale grey, males with numerous rather large dark scattered dots on dorsal, anal, and caudal fins, which are absent or rarely seen in females (Fig. 3). Dorsal fin of breeding females with distinctive large black blotch (Fig. 3). Females additionally with more yellow tones below midlateral band, in reproductive females likely more intensified into an orange band from pectoral-fin cleft to mid-body. Caudal fin with a black subcircular spot, rarely bearing a partial irregular pale ring, just above of midline of caudal fin. Colour in alcohol Similar to that of live specimens apart from lack of carotenoid pigments, most importantly of orange area on flank of females. Main diagnostic characters, i.e., black continuous midlateral band with spotshaped irregularities on ventral margin, and shape and configuration of suborbital stripe well preserved (Fig. 1B) and as in live animals (Fig. 3). Distribution Crenicichla aravera sp. nov. is endemic to the arroyo Piray Guazú Basin, Paraná River Basin, in Misiones Province, Argentina (Fig. 5). Habitat Crenicichla aravera sp. nov. is only found in the Piray Guazú Basin above the rapids separating it from the Paraná River, but has so far only been collected from the main river channel (Fig. 5), unlike C. ama sp. nov., which is also known from tributaries. Crenicichla aravera is syntopic with C. ama. Other details concerning the arroyo Piray Gauzú as given for C. ama., Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on pages 47-53, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480, {"references":["Pialek L., Rican O., Casciotta J., Almiron A. & Zrzavy J. 2012. Multilocus phylogeny of Crenicichla (Teleostei: Cichlidae), with biogeography of the C. lacustris group: species flocks as a model for sympatric speciation in rivers. Molecular Phylogenetics and Evolution 62: 46 - 61. https: // doi. org / 10.1016 / j. ympev. 2011.09.006","Rican O., Dragova K., Almiron A., Casciotta J., Gottwald J. & Pialek L. 2021 a. MtDNA species-level phylogeny and delimitation support significantly underestimated diversity and endemism in the largest Neotropical cichlid genus (Cichlidae: Crenicichla). PeerJ 9: e 12283. https: // doi. org / 10.7717 / peerj. 12283","Casciotta J., Almiron A., Pialek L., Gomez S. & Rican O. 2010. Crenicichla ypo (Teleostei: Cichlidae), a new species from the middle Parana basin in Misiones, Argentina. Neotropical Ichthyology 8: 643 - 648. https: // doi. org / 10.1590 / S 1679 - 62252010000300009"]}

Crenicichla ama sp. nov. urn:lsid:zoobank.org:act: 9C8E18CF-8833-4AAA-9395-5B2D205F1997 Figs 1A, 2, 4A, 5–8; Table 1 Crenicichla sp. Piray Guazú – Piálek et al. 2012 (molecular phylogeny); 2019a (molecular phylogeny and genetic delimitation, photo of live specimen); 2019b (molecular phylogeny and genetic delimitation). — Říčan et al. 2021a (molecular phylogeny and genetic delimitation); 2021b (molecular phylogeny, genetic and morphological delimitation, photo of live specimens). Diagnosis Crenicichla ama sp. nov. is diagnosed from the here described sympatric C. aravera sp. nov. (and from C. yjhui, C. vittata and C. mandelburgeri) in the main colouration pattern, chiefly composed of midlateral blotches along the body side associated with variably developed vertical bars vs composed chiefly of a dominant continuous midlateral band. Crenicichla ama sp. nov. is distinguished from all other species of the C. mandelburgeri species complex, i.e., those with the main body colouration chiefly composed of midlateral blotches along the body side, by having small, black spots below the midlateral blotches on occasional scales in the series 0–E3 (i.e., generally between the lower lateral line and ventral border of the midlateral blotches) vs absence of such small, black spots on the flank in C. aravera sp. nov., C. hu, C. mandelburgeri, C. taikyra, C. yaha, C. yjhui, and C. ypo vs similar small dark spots however all over the flank in the Iguazú species C. iguassuensis, C. tesay, C. tuca and males of C. tapii (only on posterior part of body and caudal peduncle) vs much larger dark spots and blotches all over the flank, evidently originating from the fragmentation of the flank bars in C. gillmorlisi. Crenicichla ama is further distinguished from C. gillmorlisi in having the dots and blotches below the midlateral blotches small and sharply defined (as in the Iguazú species) vs much larger with softer borders, and in always lacking any traces of a continuous midlateral band (vs present in most juveniles and as a trace in many adults). Crenicichla ama sp. nov. is further distinguished from all these species (except C. aravera sp. nov., C. mandelburgeri, and C. gillmorlisi) by the colouration of the dorsal fin in breeding females which features a distinct dark white-margined blotch in the dorsal fin in C. ama (vs a long dark white-margined band in the rest). Crenicichla ama sp. nov. is additionally distinguished from the Iguazú species C. tapii and C. tuca by having a complete series of midlateral blotches (vs only the anterior one or two blotches), and from C. tapii, C. tuca, C. tesay, C. yaha and C. taikyra by a different head and mouth morphology (lower jaw prognathous with normal lips vs lower jaw hypognathous with thick lips in C. tuca) and by much less robust LPJ and LPJ teeth. Crenicichla ama is distinguished from the Iguazú species C. iguassuensis and C. tesay, and from C. gillmorlisi, all with the same complete series of midlateral blotches, by having the spots on body usually limited only to around the ventral borders of the midlateral blotches (vs found throughout the body including the dorsum and belly in same intensity; also in C. tuca). Crenicichla ama sp. nov. is most similar to C. ypo and C. hu who share the same main colouration pattern and the generalistic predatory ecomorphology (intermediate in oral jaw, LPJ jaws and teeth, and head and body proportions between all five ecomorphs of Crenicichla) vs C. aravera sp. nov., C. yjhui, C. iguassuensis and C. mandelburgeri which are specialized piscivorous species. Crenicichla ama is thus readily distinguished from C. aravera in lower jaw length (mean 15.7%, SD 0.63 vs 16.8% of SL, SD 0.71), upper jaw length (mean 11.7%, SD 0.38 vs 12.8% of SL, SD 0.86), head depth (mean 16.9%, SD 0.70 in C. ama vs 15.8% of SL, SD 1.39), more robust LPJ and LPJ teeth, body depth (mean 23.1%, SD 1.10 vs 19.8% of SL, SD 1.09, caudal peduncle depth (mean 12.8% of SL, SD 0.37 vs 11.8% of SL, SD 0.38), and pectoral fin length (mean 22.9% of SL, SD 0.87 vs 20.6% of SL, SD 0.86). In morphometric characters Crenicichla ama sp. nov. is distinguished from the two most similar species C. ypo and C. hu by tendency towards longer lower jaw (C. ama 15.7% in SL, SD 0.63 vs C. ypo 14.5%, SD 2.0, vs C. hu 14.3%, SD 4.55), longer pectoral fin (C. ama 22.8% in SL, SD 1.03 vs C. ypo 20.1%, SD 0.84, vs C. hu 19.5%, SD 5.84), and larger orbit (C. ama 7.7% in SL, SD 0.47 vs C. ypo 6.4%, SD 0.37 vs C. hu 6.4%, SD 1.93). Etymology The specific epithet ‘ ama ’ is a Guaraní word for ‘rain’ and is used as a noun in apposition. The name is given in allusion to the small diagnostic spots below midlateral blotches which appear as if rain (spots) is falling from clouds (midlateral blotches). Type material Holotype ARGENTINA • ♂, 105.8 mm; Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin, upper arroyo Piray Guazú on RP20; 26°26′34.1″ S, 54°08′29.4″ W; 18 Feb. 2012; Casciotta et al. leg.; MLP 11446 (Fig. 1A). Paratypes All from Argentina, Misiones Province, Paraná River Basin, arroyo Piray Guazú Basin. ARGENTINA – Misiones • 16 ex., 83.6–111.2 mm; same collection data as for holotype; MLP 11182 (Fig. 2) • 11 ex., 67.3–103.9 mm; same collection data as for holotype; 30 Nov. 2007; Říčan et al. leg.; MLP 11180 (Fig. 2) • 2 ex., 75.6–84.0 mm; same collection data as for holotype; MLP 11447 • 8 ex., 77.5–150.8 mm; Paraná River Basin, Piray Guazú River Basin, tributary to upper arroyo Piray Guazú on RP16, 23 km from San Pedro; 26°35′45.90″ S, 54°16′59.96″ W; 26 Nov. 2016; Říčan leg.; MLP 11448 (Fig. 2). Description Body elongate, depth 4.0 to 4.8 times in SL (mean body depth 23.1% of SL, SD 1.10) (Figs 1A, 2). Head as deep as wide or slightly deeper (mean head depth16.9% of SL, SD 0.70). Snout bluntly pointed in lateral view, 2.8 to 3.1 times in head length (mean snout length 11.4% of SL, SD 0.46). Lower jaw slightly prognathous. Tip of maxilla reaching anterior margin of orbit in most specimens. Lower lip folds widely interrupted medially. Nostrils dorsolateral. Posterior margin of preopercle serrated (27 ex.) or smooth (3 ex.). Scales on flank strongly ctenoid. Head scales cycloid. Predorsal scales small. Interopercle naked. Cheek scaled, 7 to 9 scales below eye embedded in skin. Scales in E1 row 46 (2), 47 (3), 48 (4), 49 (4), 50 (3), 51 (5), 52 (5*), 53 (2), 55 (2). Scales in transverse row 9/11 (2), 9/12 (4*), 9/13 (2), 10/11 (1), 10/12 (13), 10/13 (7), 10/14 (1). Two to three scale rows between lateral lines. Upper lateral line scales 21 (1), 22 (4), 23 (10*), 24 (9), 25 (4), 26 (2). Lower lateral line scales 8 (2), 9 (8), 10 (16), 11 (3*), 13 (1). Dorsal, anal, pectoral and pelvic fins naked. Dorsal fin XXI,9 (1); XXI,11 (12); XXI,12 (4); XXII, 9 (1); XXII,10 (1); XXII,11 (9*); XXII,12 (1); XXIII,11 (1). Anal fin III,8 (11); III,9 (18*); III,10 (1). Pectoral fin 15 (7), 16 (16*), 17 (6). Caudal-fin squamation not reaching the middle of fin. Soft-dorsal fin rounded or pointed, extending beyond caudal-fin base. Tip of anal fin not reaching caudal-fin base. Caudal fin rounded. Pectoral fin rounded, not reaching tip of pelvic fin. Microbranchiospines present on second through fourth gill arches. Gill rakers externally on first gill arch: 3 on epibranchial, 1 on angle, and 8 on ceratobranchial. Two to five patches of unicuspid teeth on fourth ceratobranchial. Lower pharyngeal tooth plate with unicuspid recurved and bicuspid crenulated curved teeth, those of posterior and medial row larger than remaining ones (Fig. 4). Upper pharyngeal tooth plate with unicuspid and bicuspid teeth. Frayed zone bearing one concavity with small unicuspid teeth. Premaxillary ascending process longer than dentigerous process. Premaxilla with unicuspid teeth on outer row, larger than inner ones. Five teeth rows near symphysis. Dentary with unicuspid teeth on outer row. Premaxillary and dentary outer row teeth slightly movable, inner ones fully depressible. Colour in life Background colour of body grey to greenish-grey, darker on dorsum, lighter on venter. Grey preorbital stripe between anterior margin of orbit and snout tip. Postorbital stripe between posterior margin of orbit and preopercle or opercle distal margin grey. Flanks with 6 to 8 irregular black blotches between the upper and lower lateral lines, with fainter double-bar extensions above the upper lateral line up to the dorsal-fin base. Distinct black sharply defined spots below the midlateral blotches are diagnostic for the species. Suborbital stripe black, very narrow and pointed, not reaching ventral margin of cheek. Dorsal, anal, and caudal fins pale grey, males with numerous small dark scattered dots on dorsal, anal, and caudal fins, which are absent or rarely seen in females (Fig. 2). Dorsal fin of breeding females with a distinctive large black blotch margined with a white ring (Fig. 2). Reproductive females additionally with an orange band from pectoral-fin cleft to mid-body below the black midlateral markings. Caudal fin with a black subcircular spot, rarely bearing partial irregular pale ring, just above of midline of caudal fin. Colour in alcohol Similar to that of live specimens apart from lack of carotenoid pigments, most importantly concerning the orange area on flank of females. Main diagnostic characters, i.e., dark spots below midlateral blotches, and shape and configuration of suborbital stripe well visible (Fig. 1A) and as in live animals (Fig. 2). Distribution Crenicichla ama sp. nov. is endemic to the arroyo Piray Guazú Basin, Paraná River Basin, Misiones Province, Argentina (Fig. 5). Habitat Crenicichla ama sp. nov. is found throughout the Piray Guazú River Basin above the rapids separating it from the Paraná River (Fig. 5). The arroyo Piray Guazú is a moderately fast flowing river. It is a clear-water basin with a predominantly bouldery stream bed and running shallow water interspersed with pools (Figs 6–7). In the pools the bottom includes more silt accumulation and the pools bear macrophytes such as Echinodorus uruguayensis Arechav. and Potamogeton pseudopolygonus Hgstr., Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on pages 42-47, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480, {"references":["Pialek L., Rican O., Casciotta J., Almiron A. & Zrzavy J. 2012. Multilocus phylogeny of Crenicichla (Teleostei: Cichlidae), with biogeography of the C. lacustris group: species flocks as a model for sympatric speciation in rivers. Molecular Phylogenetics and Evolution 62: 46 - 61. https: // doi. org / 10.1016 / j. ympev. 2011.09.006","Rican O., Dragova K., Almiron A., Casciotta J., Gottwald J. & Pialek L. 2021 a. MtDNA species-level phylogeny and delimitation support significantly underestimated diversity and endemism in the largest Neotropical cichlid genus (Cichlidae: Crenicichla). PeerJ 9: e 12283. https: // doi. org / 10.7717 / peerj. 12283"]}

