Caleb D. Mcmahan

Phylogenomic Systematics of Ostariophysan Fishes: Ultraconserved Elements Support the Surprising Non-Monophyly of Characiformes

Abstract.— Ostariophysi is a superorder of bony fishes including more than 10,300 species in 1100 genera and 70 families. This superorder is traditionally divided into five major groups (orders): Gonorynchiformes (milkfishes and sandfishes), Cypriniformes (carps and minnows), Characiformes (tetras and their allies), Siluriformes (catfishes), and Gymnotiformes (electric knifefishes). Unambiguous resolution of the relationships among these lineages remains elusive, with previous molecular and morphological analyses failing to produce a consensus phylogeny. In this study, we use over 350 ultraconserved element (UCEs) loci comprising 5 million base pairs collected across 35 representative ostariophysan species to compile one of the most data‐rich phylogenies of fishes to date. We use these data to infer higher level (interordinal) relationships among ostariophysan fishes, focusing on the monophyly of the Characiformes—one of the most contentiously debated groups in fish systematics. As with most previous molecular studies, we recover a non‐monophyletic Characiformes with the two monophyletic suborders, Citharinoidei and Characoidei, more closely related to other ostariophysan clades than to each other. We also explore incongruence between results from different UCE data sets, issues of orthology, and the use of morphological characters in combination with our molecular data. [Conserved sequence; ichthyology; massively parallel sequencing; morphology; next‐generation sequencing; UCEs.]

Validity and Redescription of Profundulus balsanus Ahl, 1935 (Cyprinodontiformes: Profundulidae)

Profundulus balsanus was described by Ahl in 1935 and later placed in synonymy with P. punctatus. Recent molecular studies have demonstrated that specimens identified as P. balsanus belong to a distinct taxon. Herein we used a combination of morphometric, meristic, coloration and osteological data in order to demonstrate the distinctiveness of P. balsanus. Additionally, we used molecular data to reconstruct a phylogeny to place P. balsanus in a systematic context. Profundulus balsanus belongs to the punctatus clade within the genus Profundulus. It can be distinguished from P. oaxacae based on scale count in the lateral line and from P. punctatus based on differences in coloration in the anal fin. We provide a taxonomic key to all Mexican species of Profundulus.

Pleistocene to holocene expansion of the black-belt cichlid in Central America, Vieja maculicauda (Teleostei: Cichlidae)

The distributions of many Northern Hemisphere organisms have been influenced by fluctuations in sea level and climatic conditions during Pleistocene interglacial periods. These cycles are associated with range contraction and refugia for northern-distributed organisms as a response to glaciers. However, lower sea levels in the tropics and sub-tropics created available habitat for expansion of the ranges of freshwater organisms. The goal of this study was to use ecological niche modeling to test the hypothesis of north to south range expansion of Vieja maculicauda associated with Pleistocene glacial cycles. Understanding the biogeography of this widespread species may help us better understand the geology and interconnectivity of Central American freshwaters. Occurrence data for V. maculicauda was based on georeferencing of all museum records of specimens recovered from FishNet2. General patterns of phylogeographic structure were assessed with mtDNA. Present day niche models were generated and subsequently projected onto paleoclimatic maps of the region during the Last Interglacial, Last Glacial Maximum, and mid-Holocene. Phylogenetic analysis of mtDNA sequence data showed no phylogeographic structure throughout the range of this widespread species. Present day niche models were congruent with the observed distribution of V. maculicauda in Central America. Results showed a lack of suitable freshwater habitat in northern Central America and Mexico during the Last Interglacial, with greatest range expansion during the Last Glacial Maximum and mid-Holocene. Results support the hypothesis of a north to south range expansion of V. maculicauda associated with glacial cycles. The wide distribution of this species compared to other closely related cichlids indicates the latter did not respond to the degree of V. maculicauda in expansion of their distributions. Future work aimed at comparisons with other species and modeling of future climatic scenarios will be a fruitful area of investigation.

Redescription of Vieja hartwegi (Taylor & Miller 1980) (Teleostei: Cichlidae) from the Grijalva River basin, Mexico and Guatemala, with description of a rheophilic morph

Vieja hartwegi was described by Taylor and Miller in 1980 based on 45 juvenile and sub-adult specimens, but this species shows an enormous variation in coloration patterns during ontogenic development and between individuals. Additionally, morphological data have demonstrated the existence of two forms in V. hartwegi: the typical morph of a moderately deep to slightly elongated body shape with midlateral band, and a rheophilic morph with an elongated to slightly deep body shape and lack of midlateral band. Herein we redescribe V. hartwegi adding adults and individuals from broad geographic ranges, as well as a rheophilic morph recently discovered in the upper reaches of the Grijalva River basin in Mexico. The set of characters include coloration, meristic, morphometric and osteological features. Vieja hartwegi is a polymorphic species whose forms may be segregating because of habitat preferences and food habits.

