<Emphasis Type="Italic">Sinocyclocheilus guilinensis</Emphasis>, a new species from an endemic cavefish group (Cypriniformes: Cyprinidae) in China

The endemic Chinese cavefish genus Sinocyclocheilus possesses very rich species diversity. Specimens of this genus collected from Guilin, Guangxi, China, were originally described in an unpublished document in 1982. Later, they were recognized as Sinocyclocheilus jii. Comparison to the type specimens of S. jii leads to the conclusion that, though similar, they represent two different species. Herein we describe them as a new species, Sinocyclocheilus guilinensis sp. nov. It can be distinguished from all congeners by its possession of soft, unbranched dorsal ray without serrations, 19–20 (11–12) scale rows above (below) the lateral line, 34–36 circumpeduncular scales and 8–9 pre-dorsal vertebrae. Sinocyclocheilus guilinensis sp. nov. is most similar to S. jii. In addition to the scale count differences, a morphometrics-based principal component analysis (PCA) also supports designation of S. guilinensis and S. jii as two distinct species.     

Population Biology and Growth of Ozark Cavefish in Logan Cave National Wildlife Refuge, Arkansas

Ozark cavefish, Amblyopsis rosae, is a threatened species endemic to the Springfield Plateau of the Ozark Highlands in Arkansas, Missouri, and Oklahoma. One of the largest known Ozark cavefish populations, located in Logan Cave, Arkansas, was surveyed 25 times over a two-year period between 1993 and 1995. During the study, 147 Ozark cavefish > 30 mm (TL) were marked with visual implant tags and 140 Ozark cavefish were available for recapture; 68 were recaptured 189 times and the rest (72) were never recaptured. Individual Ozark cavefish persisted in Logan Cave for a relatively short time. Only 14% of 80 fish tagged during a previous study in 1992 were recaptured during this study, and half of all recaptured fish disappeared within three months. However, if a fish persisted for at least seven months in the cave, its probability of being recaptured over an additional year was high. Maximum persistence of a tagged fish was 28 months, suggesting these fish have a maximum life-span of 4–5 years. Growth averaged 0.6 mm per month, with maximum recorded growth of 6 mm per month and a maximum size of 65 mm TL. Smaller fish grew faster than larger fish but growth rates were sporadic, with several mid-sized fish (45–49 mm) showing little growth (0–3 mm year^-1) while some fish > 50 mm grew up to 12 mm year^-1. Most fish gained in length during April–October, the same period a maternity colony of gray bats occupied the cave. Gross Ozark cavefish movement over the study period ranged up to 1002 m, with a mean movement of 1.2 m day^-1; movement was positively correlated with Ozark cavefish total length. Death seemed the most likely explanation for loss of tagged Ozark cavefish, including fish that emigrated out of the cave. Little up-stream movement was recorded between reaches and did not account for loss of tagged fish. Reproduction within the cave and immigration from the aquifer accounted for persistence of Ozark cavefish in Logan Cave.     

Molecular phylogeny of the blind cavefish <Emphasis Type="Italic">Phreatichthys andruzzii</Emphasis> and <Emphasis Type="Italic">Garra barreimiae</Emphasis> within the family Cyprinidae

The phylogenetic relationships of two cavefish, Phreatichthys andruzzii and Garra barreimiae, belonging to the family Cyprinidae, were investigated by sequencing the mitochondrial cytochrome b gene. These cavefish species are native to Somalia (eastern Africa) and Oman (southeastern Arabian peninsula), respectively, and so far no molecular support to their taxonomy and phylogenetic position was ever provided. The analysis of cytochrome b sequences showed that the species are monophyletic taxa, closely related to each other and to other species of the genus Garra. Molecular clock calculations allowed to date the origin of these hypogaean species back to the Plio-Pleistocene and support the hypothesis that African cyprinids originated from Miocenic immigrations of Asian ancestors.     

Length–weight relationships of four cavefish species from China

Length‐weight relationships (LWRs) were estimated for four cavefish species belonging to the genus Sinocyclocheilus (Cypriniformes: Cyprinidae) occurring in the Nanpanjiang River and Hongshuihe River, which constitute the upstream of the species, S. angularis, S. furcodorsalis, S. rhinocerous, and S. tileihornes, were collected from 2011 to 2017. The b values of LWRs of these four species ranged from 2.830 to 3.719 and the correlation coefficient values (R²) were high for all species. This is the first record of LWRs for cavefish species with the horn in FishBase database.     