Crenicichla ocellata (Perugia, 1897) PARAGUAY • holotype, 257.5 mm SL; Paraguay, Puerto 14 de Mayo, Bahía Negra, Chaco Boreal; MSNG 33700., Published as part of Říčan, Oldřich, Piálek, Lubomír, Almirón, Adriana & Casciotta, Jorge, 2023, Description of two new species forming a sympatric species pair of Crenicichla (Teleostei: Cichlidae) endemic to the Piray Guazú River in the Paraná River Basin, Misiones, Argentina and belonging to the C. mandelburgeri species complex, pp. 38-63 in European Journal of Taxonomy 879 on page 63, DOI: 10.5852/ejt.2023.879.2159, http://zenodo.org/record/8135480

Genus Cichla Bloch & Schneider, 1801 TYPE SPECIES. — Cichla ocellaris Bloch & Schneider, 1801 (type species by original designation)., Published as part of Carrillo-Briceño, Jorge D., Mora-Rojas, Laura, Hendricks, Kimberly, Vanegas, Andrés & Aguilera, Orangel, 2023, New clues on the palaeodiversity of the middle Miocene freshwater ichthyofauna from the Tatacoa Desert, Colombia, pp. 327-351 in Geodiversitas 45 (10) on page 334, DOI: 10.5252/geodiversitas2023v45a10, http://zenodo.org/record/8056044, {"references":["BLOCH M. E. & SCHNEIDER J. G. 1801. - M. E. Blochii [...] Systema ichthyologiae iconibus ex illustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit. Saxo, J. G. Schneider, 584 p. https: // doi. org / 10.5962 / bhl. title. 5750"]}