Redescription of Trichomycterus striatus (Meek & Hildebrand, 1913) (Siluriformes: Trichomycteridae), with notes on its geographic distribution

Trichomycterus striatus is herein redescribed, based on examination of the types and recently collected specimens, and its geographic distribution is updated. This species can be diagnosed from all other northeastern South American congeners by its variable coloration pattern consisting of a yellowish to light brown background with a black lateral band and/or small dark brown spots on sides or uniformly light brown and by the following combination of characters: teeth conical arranged in three to four irregular rows in both jaws; anterior section of infraorbital canal (sensory pores i1 and i3) present; sensory pores s6 paired, 11-23 opercular odontodes; 27-44 interopercular odontodes; seven to eight pectoral-fin branched rays; 36-37 free vertebrae; 12-14 ribs; cleithrum pierced by several foramina; and caudal fin truncate to rounded. Trichomycterus striatus occurs from southern Costa Rica [from the Pirrís (herein reported for the first time), Térraba and Coto River basins] to eastern Panama (in most of the main river basins in both the Pacific and Atlantic versants), being the sole representative of the family in lower Central American waters.

A new species of Astyanax (Characiformes: Characidae) from Costa Rica and Panama, with a key to the lower Central American species of the genus: a new astyanax from central america

Astyanax anai, a new species of characid fish, is described from the Sixaola River basin, eastern Costa Rica-western Panama, Central America. The new species can be distinguished from all other congeners by the following combination of characters: premaxillary teeth 4-5 at the inner series and 4-6 at the outer series; maxillary teeth tricuspid, 2-4; predorsal scale series irregular and incomplete, with an unscaled space behind tip of supraoccipital process and 12-14 scales; lateral line scales 34-39; humeral region with a conspicuous black and rounded to horizontally ovate spot and two diffuse brown and vertically elongate bars (the first through the rounded to horizontally ovate spot, the second 2-4 scales behind the first); body depth 36·6-42·3% of standard length (LS ); midlateral stripe formed by a series of 10-14 anteriorly-directed dermal herringbone, or chevron-shaped, marks, most apparent in juveniles and in preserved specimens, extending above the lateral line from the black humeral spot or just behind it (from the second vertical bar) to the caudal peduncle; scale rows from lateral line to base of first dorsal-fin ray 8-9; scale rows from lateral line to base of pelvic fin 7-8; pre-anal distance 53·9-61·9% of LS ; total anal-fin elements 29-33; caudal spot elongated, rhomboid or rectangular, with its anterior margin surpassing the middle of the caudal peduncle, usually reaching the anal-fin insertion, posteriorly covering 4-7 principal caudal-fin rays and not extending onto the ventral and dorsal margins of the caudal peduncle, covering 3-5 horizontal scale rows. In order to test the phylogenetic relationships of the new taxon in relation to the other North and Central American species of the genus, a new phylogenetic hypothesis based on a reanalysis of the morphological matrix by Schmitter-Soto (2016) is proposed. A key to the lower Central American (southern Nicaragua to eastern Panama) species of Astyanax is also provided.

Assessing population-level morphometric variation of the Mountain Mullet Agonostomus monticola (Teleostei: Mugilidae) across its Middle American distribution

Population-level morphometric variation of the Mountain Mullet (Agonostomus monticola) was assessed in 419 adult specimens from 25 sample sites (river basins) across its Middle American distribution (Pacific and Atlantic-Caribbean drainages). This analysis was based on 36 standardized linear measurements and 19 landmarks on geometric morphometrics approach. Discriminant function analysis (DFA) revealed 19 linear morphological characters with significant variation among groups. Geometrically, the most notable changes were associated to the curvature of the frontal region of the head, the anterior and posterior insertion of the first dorsal and anal fins. The resulting grouping based on the DFA and geometric morphometrics techniques (Pacific-A, Pacific-B and NE México-Caribbean) were similar to those previously recovered by genetic techniques, where the Pacific-B (Ayuquila river basin) was the most different group. Our results provide morphological evidence for considering Agonostomus monticola as a complex of evolutionary entities, represented by two forms in the Pacific Ocean and another in the Atlantic Ocean.

Clarification of type material for Bornean species of the genus Osteochilus (Cyprinidae)

Two species and one subspecies of the genus Osteochilus (Cypriniformes: Cyprinidae) were described from Borneo in a revision of the group by Karnasuta in 1993. Examination of type material has illuminated points of conflict regarding some of the type specimens listed in original descriptions. Based on archived correspondence and the authors original thesis, we were able to determine the disposition of specimens and revise collection information for type material accordingly.