Siphonodella leiosa (Conodonta), a new unornamented species from the Tournaisian (lower Carboniferous) of Puech de la Suque (Montagne Noire,France)

A new conodont species, Siphonodella leiosa, is described from the lower Carboniferous pelagic limestones of the Montagne Noire (France), deposited in North Gondwana on a outer platform environment. Specimens were obtained from one level dated to the Siphonodella jii conodont Zone. The major difference from other siphonodellid conodonts known in this area is that the elements of this new species have a practically entirely smooth and unornamented platform, apart from the development of one or two low rostral ridge-like nodes. Similar morphologies were generally observed in shallow marine deposits of the same time frame from China, Russia and East and Central European areas. The new discovery reinforces the idea that ornamentation of siphonodellids is not only related to bathymetry, but that temperature could play an important role in the diversification and radiation of unornamented species during the Siphonodella jii conodont Zone.     

Prox 1 in eye degeneration and sensory organ compensation during development and evolution of the cavefish Astyanax

We have investigated expression of the homeobox gene Prox 1 during eye degeneration and sensory organ compensation in cavefish embryos. The teleost Astyanax mexicanus consists of sighted surface-dwelling forms (surface fish) and several populations of blind cave-dwelling forms (cavefish), which have evolved independently. Eye formation is initiated during cavefish development, but the lens vesicle undergoes apoptosis, and the eye subsequently arrests and degenerates. The requirement of Prox 1 for lens fiber differentiation and γ-crystallin expression in the mouse suggests that changes in the expression of this gene could be involved in cavefish eye degeneration. Surface fish and cavefish embryos stained with a Prox 1 antibody showed Prox 1 expression in the lens, neuroretina, myotomes, heart, hindbrain, and gut, as reported in other vertebrates. We found that Prox 1 expression is not altered during cavefish lens development. Prox 1 protein was detected in the lens vesicle as soon as it formed and persisted until the time of lens degeneration in each cavefish population. The cavefish lens vesicle was also shown to express a γ-crystallin gene, suggesting that Prox 1 is functional in cavefish lens development. In addition to the tissues described above, Prox 1 is expressed in developing taste buds and neuromasts in cavefish, which are enhanced to compensate for blindness. It is concluded that the Prox 1 gene is not involved in lens degeneration, but that expansion of the Prox 1 expression domain occurs during taste bud and neuromast development in cavefish. Received: 31 July 1999 / Accepted: 8 November 1999     

Modularity and sense organs in the blind cavefish, Astyanax mexicanus

SUMMARY Mexican tetra (Astyanax mexicanus) exist as two morphs: a sighted (surface) form and a blind (cavefish) form. In the cavefish, some modules are lost, such as the eye and pigment modules, whereas others are expanded, such as the taste bud and cranial neuromast modules. We suggest that modularity can be viewed as being nested in a manner similar to Baupläne so that modules express unique sets of genes, cells, and processes. In terms of evolution, we conclude that natural selection can act on any of these hierarchical levels within modules or on all the sensory modules as a whole. We discuss interactions within and between modules with reference to the blind cavefish from both genetic and developmental perspectives. The cavefish represents an illuminating example of module interaction, uncoupling of modules, and module expansion.     

Status of the Federally Endangered Alabama Cavefish, Speoplatyrhinus Poulsoni (Amblyopsidae), in Key Cave and surrounding Caves, Alabama

Speoplatyrhinus poulsoni, the Alabama cavefish, is a federally endangered stygobitic fish that is restricted to Key Cave in northwestern Alabama. Due to the low abundance of this species and the continuous threats to the cave's recharge area, we surveyed Key Cave from 1992 to 1997. Alabama cavefish were present in five different pools and had numbers comparable to previous surveys performed in the 1970's and 1980's. Three different size classes were observed, indicating recruitment is occurring. These data suggest that the population is relatively stable, but future monitoring is recommended. The Alabama cavefish is an endangered species and is extremely difficult to capture, therefore any marking program to accurately estimate the population size is unlikely. A single specimen of Typhlichthys subterraneus was captured in Key Cave. This falsifies the previous assumption that these two species are allopatric. Because of this discovery, we resurveyed seven caves in the area around Key Cave for additional populations of S. poulsoni. These caves had either known populations of T. subterraneus or unidentified cavefish. No additional populations of S. poulsoni were discovered, but further investigation is warranted. The Key Cave National Wildlife Refuge was established in the high recharge area of Key Cave to protect this habitat, yet threats to the groundwater continue from encroaching urbanization.     