cf. Astronotus sp. (Fig. 2 H1-H4). MATERIAL EXAMINED. — Isolated dentary bone (VPPLT-668A) LOCALITY. — La Victoria Fm. Tatacoita (see Fig. 1 B; Appendix 1). DESCRIPTION VPPLT-668A is a short and high left dentary bone, 22 mm in length and missing the posterior part of the mandibular sensory canal and the coronoid process (Fig. 2 H1-H4). The dorsal margin is well preserved, but all teeth are missing. There are approximately six circular tooth implantation rows, with the first three from the outer border best defined (Fig. 2 H3). Enlarged bases characterize the outer tooth row, reaching up to twice the diameter of those bases of the other inner rows. The external face of the dentary has three well-developed foramina on the bony mandibular canal. The largest one is oval in shape (Fig. 2 H1) and corresponds with the foramen “f.5” referred by Casciotta & Arratia (1993). In the ventral section of the dentary, abundant small foramina can be observed. REMARKS Our comparison (based on osteological collections and literature review) suggests that VPPLT-668A is clearly different from other medium/large sizes South American cichlids such as Acaronia Myers, 1940, Aequidens Eigenmann & Bray, 1894, Caquetaia Fowler, 1945, Cichla Bloch & Schneider, 1801, Cichlasoma Swainson, 1839, Crenicichla Heckel, 1840, Geophagus Heckel, 1840, Retroculus Eigenmann & Bray, 1894, Satanoperca Günther, 1862, Symphysodon Heckel, 1840, and Uaru Heckel, 1840. The dentary shape and the pattern and size of the tooth implantation rows in VPPLT-668A resemble the jaw morphology of Astronotus (see Fig. 2 I1-I2). These are features that are “apparently” observed only in this genus; although this should be confirmed in the future with more detailed studies on cranial elements of the South American cichlids. Our osteological comparisons with specimens of Astronotus were restricted to Astronotus ocellatus (Agassiz, 1831). The genus Astronotus is represented by three extant species, Astronotus ocellatus, Astronotus crassipinnis (Heckel, 1840), and Astronotus mikoljii (Pérez-Lozano et al. 2022). The natural distribution of Astronotus includes the Amazon, Orinoco, upper Paraná, and French Guiana basins (Pérez-Lozano et al. 2022), with habitat preference of quiet shallow waters (van der Sleen & Albert 2018). Due to the state of preservation of VPPLT-668A, and the absence of more specimens with other diagnostic characters, we are unable to classify this fossil beyond cf. Astronotus sp. The presence of cf. Astronotus in the middle Miocene of the Honda Group represents the first fossil record for the genus. Neotropical cichlids, with at least 44 genera, constitute the third most speciose group of freshwater fishes in South America (van der Sleen & Albert 2018). Despite the high species richness, few studies based on comparative descriptions of cranial elements (e.g., dentaries, maxillary, and premaxillary bones) have been carried out (e.g., Casciotta & Arratia 1993). One of the problems added to the study of isolated fossil cichlid dentaries is that the morphology of dentary bone is similar among Neotropical cichlids (Casciotta & Arratia 1993). Nevertheless, morphological features can allow some tentative assignments as long as the comparisons are supported by diversity in comparative osteological material., Published as part of Carrillo-Briceño, Jorge D., Mora-Rojas, Laura, Hendricks, Kimberly, Vanegas, Andrés & Aguilera, Orangel, 2023, New clues on the palaeodiversity of the middle Miocene freshwater ichthyofauna from the Tatacoa Desert, Colombia, pp. 327-351 in Geodiversitas 45 (10) on page 334, DOI: 10.5252/geodiversitas2023v45a10, http://zenodo.org/record/8056044, {"references":["CASCIOTTA J. R. & ARRATIA G. 1993. - Jaws and teeth of American cichlids (Pisces: Labroidei). Journal of Morphology 217 (1): 1 - 36. https: // doi. org / 10.1002 / jmor. 1052170102","MYERS G. S. 1940. - Suppression of Acaropsis and Chalcinus, two preoccupied generic names of South American fresh-water fishes. Stanford Ichthyological Bulletin 1 (5): 1 - 170.","EIGENMANN C. H. & BRAY W. L. 1894. - A revision of the American Cichlidae. Annals of the New York Academy of Science 7 (1): 607 - 624. https: // doi. org / 10.1111 / j. 1749 - 6632.1893. tb 55412. x","FOWLER H. W. 1945. - Colombian zoological survey. Part I. The fresh-water fishes obtained in 1945. Proceedings of the Academy of Natural Sciences of Philadelphia 97: 93 - 135. https: // www. jstor. org / stable / 4064382","BLOCH M. E. & SCHNEIDER J. G. 1801. - M. E. Blochii [...] Systema ichthyologiae iconibus ex illustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit. Saxo, J. G. Schneider, 584 p. https: // doi. org / 10.5962 / bhl. title. 5750","SWAINSON W. 1839. - On the Natural History and Classification of Fishes, Amphibians, & Reptiles. Longman, Orme, Brown, Green & Longmans, and John Taylor, London, 448 p.","HECKEL J. J. 1840. - Johann Natterer's neue Flussfische Brasilien's nach den Beobachtungen und Mittheilungen des Entdeckers beschrieben (Erste Abtheilung, Die Labroiden). Annalen des Wiener Museums der Naturgeschichte 2: 325 - 471. https: // doi. org / 10.5962 / bhl. title. 101457","GUNTHER A. C. L. G. 1862. - Catalogue of the Fishes in the British Museum. Vol. 5. Catalogue of the Acanthopterygii, Pharyngognathi and Anacanthini in the Collection of the British Museum. Trustees, London, 534 p. https: // doi. org / 10.5962 / bhl. title. 8809","PEREZ-LOZANO A., LASSO-ALCALA O. M., BITTENCOURT P. S., TAPHORN D. C., PEREZ N. & FARIAS I. P. 2022. - A new species of Astronotus (Teleostei, Cichlidae) from the Orinoco River and Gulf of Paria basins, northern South America. ZooKeys 1113: 111 - 52. https: // doi. org / 10.3897 / zookeys. 1113.81240","VAN DER SLEEN P. & ALBERT J. S. 2018. - Field Guide to the Fishes of the Amazon, Orinoco, and Guianas. Princenton University Press, New Jersey, 464 p."]}

cf. Cichla sp. (Fig. 2 J1-J3). MATERIAL EXAMINED. — Isolated dentary bone (VPPLT-668B). LOCALITY. — La Victoria Fm. Tatacoita (see Fig. 1 B; Appendix 1). DESCRIPTION VPPLT-668B is an incomplete dentary bone of 16 mm in length, missing the posterior part of the mandibular sensory canal and the coronoid process (Fig. 2 J1-J2). The external face of the dentary preserves four well-developed foramina on the bony mandibular canal, the two smallest being located close to the dentary symphysis. All teeth are missing, and the preserved dorsal margin is characterized by six to seven circular tooth implantations rows of small size. REMARKS VPPLT-668B is easily distinguishable from VPPLT-668A assigned to cf. Astronotus. Our osteological comparisons suggest a close resemblance between VPPLT-668B and the dentaries of extant Cichla species (Fig. 2 K1-K3) than with those of Crenicichla or other medium/large sizes South American cichlids. Although VPPLT-668B is incomplete, its dentary shape and the pattern and size of the tooth implantations rows resemble the jaw morphology of Cichla species. However, due to the state of preservation of VPPLT-668B and the absence of more specimens, for now, we prefer to refer to this fossil as cf. Cichla sp. The genus Cichla, contains the largest cichlid of South America, and is represented by at least 15 species from the Amazon and Orinoco basins, as well as rivers of the Guianas (Froese & Pauly 2022). The extant species have a wide range of habitat preferences, including rivers and floodplain lakes (van der Sleen & Albert 2018). The presence of cf. Cichla in the middle Miocene of the Honda Group represents the first fossil record for the genus. From the Honda Group, Lundberg (1997) reported three premaxillaries, three dentaries, and two anguloarticular bones in fragmentary condition as indeterminate cichlids from the Fish Bed locality, suggesting a lack of morphological characters and relation to modern cichlids. Future work with more detailed anatomical comparisons on the specimens that Lundberg (1997) reported could support or contrast these assignments. Other new cichlid material reported herein includes an isolated anal spine (VPPLT-1164) coming from the Km 121 locality (Appendix 1). The spine is 18 mm in length, robust and elongated, with a sharp end at the apical section, and typical of the anal region (Fig. 2 L1-L2); nevertheless, taxonomic identification based on isolated spines is not possible., Published as part of Carrillo-Briceño, Jorge D., Mora-Rojas, Laura, Hendricks, Kimberly, Vanegas, Andrés & Aguilera, Orangel, 2023, New clues on the palaeodiversity of the middle Miocene freshwater ichthyofauna from the Tatacoa Desert, Colombia, pp. 327-351 in Geodiversitas 45 (10) on page 334, DOI: 10.5252/geodiversitas2023v45a10, http://zenodo.org/record/8056044, {"references":["FROESE R. & PAULY D. (EDS) 2022. - FishBase. World Wide Web electronic publication. https: // www. fishbase. org (02 / 2022)","VAN DER SLEEN P. & ALBERT J. S. 2018. - Field Guide to the Fishes of the Amazon, Orinoco, and Guianas. Princenton University Press, New Jersey, 464 p.","LUNDBERG J. G. 1997. - Fishes of the La Venta fauna: Additional taxa, biotic and paleoenvironmental implications, in KAY R. F., MADDEN R. H., CIFELLI R. L. & FLYNN J. J. (eds.), Vertebrate Paleontology in the Neotropics. The Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington and London: 67 - 91."]}

Carrillo-Briceño, Jorge D., Mora-Rojas, Laura, Hendricks, Kimberly, Vanegas, Andrés, Aguilera, Orangel (2023): New clues on the palaeodiversity of the middle Miocene freshwater ichthyofauna from the Tatacoa Desert, Colombia. Geodiversitas 45 (10): 327-351, DOI: 10.5252/geodiversitas2023v45a10

FAMILY Cichlidae Amatitlania nigrofasciata (Günther, 1867); Introduced; Convict Cichlid; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014, Labatos Jr. & Briones 2014; DD Cichla ocellaris Bloch & Schneider, 1801; Introduced; Peacock Cichlid; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; NE Coptodon zillii (Gervais, 1848); Introduced; Redbelly Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Hemichromis bimaculatus (Gill, 1862); Introduced; Jewelfish; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Mayaheros urophthalmus (Günther, 1862); Introduced; Mexican Mojarra; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis aureus (Steindachner, 1864); Introduced; Blue Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis mossambicus (Peters, 1852); Introduced; Mozambique Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis niloticus (Linnaeus, 1758); Introduced; Nile Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis spilurus (Günther, 1894); Introduced; Sabaki Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis urolepis (Norman, 1922); Introduced; Wami Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Parachromis managuensis (Günther, 1867); Introduced; Jaguar Guapote; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Aquilino et al. 2011, Guerrero 2014; LC Sarotherodon melanotheron Rüppell, 1852; Introduced; Blackchin Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Aquino et al. 2011, Guerrero 2014; LC, Published as part of Jamandre, Brian Wade, 2023, Freshwater fishes of the Philippines: a provisional checklist, pp. 151-181 in Zootaxa 5301 (2) on pages 163-164, DOI: 10.11646/zootaxa.5301.2.1, http://zenodo.org/record/8030274, {"references":["Guerrero III, R. M. (2014) Impacts of introduced freshwater fishes in the Philippines (1905 - 2013): A review and recommendations. Philippine Journal of Science, 143 (1), 49 - 62.","Aquilino, S. V. L., Tango, J. M., Fontanilla, I. K. C., Pagulayan, R. C., Basiao, Z. U., Ong, P. S. & Quilang, J. P. (2011) DNA barcoding of the ichthyofauna of Taal Lake, Philippines. Molecular Ecology Resources, 11, 612 - 619. https: // doi. org / 10.1111 / j. 1755 - 0998.2011.03000. x","Aquino, L. M. G., Tango, J. M., Canoy, R. J. C., Fontanilla, I. K. C., Basiao, Z. U., Ong, P. S. & Quilang, J. P. (2011) DNA barcoding of fishes of Laguna de Bay, Philippines. Mitochondrial DNA, 22 (4), 143 - 153. https: // doi. org / 10.3109 / 19401736.2011.624613"]}

Jamandre, Brian Wade (2023): Freshwater fishes of the Philippines: a provisional checklist. Zootaxa 5301 (2): 151-181, DOI: 10.11646/zootaxa.5301.2.1, URL: http://dx.doi.org/10.11646/zootaxa.5301.2.1

FAMILY Cichlidae Amatitlania nigrofasciata (Günther, 1867); Introduced; Convict Cichlid; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014, Labatos Jr. & Briones 2014; DD Cichla ocellaris Bloch & Schneider, 1801; Introduced; Peacock Cichlid; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; NE Coptodon zillii (Gervais, 1848); Introduced; Redbelly Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Hemichromis bimaculatus (Gill, 1862); Introduced; Jewelfish; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Mayaheros urophthalmus (Günther, 1862); Introduced; Mexican Mojarra; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis aureus (Steindachner, 1864); Introduced; Blue Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis mossambicus (Peters, 1852); Introduced; Mozambique Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis niloticus (Linnaeus, 1758); Introduced; Nile Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis spilurus (Günther, 1894); Introduced; Sabaki Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Oreochromis urolepis (Norman, 1922); Introduced; Wami Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Guerrero 2014; LC Parachromis managuensis (Günther, 1867); Introduced; Jaguar Guapote; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Aquilino et al. 2011, Guerrero 2014; LC Sarotherodon melanotheron Rüppell, 1852; Introduced; Blackchin Tilapia; Introduced to river basins of Luzon, Visayas and Mindanao Island groups; Aquino et al. 2011, Guerrero 2014; LC