The 30-year recovery effort for the Ozark cavefish (<Emphasis Type="Italic">Amblyopsis rosae</Emphasis>): Analysis of current distribution,population trends,and conservation status of this threatened species

Here we review the thirty year recovery effort and conservation status of the Ozark cavefish, Amblyopsis rosae. We summarized the historic and current range of the species, and report county range extensions for both A. rosae and its confamilial Typhlichthys subterraneus. Ozark cavefish survey data spanning almost a century were analyzed for temporal trends using the Mann-Kendall Test/Sen’s Slope Estimator Method. Results were inconclusive because variance was high and the majority of data sets were not sufficiently large to detect a trend. However, the two largest populations (Cave Springs Cave and Logan Cave, Benton Co., Arkansas) have stabilizing or increasing survey counts. While the number of active cavefish sites has decreased over 50% since 1990, the number of surveyed individuals has not. Reasons for endangerment were reanalyzed since federal listing; the primary threat has shifted from overcollection to habitat degradation. We analyzed the progress of recovery task implementation, and we critically evaluated the basis of delisting criteria. Recovery Task 1, the hydrogeologic delineation of subterranean habitats, is almost complete. Recovery Task 2 prescribes protection and management for Recovery Caves, and important progress has been made. Recovery Task 3 involves the development and implementation of monitoring programs in Recovery Caves. Several important studies have been performed, and indicate that many cavefish populations are experiencing chronic, low-level exposure to a suite of anthropogenic contaminants. Delisting conditions are largely unattainable as currently worded. We suggest that recovery criteria be amended such that habitat protection goals are attainable, that the list of Recovery Caves can be periodically updated, and that the recovery population goal is increased and distributed between more sites.     

A new cavefish species, Sinocyclocheilus brevibarbatus (Teleostei: Cypriniformes: Cyprinidae), from Guangxi, China

A new species of the cavefish genus Sinocyclocheilus is described based on six specimens collected from a subterranean river in Du’an County in the Guangxi Zhuang Autonomous Region of southern China. Sinocyclocheilus brevibarbatus sp. nov. can be distinguished from all congeners by the following combination of characteristics: a completely scaled body with well-developed eyes and a strongly-humped back; an almost straight lateral line possessing 49–51 perforated scales; seven soft rays on the dorsal fin; a relatively large eyeball (3.3–5.7% of SL) and short maxillary and rictal barbels (1.7–5.7% and 4.5–7.0% of SL, respectively).     

Sinocyclocheilus zhenfengensis,a new cyprinid species (Pisces: Teleostei) from Guizhou Province,Southwest China

Sinocyclocheilus zhenfengensis sp. nov. is a new species from an underground water environment in a karst cave located in Beipanjiang River of the Pearl River Basin in Shuangrufeng (Double Breasts Hill) Scenic, Etian Village, Zhexiang Township, Zhenfeng County, Guizhou Province, Southwest China. The new species has eyes, its body is completely scaled, an anteriorly directed bulge has formed on the back of its head, and a long pectoral fin exceeds the pelvic fin insertion. The body of the new species is apparently similar to that of S. bicornutus, but differs in scale rows above the lateral line, scale rows below the lateral line, snout length, and eye diameter. Phylogenetic reconstruction of the cytochrome b gene using the maximum parsimony (MP) and Bayesian inference (BI) methods indicates that S. zhenfengensis is closely related to S. bicornutus, S. tianeensis, S. jiuxuensis, and S. altishoulderus; however, it differs in terms of eye size, lateral line scales, scale rows above and below the lateral line.     