Bujurquina omagua sp. nov. urn:lsid:zoobank.org:act: 8E478708-58F1-4E97-960C-9FCD93107C9A Figs 6–10; Table 1 Bujurquina sp. Oran Říčan et al. 2022. Bujurquina sp. Peru 1 Říčan et al. 2022. Diagnosis Bujurquina omagua sp. nov. is highly apomorphic compared to other described species in the genus because it is unique in several character states. These include: 1) shape and pattern of the suborbital stripe in adults, which is bend and with a blotch at the postero-ventral end (vs straight or absent, with or without a blotch), with an ontogenetic change with juveniles and subadults having a straight stripe running diagonally postero-ventrally and starting from below the eye in juveniles (i.e., not bend as in adults) and without a blotch, 2) orange cheek and lower head coloration with alternating opalescent lines and series of spots on snout and spotted on cheek and lower head, 3) spinous portion of dorsal fin with a single thin diagonal black line per membrane, or two lines per membrane in holotype and largest adults (vs unpatterned in all Southern group species, or with circular blotches, with several thick diagonal black lines per membrane, or two thick horizontal bands across the dorsal fin in all other Northern group species), 4) longest snout of all species without overlap (mean 15.5%, 14.4–16.4% of SL; followed by B. oenolaemus with a mean of 11.4%, 10.3–13.1 of SL), and 5) most delicate LPJ, expressed by being shortest of all species with widest horns (length/width ratio 0.42–0.45 vs 0.50–0.64 in all other species), one of the smallest dentigerous areas (length of dentigerous area/width of LPJ ratio 0.33–0.34 vs 0.34–0.44 except B. tambopatae, B. moriorum and B. huallagae where also 0.33–0.34), and the least robust dorso-ventrally (bone at posterior end of dentigerous area less thick or equal to length of largest posterior teeth vs much thicker than teeth in other species). Bujurquina omagua sp. nov. is additionally diagnosed by the following unique combination of morphometric characters, the most pronounced of which characterize the robust head of the species: 1) second to third longest head (after B. oenolaemus and together with B. labiosa; both with a mean 37.0%, 36.3–38.0% of SL in B. omagua vs mean 39.1%, 38.6–40.4% of SL), 2) third longest preorbital distance (after B. robusta and B. oenolaemus; mean 9.3%, 8.5–10.0% of SL), 3) large interocular distance (largest mean value of all species, 12.5% of SL vs 9.7–12.4% however with large overlap of ranges with the other species), 4) rather wide head (mean 19.9%, range 19.1–20.7% of SL), 5) rather long preorbital distance (mean 9.3%, range 8.5–10.0% of SL, third longest based on mean value), 6) robust head which has rather small eyes (mean 11.3%, range 10.3–12.0% of SL), 7) very deep body (mean 45.1%, range 43.2–46.8% of SL, based on mean the second after B. vitatta), but 8) with rather long and shallow caudal peduncle (CPL mean 13.8%, range 12.6–15.6% of SL; CPD mean 16.6%, range 15.5–17.1% of SL), and 9) rather short last spine in the dorsal fin (mean 15.6%, range 12.3–17.1% of SL). Etymology The specific epithet ‘ omagua ’ is a noun in apposition given after the Omagua people, which were at the time of first contact with Europeans in the 16 th century the dominant people along the banks of the Amazon River upstream from the mouth of the Negro River well into Peru. Type material Holotype PERU • adult ♂, 97.4 mm; Amazonas river basin, Department Loreto, Maynas Province, District Las Amazonas, close to Oran village, Quebrada Sabalillo, cabeceras upper locality P18-17; 3° 27´08.4˝ S, 72° 29´11.8˝ W; 97 m a.s.l. based on GPS, 115 m a.s.l. based on Google Earth; 28 Jun. 2018; Říčan leg.; clear-water stream with blackish tinge; ID tag 1189; MUSM 70225 (Figs 6, 8–9). Paratypes All from Peru, Amazonas river basin, Department Loreto, Maynas Province, District Las Amazonas, close to Oran village, 9 ex. PERU • 1 adult, 78.6 mm; same collection data as for holotype; ID tag 1188; GenBank: OP436199; MUSM 70224 • 1 adult, 98.9 mm; same collection data as for holotype; ID tag 1190; GenBank: OP436200; MUSM 70226 (Figs 7–9) • 1 adult, 69.7 mm; same collection data as for holotype; ID tag 1191; GenBank: OP436201; MUSM 70227 (Fig. 7) • 1 adult, 77.1 mm; same collection data as for holotype; ID tag 1192; GenBank: OP436202; MUSM 70228 (Fig. 7) • 1 adult, 78.7 mm; Amazonas river basin, Department Loreto, Maynas Province, District Las Amazonas, close to Oran village, Qebrada Sabalillo on trail from mouth, P18-15; 3° 27´45.6˝ S, 72° 29´22.5˝ W; 90 m a.s.l. based on GPS, 110 m a.s.l. based on Google Earth; 27 Jun. 2018; Říčan leg.; clear-water stream with blackish tinge; ID tag 1182; GenBank: OP436196; MUSM 70221 (Figs 7, 9) • 1 adult, 79.9 mm; Amazonas river basin, Department Loreto, Maynas Province, District Las Amazonas, close to Oran village, cabeceras of Quebrada Sabalillo, lower loc., P18-16; 3° 27´17.5˝ S, 72° 29´22.8˝ W; 75 m a.s.l. based on GPS, 117 m a.s.l. based on Google Earth; 28 Jun. 2018; Říčan leg.; clear-water stream with blackish tinge; ID tag 1185; GenBank: OP436197; MUSM 70222 (Figs 7, 10) • 1 adult, 92.6 mm; same collection data as for preceding; ID tag 1186; GenBank: OP436198; MUSM 70223 (Fig. 7) • 1 adult; Amazonas river basin, Department Loreto, Maynas Province, District Las Amazonas, close to Oran village, small unnamed quebrada just before entering Oran creek, P18-21; 3° 27´21.2˝ S, 72° 30´43.2˝ W; 82 m a.s.l. based on GPS, 104 m a.s.l. based on Google Earth; 3 Jul. 2018; Říčan leg.; clear-water stream with blackish tinge; ID tag 1200; GenBank: OP436204; MUSM 70229 (Fig. 7) • 1 juvenile; same collection data as for preceding; GenBank: OP436205; MUSM 70230. Description Compound, based on eight specimens from Sabalillo creek, 69.7–98.9 mm. Refer to Figs 6–10. Measurements summarized in Table 1. SHAPE. Moderately elongate, rather robust, deep bodied (body depth mean 45.1%, range 43.2–46.8% of SL). Head especially robust, long (head length mean 37.0%, 36.3–38.0% of SL), wide (head width mean 19.9%, range 19.1–20.7% of SL), with large interocular distance (mean 12.5%, range 11.6–14.2% of SL). Frontal outline straight, nape and snout curved, anterior half of dorsal-fin base contour curved. Ventral outline curved. Snout long (mean 15.5%, 14.4–16.4% of SL). Preorbital distance long (mean 9.3%, range 8.5–10.0% of SL), preorbital bone much longer than high. Robust head with rather small eyes (mean 11.3%, range 10.3–12.0% of SL). Lips rather narrow, not turgid. Corner of mouth not reaching vertical from anterior margin of orbit, especially in largest specimens. Rather long and shallow caudal peduncle (CPL mean 13.8%, range 12.6–15.6% of SL; CPD mean 16.6%, range 15.5–17.1% of SL), and rather short last spine in dorsal fin (mean 15.6%, range 12.3–17.1% of SL). SCALES. Squ. long. (E1) scales 24 (5), 25 (3); 2 scales between L1 and dorsal fin anteriorly, 2 scales between L1 and L2. Upper lateral line (L1) with 15 (1), 16 (7) scales. Lower lateral line (L2) with 9 (4), 10 (3), 11 (1) scales; canal bearing scales on caudal fin: 0 in dorsal, 0-1 in median and 0 in ventral sequence. Cheek scales in 3 (8) series. Anterior portion of caudal fin scaly. FINS. Dorsal fin XII.11 (1), XIII.10 (1), XIII.11 (1), XIV.9 (1), XIV.10 (4). Soft anal fin pointed, 3 rd ray longest, reaching to middle of caudal fin or slightly beyond. Anal fin III.8 (8). Pectoral fin with rounded tip, reaching to above genital papilla to anal-fin origin (to 2 nd spine in one specimen); Pectoral fin 13 (8). Pelvic fin pointed, first ray slightly produced, reaching to origin of anal fin or base of 1 st spine of anal fin. Caudal fin rounded, without well-developed streamers or with only short streamers. GILL-RAKERS. 1 epibranchial, 1 in angle, and 5 (3), 6 (4), 7 (1) ceratobranchial rakers externally on first arch. JAW TEETH. An outer series of stronger, conical, sharply pointed, slightly recurved teeth, slightly increasing in size anteriorly, not worn, 15–22 in outer hemiseries in upper jaw. LOWER PHARYNGEAL JAW AND TEETH. Lower pharyngeal jaw (LPJ) as examined in two specimens (79.8– 98.9 mm) (Fig. 10) very delicate, most delicate LPJ of all described species of Bujurquina, very shallow vertically dorso-ventrally (bone at posterior end of dentigerous area less thick than length of largest posterior teeth in the smaller specimen, almost so in the larger specimen vs thicker than the teeth in all other species). LPJ also shortest and widest of all species, length/width ratio 0.42–0.45 (vs 0.50–0.64 in all other species). LPJ tooth-plate (due to the longest arms within Bujurquina) also one of the shortest in genus, length of dentigerous area/width of LPJ ratio 0.33–0.34 (vs 0.34–0.44 except B. tambopatae, B. moriorum and B. huallagae where also 0.33–0.34), Teeth slightly laterally compressed (most pronounced in posterior row), slightly bicuspid with main cusp posterior, central and posterior teeth largest, smaller towards outside, central teeth with flat-worn apex, others with pronounced tip. Coloration in alcohol Coloration limited to melanin patterns in preserved state, ground colour light yellowish-brownish, lighter towards whitish on chest and anterior abdomen, dusky greyish on anterior head, snout, and dorsum. Dark diffuse and wide interorbital band. Operculum silvery greyish. Up to four clearly defined grey lines between eye and mouth corner and on anterioir cheek interspersed with spots, rest of cheek as well as opercular series richly spotted. Suborbital stripe rather marrow and of uniform width, diffuse and ghost-like, bend from behind eye forward to cheek and then postero-ventrally to a variously developed suborbital stripe blotch in adults (never ocellated and with diffuse borders). In one collected juvenile specimen and in subadults straight (i.e., not bend), running from below eye (vs from behind eye) diagonally postero-ventrally. Midlateral band continuous from head to penultimate vertical bar on body, strongly pigmented and black, very wide anteriorly and gradually narrowing posteriorly. No interruption in vertical bar 5 (sensu Říčan et al. 2005), midlateral blotch (usually present and dominant in bar 5 in cichlasomatine cichlids) is absent. Lateral band continued on head across nape close behind eye. Body with five vertical bars plus one caudal peduncle bar and a caudal peduncle blotch (usually vertically elongated). The five body bars correspond to ontogenetic bars 2–5 with the anteriormost ontogenetic bars 6–7 fused into one wide bar below midlateral band but visible as two above it. Posterior body bars (2–4) narrow and tightly spaced, anterior bars (5, 6+ 7) wide and widely spaced. Bars darkest dorsally, reaching ventrally approximately to level of lower edge of caudal peduncle. Bars do not extend onto dorsal fin. Body scale centres (below midlateral band) with dark blotches, more pronounced in zones of vertical bars. Dorsal fin light grey, lappets indistinct, with a single narrow grey diagonal stripe per membrane in spinous part, with blotches on ventral portion of soft part. Anal fin light grey, with spots on last about four membranes. Caudal fin light grey with minute slightly elongate dark dots on anterior half, distally membranes hyaline with grey edges. Pelvic fin light grey, edge and produced portion of first ray white. Coloration in life Melanin patterns as in preserved. Lower head, cheek, and opercular series yellow-orange, with opalescent golden to azure blue lines and spots, arranged as up to four clearly defined lines between eye and mouth corner and on anterioir cheek interspersed with spots, rest of cheek as well as opercular series richly spotted. Fins orange, pelvic fin yellow, edge and produced portion of first ray white (opalescent). Spinous portion of dorsal fin with a single thin diagonal black line per membrane in juveniles and subadults, and with two lines per membrane in holotype and large adults. Lower lip pale greyish to indistinctly light azure blue. Midlateral blotch present only as a paler reflective area within the dark midlateral band (hence absent from coloration of preserved specimens). Habitat Bujurquina omagua sp. nov. was found only in small rainforest streams within the basins of the Sabalillo and Oran creeks. The localities (Fig. 11) were around 2 m wide streams, with shallow water (less than 1 m except in deeper pools) but deep mud and accumulated debris, blackish water, and only weak or moderate current. Bujurquina omagua has not been collected from the main streams of the Oran creek or from the lower sections of the Oran and Sabalillo creeks (Fig. 12), which have muddy water and where B. syspilus was found. Distribution Bujurquina omagua sp. nov. is only known from the Sabalillo and Oran creeks which empty into the Amazon River on either side of the village of Oran located on the first high ground just downstream from the mouth of the Napo into the Amazon, District Las Amazonas, Maynas Province, Department Loreto, Perú (Figs 1, 13). Morphometric differentiation of Bujurquina omagua sp. nov. Bujurquina omagua sp. nov. has a highly apomorphic head and body shape that sets it apart from the sympatric/parapatric species. When analyzed through PCA these species are arranged along the main tentic-lotic axis as corresponding to their lowland-upland habitats (Figs 14–15). The lotic/lentic ecomorphology dichotomy is in the PCA analyses best correlated with caudal peduncle length (long vs short), caudal peduncle depth (narrow vs deep), body depth (narrow vs deep), the size of the eye (orbital diameter; small vs large), pectoral fin length (short vs long), and last dorsal fin spine length (short vs long). The remaining characters are not correlated with the main ecomorphological dichotomy (i.e., interocular distance, head width, head length, preorbital distance, snout length, and first ventral fin ray length) (Figs 14–15). The lotic/lentic ecomorphology dichotomy is best visible in the analysis including all central western Amazonian species (Figs 1 and 14), while the diagnostic characters of Bujurquina omagua sp. nov. are best visible in the analysis excluding the westernmost and most highland species B. zamorensis (Fig. 15). Bujurquina omagua is in the PCAs distinguished from the sympatric/parapatric species predominantly by characters not correlated with the lotic-lentic, namely large head, wide head, long snout, and long preorbital dostance. All these diagnostic characters are in line with the diagnosis of the species based on means and ranges of the morphometric characters (see above). Bujurquina omagua sp. nov. has an outlying LPJ shape that can be described as the most delicate LPJ found among the valid species of Bujurquina (Fig. 16). The LPJ of Bujurquina omagua (studied in two specimens, see above) as expressed by proportional measurements is the shortest and widest of all species (L/W ratio) and the LPJ tooth-plate is the shortest in the genus (Ld/W ratio). Additionaly to this the LPJ is very shallow vertically dorso-ventrally, but this character was scored as binary, in comparison to the length of teeth (see Results), and thus not included in the PCA analysis. Paradoxically, the most delicate LPJ of B. omagua sp. nov. is associated with one of the largest heads among species of Bujurquina (together with B. labiosa) and with the longest snout among species of Bujurquina. A similar head and snout shape in B. oenolaemus is however associated with the most robust LPJ (Fig. 16) of this molluscivorous species. A similar combination of a long snout and large head with a very delicate LPJ as in B. omagua sp. nov. is however also known from the cichlid genus Australoheros Říčan & Kullander, 2006 in the species A. forquilha Říčan & Kullander, 2008 and A. ykeregua Říčan, Piáleck, Almirón & Casciotta, 2011 (Říčan & Kullander 2008; Říčan et al. 2011). Phylogeny of Bujurquina and phylogenetic position of Bujurquina omagua sp. nov. Phylogenetic analyses of the 1055 characters cytb data matrix performed with MP in PAUP* (including all ingroup sequences) and BI analyses in MrBayes and BEAST (performed with the corresponding haplotypes data matrix) provided robust and very similar results. BI runs in independent analyses with the best-fitting model of evolution (GTR+I+G) converged well (ESS>200 for all parameters; burn-in 10%) and led to identical topologies under the same settings. The NJ and MP analyses were done with the complete dataset, the BI and BEAST analyses with the haplotype dataset. No phylogenetic differences were found between the MP, BI and BEAST analyses, and hence only the BEAST analysis is shown in Fig. 17. The phylogenetic reconstructions divide Bujurquina into two main clades that can be diagnosed by patterned vs unpatterned dorsal fin, corresponding in geographical terms to Northern vs Southern group (Figs 1 and 17). The main biogeographic dichotomy within the genus between the Southern and Northern groups is within the present upper Ucayali basin in Peru (Fig. 1). The two main clades of Bujurquina are dated as having diverged at 16.6 Ma. Within the Southern group the two southern-most species from Argentina, Paraguay, Brazil and Bolivia are sister species (B. vittata and B. oenolaemus), and they are a sister group to the clade composed of the Peruvian B. eurhinus and B. tambopatae as sister species, followed by the Bolivian B. sp. Bolivia and then the Peruvian B. robusta. The phylogenetically basal-most species of the Southern group is the Peruvian B. labiosa. The Peruvian species are thus in basal positions within the Southern group, and they also have the longest internodes and the earliest dates of divergence based on molecular clock dating, with the time frame of divergence among the Peruvian species dated between 12.4 Ma and 3.0 Ma (Fig. 17). The Northern group of Bujurquina is made up of two highly divergent clades (Fig. 17) unlike the situation in the Southern group. The first clade within the Northern group shows a younger basal divergence at 3.9 Ma and includes, as a basal species, the here described B. omagua sp. nov., and then, among the Peruvian species, B. huallagae and B. syspilus. Samples of the morphologically and biogeographically Southern group species B. megalospilus are found within B. syspilus together with some samples of the Southern gro

Říčan, Oldřich, Říčanová, Štěpánka (2023): A new highly apomorphic species of Bujurquina (Teleostei: Cichlidae) from a reverse flowing river in the Peruvian Amazon, with a key to the species in the genus. European Journal of Taxonomy 870: 167-201, DOI: https://doi.org/10.5852/ejt.2023.870.2127, URL: http://dx.doi.org/10.5852/ejt.2023.870.2127

To date, 41 species of Gyrodactylus have been described from Africa. However, none of these have been reported in Morocco. After identifying and examining 738 cyprinid host specimens, 26 specimens belonging to Gyrodactylus were found to parasitize the gills of nine species of Luciobarbus, Carasobarbus, and Pterocapoeta. The current study provides new information about the presence of a new parasitic species in Morocco, the first to be characterized on a species level in the Maghreb region. It describes in detail 12 specimens of Gyrodactylus isolated from the gills of Luciobarbus pallaryi (Pellegrin, 1919) and Luciobarbus ksibi (Boulenger, 1905). Based on morphoanatomical observations, the characterization of the specimens collected indicates a species of Gyrodactylus that is new to science, described here as Gyrodactylus nyingiae n. sp. The new species is different from previously described gyrodactylids infecting African cyprinid hosts because it has a longer hamulus total length, a longer hamulus root, a downward projecting toe of the marginal hook, and a trapezium-shaped ventral bar membrane with a slightly striated median portion and small rounded anterolateral processes. This study increases the total number of Gyrodactylus spp. found in African cyprinids to four. This research was funded by the Special Research Fund (BOF) of Hasselt University, grant number BOF22DOCLI04, to M.I.S., BOF20TT06 and BOF21INCENT09 to M.P.M.V and by research grant 1513419N of the Research Foundation—Flanders (FWO-Vlaanderen) The authors would like to sincerely thank Abdelaziz Benhoussa for allowing us to work in the Biodiversity, Ecology and Genome Laboratory (Mohammed V University in Rabat). Bouabid Badaoui and Hocein Bazairi are thanked for their statistical advice. Nikol Kmentová, Walter Boeger, Daniel Mungai Ndegwa, Kelly Thys, Tiziana Gobbin and Hervé Mandtoumbi are thanked for their laboratory assistance and help with statistical analysis

Based on morphological and molecular analysis of Astronotus species, a new species is described from the Orinoco River and Gulf of Paria basins in Venezuela and Colombia. Morphologically, it differs from Astronotus crassipinnis and Astronotus ocellatus in pre-orbital depth, caudal peduncle depth, head width, and caudal peduncle length, with significant differences in average percentage values. Osteologically, it differs from the two described species by lacking a hypurapophysis on the parahypural bone (hypural complex) and having two or three supraneural bones. Another characteristic that helps diagnose the new species is the morphology of the sagitta otolith, which is oval with crenulated dorsal and ventral margins and a rounded posterior edge. Genetically, the new species is distinct from all the other lineages previously proposed for the genus, delimited by five single locus species delimitation methods, and also has unique diagnostic nucleotides. Phylogenetic analyses support the monophyly of the new species as well as all other species/lineages. Astronotus species have considerable genetic, anatomical, and sagitta otolith shape differences, but have few significant traditional morphometric and meristic differences, because there is high variability in counts of spines, soft dorsal-fin rays, and lateral-line scales. It is clear that this new species is genetically and anatomically differentiated from all other species within the genus, and deserves recognition as a new valid species.