Albinism in phylogenetically and geographically distinct populations of Astyanax cavefish arises through the same loss-of-function Oca2 allele

The Mexican tetra, Astyanax mexicanus, comprises 29 populations of cave-adapted fish distributed across a vast karst region in northeastern Mexico. These populations have a complex evolutionary history, having descended from ‘old'' and ‘young'' ancestral surface-dwelling stocks that invaded the region ∼6.7 and ∼2.8 MYa, respectively. This study investigates a set of captive, pigmented Astyanax cavefish collected from the Micos cave locality in 1970, in which albinism appeared over the past two decades. We combined novel coloration analyses, coding sequence comparisons and mRNA expression level studies to investigate the origin of albinism in captive-bred Micos cavefish. We discovered that albino Micos cavefish harbor two copies of a loss-of-function ocular and cutaneous albinism type II (Oca2) allele previously identified in the geographically distant Pachón cave population. This result suggests that phylogenetically young Micos cavefish and phylogenetically old Pachón cave fish inherited this Oca2 allele from the ancestral surface-dwelling taxon. This likely resulted from the presence of the loss-of-function Oca2 haplotype in the ‘young'' ancestral surface-dwelling stock that colonized the Micos cave and also introgressed into the ancient Pachón cave population. The appearance of albinism in captive Micos cavefish, caused by the same loss-of-function allele present in Pachón cavefish, implies that geographically and phylogenetically distinct cave populations can evolve the same troglomorphic phenotype from standing genetic variation present in the ancestral taxon.     

Sinocyclocheilus donglanensis, a new cavefish (Teleostei: Cypriniformes) from Guangxi, China

Sinocyclocheilus donglanensis, a new cyprinid species from a subterranean river in Donglan County in the Guangxi Zhuang Autonomous Region of southern China, is described. It is distinguished from all congeners by the following combination of characteristics: a completely scaled body with well-developed eyes; a curved lateral line possessing 57–64 scales; pectoral fin not reaching pelvic fin origin and last unbranched ray of the dorsal fin clearly serrated along its posterior edge; 8–9 predorsal vertebrae; 8–9 gill rakers; joints of dentary-angulars not close to each other at the isthmus; and a slightly inferior mouth with the upper jaw (6.2–7.4% in standard length: SL) protruding slightly beyond the lower one (5.7–6.7% SL). Sinocyclocheilus donglanensis is sympatric with the peculiarly shaped, hunchbacked S. altishoulderus.     

The Hoosier cavefish,a new and endangered species (Amblyopsidae,Amblyopsis) from the caves of southern Indiana

We describe a new species of amblyopsid cavefish (Percopsiformes: Amblyopsidae) in the genus Amblyopsis from subterranean habitats of southern Indiana, USA. The Hoosier Cavefish, Amblyopsis hoosieri sp. n., is distinguished from A. spelaea, its only congener, based on genetic, geographic, and morphological evidence. Several morphological features distinguish the new species, including a much plumper, Bibendum-like wrinkled body with rounded fins, and the absence of a premature stop codon in the gene rhodopsin. This is the first new cavefish species described from the United States in 40 years and exemplifies how molecular data can alert us to the presence of otherwise cryptic biodiversity.     

Metabolic Rate and Natural History of Ozark Cavefish, Amblyopsis Rosae, in Logan Cave, Arkansas

We investigated the effects of mass, season, and activity on oxygen consumption of the federally threatened Ozark cavefish, Amblyopsis rosae, at Logan Cave National Wildlife Refuge, Benton County, Arkansas. We used an acclimatization protocol to measure in-situ metabolic rates. There was a significant effect of mass on the metabolic rate of Ozark cavefish and this relationship differed significantly among seasons. We observed a positive relationship between mass and oxygen consumption during summer and autumn but a negative relationship for winter and spring. There was a 1°C water temperature change between summer/autumn sampling and winter/spring sampling, but it is not likely temperature alone accounted for seasonal variation in mass-oxygen consumption relationships. Activity in the respirometer did not vary significantly among seasons and was not correlated with fish mass or time of day. Seasonal relationships between mass and oxygen consumption may therefore reflect alterations in environmental conditions (i.e. food availability, ambient dissolved oxygen), condition, changes in susceptibility to handling stress, or may reflect low numbers of fish tested. Natural history observations were made throughout the study. Four females were seen in late August with ova visible in the body cavity. On 20 June 1996 we first observed five small cavefish (approximately 10 mm TL) in a pool just upstream of the sinkhole entrance that were likely less than a month old. One large adult (55 mm TL) was seen in the same pool with the young fish for six weeks, until two small fish were displaced by a storm event. Although this does not provide a clear sign of parental care, it does suggest that cannibalism is not always the rule for this species as previously suggested.     