Vieja bifasciata (Steindachner) (Fig. 38, [LU]) Distribution. Tai Tam Reservoir, Tai Tam Byewash Reservoir (GBIF.org, 2021; current survey); Hok Tau Reservoir; Tai Tam Tuk Reservoir (current survey). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on pages 147-148, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Hemichromis fasciatus (Peters) Distribution. Tai Lam Chung Reservoir (AFCD, 2021c). Native range. Africa., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 144, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Vieja fenestrata (Günther) (Fig. 39, [LU]), Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 148, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Chan, Jeffery C.F., Tsang, Alphonse H.F., Yau, Sze-man, Hui, Tommy C.H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David, Liew, Jia Huan (2023): The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins. Raffles Bulletin of Zoology 71: 128-168, DOI: 10.26107/RBZ-2023-0012

Pelmatolapia mariae (Boulenger) Distribution. Tai Lam Chung Reservoir (AFCD, 2021c); Pak Ngan Heung Stream (DSD, 2021); Ting Kok area (GBIF. org, 2021; current survey). Native range. Africa., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 147, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Labidochromis caeruleus (Fryer) Distribution. Plover Cove Reservoir; Kowloon Reservoir (current survey). Native range. Africa (Lake Malawi endemic). Remarks. One of the most commonly sold ornamental cichlids in Hong Kong. However, they are rarely recorded in the wild., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Oreochromis niloticus (Linnaeus) (Fig. 35, [LU]) Distribution. Tai Chung Hau area, Lam Tsuen River, Kau Lung Hang Lo Wai area, Shuen Wan area, Sha Ling area, Tan Shan River, Pui O Stream, Tong Fuk River, Tai O River, Tai Ho River (Chan, 2001); Lung Mei area (Halcrow China Limited, 2007); Fung Lok Wai area (CH 2M HILL Hong Kong Limited, 2008); Tsang Tsui area (Metcalf & Eddy Limited, 2008); Sha Lo Wan area (Ove Arup & Partners Hong Kong Limited, 2009); Big Wave Bay Stream (DSD, 2010; current survey); Kong Nga Po area (Mott MacDonald Hong Kong Limited, 2013); Fanling area, Kwu Tung area (Ove Arup & Partners Hong Kong Limited, 2013b); Ho Pui Reservoir, Hok Tau Reservoir, Shing Mun Reservoir, Kowloon Reservoir, Plover Cove Reservoir, Pok Fu Lam Reservoir, Kowloon Reception Reservoir, Shek Lei Pui Reservoir (Lai, 2011; current survey); Sandy Ridge Cemetery (Ove Arup & Partners Hong Kong Limited, 2016); Mui Tsz Lam stream (AECOM Asia Company Limited, 2016a); Hung Shui Kiu area (AECOM Asia Company Limited, 2016b); Shan Ha Tsuen area, Muk Kiu Tau Tsuen area, Tai Tong area (Ove Arup & Partners Hong Kong Limited, 2017); Sung Shan New Village, Lin Fa Tei area, Ha Che area (Atkins China Limited, 2020); Yuen Long Town Nullah (Black & Veatch Hong Kong Limited, 2020a); Hong Po Road area (Black & Veatch Hong Kong Limited, 2020b); Nam Wa Po area (DSD, 2020); Shek Kiu Tau area (Mott MacDonald Hong Kong Limited, 2020); Pak Tin area (Tai Wai) (AECOM Asia Company Limited, 2021a); Lau Shui Heung Reservoir (AFCD, 2021c); Ma Wat River, Ping Yuen River, Pak Ngan Heung Stream, Ma Tso Lung Stream, Ngau Tam Mei River, Yuen Long Bypass Floodway, Ho Chung River, Kam Tin River, Ping Kong Stream, Cheung Po Stream, Wong Lung Hang Stream, Tai Tei Tong Stream, Shek Kong Stream, Ho Pui River, Deep Water Bay Stream (DSD, 2021); Wetland Park, Hang Tau area, Tin Shui Wai Park, Tai Po Kau Nature Reserve, Sheung Yue River, Tin Shui Wai River, She Shan River, Ting Kok area, Nam Sang Wai area, San Tin area, Kai Tak River, High Island Reservoir, Hung Shui Hang Irrigation Reservoir, Shan Pui River (GBIF.org, 2021); Tung Chung River (Green Power, 2021; current survey); Mai Po Nature Reserve (WWF, 2021); Kowloon Byewash Reservoir; Lam Tei Irrigation Reservoir, Tai Shui Hang area, Aberdeen Lower Reservoir, Aberdeen Upper Reservoir, Tai Tam Reservoir, Tai Tam Tuk Reservoir, Tai Tam Byewash Reservoir, Tai Tam Intermediate Reservoir (current survey). Native range. Africa. Remarks. The introduction of this species has not been as well documented as that of O. mossambicus. However, this species may have been misidentified as O. mossambicus in earlier studies. Presently, O. niloticus appears to be relatively abundant and has replaced O. mossambicus in numerous habitats. It differs from other Oreochromis spp. in that the caudal fin has black bars, and the upper margin of the dorsal fin is black or grey and never entirely fringed with red as in O. mossambicus, O. aureus, or hybrids (Trewavas, 1983)., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on pages 146-147, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765, {"references":["Chan BPL (2001) Sustainability and biodiversity: the impact, alternative design and prospects of restoration of channelized lowland streams in Hong Kong. Unpublished PhD Thesis. University of Hong Kong, Hong Kong, pp. 169 - 211.","CH 2 M HILL Hong Kong Limited (2008) Proposed Development at Fung Lok Wai, Yuen Long at Lot 1457 R. P. in D. D. 123. Environmental Impact Assessment Report prepared for the Environmental Protection Department, Hong Kong Government. https: // www. epd. gov. hk / eia / register / report / eiareport / eia _ 1492008 / EIA % 20 Report / html / FLW _ EIA. htm (Accessed 21 October 2021).","DSD (2010) Drainage Improvement in Big Wave Bay. Project Profile for Environmental Permit prepared for the Environmental Protection Department, Hong Kong Government, 22 pp.","Lai SYH (2011) Reservoir fishes of Hong Kong with remarks on conservation options. Memoirs of the Hong Kong Natural History Society, (27): 63 - 82.","Green Power (2021) Ecological Baseline Study of Tung Chung River Catchment. Green Power, Hong Kong, pp. 60 - 62.","Trewavas E (1983) Tilapiine fishes of the genera Sarotherodon, Oreochromis and Danakilia. Bulletin of the British Museum (Natural History), 878: 1 - 583."]}

Astronotus ocellatus (Agassiz) (Fig. 27) Distribution. Black Hill area, Kowloon Reservoir, Plover Cove Reservoir (GBIF.org, 2021); Shing Mun Reservoir; Lai King area (current survey). Native range. Central and South America. Remarks. Popular in the aquarium trade; frequently released in urban ponds. However, breeding populations have never been observed in Hong Kong., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 143, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Amphilophus labiatus (Günther) Distribution. Plover Cove Reservoir (GBIF.org, 2021; current survey). Native range. Central and South America. Remarks. Breeding populations found in Plover Cove Reservoir but not as abundant as Amphilophus citrinellus., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 143, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Maylandia lombardoi (Burgess) Distribution. Shing Mun Reservoir (current survey). Native range. Africa (Lake Malawi endemic). Remarks. As with Labidochromis caeruleus, it is one of the most commonly sold ornamental cichlids in Hong Kong, but is rarely recorded in the wild., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Amphilophus citrinellus (Günther) (Fig. 26, [LU]) Distribution. Plover Cove Reservoir (GBIF.org, 2021; current survey). Native range. Central and South America. Remarks. Common in the aquarium trade., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 143, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Heterotilapia buttikoferi (Hubrecht) (Fig. 32, [LU]) Distribution. Kowloon Byewash Reservoir (Mott MacDonald Hong Kong Limited, 2008 as Tilapia joka; current survey); Plover Cove Reservoir (Lai, 2011 as Tilapia buttikoferi; GBIF.org, 2021; current survey); Lai Chi Kok Park, Aberdeen Lower Reservoir, Tai Tam Tuk Reservoir, Shing Mun Reservoir (GBIF.org, 2021; current survey); Lok Ma Chau area, Aberdeen Upper Reservoir (current survey). Native range. Africa., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765, {"references":["CH 2 M HILL Hong Kong Limited (2008) Proposed Development at Fung Lok Wai, Yuen Long at Lot 1457 R. P. in D. D. 123. Environmental Impact Assessment Report prepared for the Environmental Protection Department, Hong Kong Government. https: // www. epd. gov. hk / eia / register / report / eiareport / eia _ 1492008 / EIA % 20 Report / html / FLW _ EIA. htm (Accessed 21 October 2021).","Lai SYH (2011) Reservoir fishes of Hong Kong with remarks on conservation options. Memoirs of the Hong Kong Natural History Society, (27): 63 - 82."]}

Vieja melanurus (Günther) (Fig. 40), Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 148, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Heros efasciatus (Heckel) Distribution. Aberdeen Lower Reservoir (GBIF.org, 2021). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Mesonauta festivus (Heckel) Distribution. Aberdeen Lower Reservoir (GBIF.org, 2021). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Coptodon zillii (Gervais) (Fig. 28, [LU]) Distribution. Shing Mun River (Chan, 2001 as Tilapia zillii); High Island Reservoir, Hok Tau Reservoir, Kowloon Reservoir, Lau Shui Heung Reservoir, Plover Cove Reservoir, Pok Fu Lam Reservoir, Kowloon Reception Reservoir, Shek Lei Pui Reservoir, Tai Lam Chung Reservoir (Lai, 2011 as Tilapia zillii); Lok Ma Chau Loop, Tse Koo Hang area (Ove Arup & Partners Hong Kong Limited, 2013a as Tilapia zillii); Sham Wat River (Mott MacDonald Hong Kong Limited, 2014 as Tilapia zillii); Sandy Ridge Cemetery (Ove Arup & Partners Hong Kong Limited, 2016 as Tilapia zillii); Lin Fa Tei area (Atkins China Limited, 2020); Nam Sang Wai area (Black & Veatch Hong Kong Limited, 2020a; current survey); Shek Pik Reservoir (AFCD, 2021c); Ma Tso Lung Stream, Ping Yuen River, Yuen Long Bypass Floodway, Ho Chung River, Ma Wat River, Lam Tsuen River, Pak Ngan Heung Stream (DSD, 2021 as Tilapia zillii); Tung Chung River (Green Power, 2021); Wetland Park, Shui Lo Cho Stream, Pui O Stream, Shek Lei Pui Reservoir, Shap Long Irrigation Reservoir, Lamma Island area, Ting Kok area, Tai Tam Byewash Reservoir, Tan Shan River, Tai Ho River, Hang Hau Man Kuk Lane Park (GBIF.org, 2021); Deep Water Bay Stream, San Tin area, Ng Tung River, Sheung Yue River, Kam Tin River, Tsing Tai Stream, Shing Mun Reservoir, Lower Shing Mun Reservoir, Lam Tei Irrigation Reservoir, Hung Shui Hang Irrigation Reservoir, Kowloon Byewash Reservoir, Tai Tam Tuk Reservoir, Tai Tam Reservoir, Tai Tam Intermediate Reservoir (current survey). Native range. Africa. Remarks. One of the most widespread cichlids in Hong Kong, thriving in both degraded lowland streams and pristine hill streams. Abundant in reservoirs where they are now found in greater numbers than Oreochromis spp., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on pages 143-144, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765, {"references":["Chan BPL (2001) Sustainability and biodiversity: the impact, alternative design and prospects of restoration of channelized lowland streams in Hong Kong. Unpublished PhD Thesis. University of Hong Kong, Hong Kong, pp. 169 - 211.","Lai SYH (2011) Reservoir fishes of Hong Kong with remarks on conservation options. Memoirs of the Hong Kong Natural History Society, (27): 63 - 82.","Green Power (2021) Ecological Baseline Study of Tung Chung River Catchment. Green Power, Hong Kong, pp. 60 - 62."]}

Melanochromis auratus (Boulenger) Distribution. Unknown. Native range. Africa (Lake Malawi endemic). Remarks. No location data was provided by AFCD (2021a)., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Herichthys cyanoguttatus (Baird & Girard) Distribution. Kai Tak River (GBIF.org, 2021). Native range. Texas, North America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 145, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Geophagus brasiliensis (Quoy & Gaimard) Distribution. Shing Mun Reservoir (GBIF.org, 2021). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 144, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Cichla temensis (Humboldt) Distribution. Shing Mun Reservoir (current survey). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 143, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Cichla monoculus (Spix & Agassiz) Distribution. Shing Mun Reservoir; Kowloon Reservoir (GBIF.org, 2021). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 143, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Parachromis managuensis (Günther) (Fig. 37, [LU]) Distribution. Plover Cove Reservoir (GBIF.org, 2021; current survey). Native range. Central and South America., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 147, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765

Hemichromis cf. camerounensis (Bitja-Nyom, Agnèse, Pariselle, Bilong-Bilong, Gilles & Snoeks) (Fig. 29, [LU]) Distribution. Lower Aberdeen Reservoir (GBIF.org, 2021; current survey); Upper Aberdeen Reservoir (current survey). Native range. Africa. Remarks. Identified as the newly described Hemichromis cf. camerounensis due to the presence of two red opercular spots accompanied by a large black spot, separating it from all other Hemichromis species (Bitja-Nyom et al., 2021). However, DNA barcoding of the cytochrome b gene is necessary to confirm its identity, as some populations of Hemichromis elongatus are known to have the same feature (Bitja-Nyom et al., 2021)., Published as part of Chan, Jeffery C. F., Tsang, Alphonse H. F., Yau, Sze-man, Hui, Tommy C. H., Lau, Anthony, Tan, Heok Hui, Low, Bi Wei, Dudgeon, David & Liew, Jia Huan, 2023, The non-native freshwater fishes of Hong Kong: diversity, distributions, and origins, pp. 128-168 in Raffles Bulletin of Zoology 71 on page 144, DOI: 10.26107/RBZ-2023-0012, http://zenodo.org/record/7815765, {"references":["Bitja-Nyom AR, Agnese JF, Pariselle A, Bilong-Bilong CF, Gilles A & Snoeks J (2021) A systematic revision of the five-spotted Hemichromis complex (Cichliformes: Cichlidae) from West Africa and Lower Guinea, with the description of a new species from Cameroon. Hydrobiologia. 848 (16): 3779 - 3803."]}

The East African Great Lakes are inhabited by impressive radiations of cichlids that display a large variation in morphology, diet, colour pattern, and behaviour and have adapted to a large array of niches. Whilst most of these cichlids are bottom dwellers, a number of species have adapted to a pelagic environment and inhabit open waters. From the Lake Edward system, one pelagic species had been previously described, the zooplanktivorous Haplochromis pappenheimi. Our sampling revealed the Lake Edward system to be inhabited by two more pelagic species that were unknown to science and are formally described here, the zooplanktivorous Haplochromis pelagicus sp. nov. and the insectivorous H. aureus sp. nov. All three species seem mostly restricted to deepwater regions where the depth exceeds six metres in Lake Edward, except for H. pelagicus sp. nov., which also occurs in shallower regions of Lakes Edward and George and of the Kazinga Channel. Sexual dimorphism, mainly in head shape, was discovered in H. pappenheimi and H. pelagicus sp. nov. These differences suggest that females of these species have larger buccal cavities than males, which is most likely linked to female mouth brooding behaviour. This research was conducted within the framework of the BELSPO (Belgian Science Policy Ofce) funded BRAIN projects HIPE (Human impacts on ecosystem health and resources of Lake Edward) and KEAFish (The biodiversity, biogeography and evolutionary history of the northern basins of the Great African Lakes: the enigmatic fsh faunas of Lakes Kivu, Edward and Albert revisited) and an FWO (Research Foundation Flanders) funded PhD fellowship to NV (11E0520N). MM beneftted from a FishBase and Fish taxonomy training (2016) through the framework agreement of the RMCA with the Directorate-General for Development Cooperation and Humanitarian Aid. The feldwork by NV, MVS, and ED was supported by the FWO and feldwork by MVS by the King Leopold III Fund for Nature Exploration and Conservation. We thank L. Wasswa (Ugandan Fisheries Department) and M. Bifamengo (NaFIRRI, Uganda) for their help in collecting specimens, and W. Okello (NaFIRRI, Uganda) for valuable logistic support. We are grateful to M. Parrent (RMCA), O. Pauwels (IRSNB), J. Maclaine (NHMUK), and E. Aßel (ZMB) for curatorial services, E. Aßel for the provided photograph and X-ray image of the lectotype of H. pappenheimi, and support from the SYNTHESYS+ Project, http://www.synthesys.info/, which is fnanced by European Community Research Infrastructure Action under the H2020 Integrating Activities Programme, Project number 823827.

Understanding the interspecific morphological variability of Caquetaia kraussii (Perciformes: Cichlidae) between different localities in its distribution range is becoming essential, as this species constitutes a valuable resource for the economy and subsistence of the local human communities where it is endemic in Colombia and Venezuela. In order to develop efficient farming and handling plans for this species, a deep understanding of the factors and mechanisms generating morphological variability is crucial. This study analyzes the morphological variability of C. kraussii by using geometric morphometrics in four localities distributed between the Dique and North channels, which are part of the Bolívar department in Colombia. Likewise, the effect of environmental variables such as temperature (T°), dissolved oxygen (OD) and pH on morphological variability was analyzed using a partial least squares approach. The results show that environmental stress has an influence on ~10% of the body shape of C. kraussii, whereas ~90% of the body shape is not directly influenced by environmental parameters, suggesting an effect from stress related to sexual dimorphism. Similarly, the analyses show shape variation among localities, mainly between populations of lotic environments and those of lentic environments. This morphological disparity seems to be subject to environmental and sexual stresses in the different localities.