Phototactic response and light sensitivity in an epigean and a hypogean population of a barb (Garra barreimiae, Cyprinidae)

We investigated the response of two populations of the barb Garra barreimiae to different light intensities (0.5–2000 lx) from a light source. Adults of both the surface (epigean) and cave (hypogean) G. barreimiae populations show photophobic behavior. A photophobic response in the cave form was seen only at higher light intensities because the cavefish are eyeless and rely on extra-retinal light receptors to detect light. In contrast, juveniles (surface and cave) showed photophilic behavior, and their preference for the photic zone of the test tank decreased with increasing age. We discuss the potential role played by photophobic behavior for the colonization of caves by previously surface-dwelling fishes.     

A new cavefish of Sinocyclocheilus (Teleostei: Cypriniformes: Cyprinidae) from the Nanpanjiang River in Guizhou,China

A new species, Sinocyclocheilus xingyiensis, is described based on specimens collected from a karst cave in Guizhou Province, China. The authors used an integrated taxonomic approach, including morphological and molecular data, to identify the new species as a member of the Sinocyclocheilu angularis group, and it can be distinguished from all other members of this group by a combination of the following features: two pairs of long barbels and long pectoral fins, 42–46 lateral-line scales, 7 (13–14) on outer (inner) side of the first gill arch and 35 (14–15 + 4 + 16 − 17) vertebrae. Phylogenetic analyses based on the cytochrome b (cyt b) gene fragment suggest that S. xingyiensis is a sister lineage to Sinocyclocheilus flexuosdorsalis. The genetic distance (Kimura 2-parameter) between the S. xingyiensis and S. angularis groups of Sinocyclocheilus species based on cyt b gene fragment ranged from 1.2% to 15.4%.     

Length-to-weight and length-to-length ratios of six cavefish species from the Pearl River Basin,South China

Length-to-weight and length-to-length ratios were estimated for six cavefishes including Yunnanilus longibarbatus, Yunnanilus bailianensis, Yunnanilus jinxiensis, Oreonectes polystigmus, Paranemachilus pingguoensis, and Paranemachilus genilepis from the upper reaches of the Pearl River, Guangxi Province, China (between 106.26E, 23.14N and 111.42E, 24.85N). The fish were captured between May and October in 2018 and 2019 using trap nets (mesh size 7 mm) set in front of selected caves at 6:30–7:30 p.m. and were hauled out at 6:30–7:30 a.m. the next day. After removal, the fish were measured and weighed immediately to the nearest 0.1 cm and 0.1 g, respectively. The regression parameter r² was between .95 and .98. The total length-to-standard length ratios were also calculated with r² ≥ .98. This study should provide a useful reference for biological studies and biodiversity conservation of cavefish.     

Active wall following by Mexican blind cavefish (Astyanax mexicanus)

When introduced into a novel environment that limits or prevents vision, a variety of species including Mexican blind cavefish (Astyanax mexicanus) exhibit wall-following behaviors. It is often assumed that wall following serves an exploratory function, but this assertion remains untested against alternative artifactual explanations. Here, we test whether wall following by cavefish is a purposeful behavior in which fish actively maintain a close relationship with the wall, or an artifactual consequence of being enclosed in a small concave arena, in which fish turn slightly to avoid the wall whenever it impedes forward movement. Wall-following abilities of fish were tested in a large, goggle-shaped arena, where forward motion along the convex wall was unimpeded. In this circumstance, cavefish continued to follow the wall at frequencies significantly above chance levels. Lateral line inactivation significantly reduced the ability of fish to follow convex, but not concave or straight, walls. Wall-following abilities of normal fish decreased with decreasing radius of wall convex curvature. Our results demonstrate that cavefish actively follow walls of varying contours. Radius-of-curvature effects coupled with the difficulties posed by convex walls to lateral line-deprived fish suggest a partially complementary use of tactile and lateral line information to regulate distance from the wall.