Population genetics of grotto sculpin (Cottus specus), a new cave-adapted fish species

The grotto sculpin (Cottus specus) is a troglomorphic fish endemic to cave systems in Perry County, Missouri. These fish are state-threatened and were recently listed as federally endangered under the Endangered Species Act (ESA). Due to the unstable nature of the cave environment, grotto sculpins are highly susceptible to pollution via agricultural and waste runoff. Here we provide a summary of population genetic diversity within the range of the grotto sculpin. Samples from twenty-one cave, surface, and resurgence sites within and surrounding the Bois Brule drainage in Perry County, southeast Missouri, USA were collected and sequence data from the mitochondrial control were analyzed using analysis of molecular variance, maximum likelihood, parsimony, and Bayesian techniques. We found a substantial degree of divergence from surface populations of Midlands and Black River banded sculpin races (7.7–8.0 %). Within the Bois Brule drainage, two distinct lineages of grotto sculpin were identified which correspond with northern and southern cave system boundaries and may represent independent cave invasions.     

Phylogeography of cave pseudoscorpions in southern Australia

Aim  To investigate molecular phylogenetic divergence and historical biogeography of the cave-dwelling pseudoscorpion genus Protochelifer.
Location  Caves and nearby epigean habitats in southern Australia were sampled from western Victoria, Naracoorte Caves, Flinders Ranges, Kangaroo Island, Nullarbor Plain and south-west Western Australia.
Methods  Allozyme electrophoresis (57 individuals) and a 569-base-pair section of the mtDNA COI gene (22 individuals) were used to reconstruct phylogenetic relationships among four cave species and three epigean species from 13 locations.
Results  Phylogenetic reconstruction using the allozyme and mtDNA sequence data revealed a similar topology, showing recent speciation of several Protochelifer populations in caves from Naracoorte to the Nullarbor Plain. Naracoorte Caves contained a single species, Protochelifer naracoortensis , found in four separate caves, while all other cave species appear to be restricted to single caves.
Main conclusions  At a local scale, as indicated by the four Naracoorte caves, dispersal is thought to occur via micro- and mesocaverns, and possibly by phoresy using insect or bat vectors. With current data we are unable to determine if cavernicolous species of Protochelifer have arisen from a single cave colonization event followed by phoretic dispersal on bats to other caves, or multiple cave-invasion events from independent epigean ancestors. Genetic heterogeneity among Protochelifer populations from Nullarbor caves suggest that P. cavernarum , the only species presently recorded from this region, is likely to constitute a species complex requiring further study to fully resolve its relationships.     

Shape variation in surface and cave populations of the armoured catfishes Ancistrus(Siluriformes: Loricariidae) from the São Domingos karst area, upper Tocantins River, Brazil

Body shape of four distinct populations of Ancistrus cryptophthalmus from different caves in the São Domingos karst area and the local epigean (surface) Ancistrus species were compared using geometric morphometrics. Angélica and Bezerra catfishes were closer to each other and successively to the studied sample of epigean catfishes than to Passa Três and São Vicente I catfishes. Greater similarity between catfishes from the Angélica‐Bezerra cave system to epigean catfishes than to other cave catfishes may represent a plesiomorphic feature. On the other hand, the present analyses allowed the distinction between Passa Três and São Vicente I catfishes, which inhabited the same cave system. Topographic isolation due to the presence of several waterfalls interposed between the stream reaches where these catfishes live may explain such morphological differentiation. Also, the presence of a waterfall downstream of the reach where the Angélica sample was obtained may have contributed to the partial isolation of these catfishes from those living in Bezerra Cave. Differences in population sizes probably influenced the degrees of morphological differentiation observed among the cave populations. The mosaic distribution of character states indicated that the four populations of A. cryptophthalmus have evolved rather independently, but the continuous variation suggests a recent connection between these populations, perhaps with a limited gene flow. Considering other features such as reduction of eyes and pigmentation, and changes in body shape, a separate species status for the epigean and hypogean Ancistrus species from the São Domingos karst area has been maintained.     

Epigean and subterranean ichthyofauna from the São Domingos karst area, Upper Tocantins River basin, Central Brazil

In a survey of epigean and cave streams in the São Domingos karst area, 38 species were present in the stream reaches, including 10 characiforms, 19 siluriforms, seven gymnotiforms and two perciforms. One species of Astyanax and the new armoured catfish species Parotocinclus were the most common epigean fishes in São Domingos. The most conspicuous non-troglomorphic cave fishes were Hoplerythrinus unitaeniatus , Astyanax sp., Brycon sp. and two species of Imparfinis , with I. hollandi being the most common fish in most caves. São Domingos karst area has the most diverse and abundant Brazilian cavefish fauna, not only in terms of troglobitic species but also in general fish richness, with 22 non-troglomorphic species recorded in caves in addition to five troglobitic ones. Most fishes examined for stomach contents had at least partially full stomachs. The studied fishes were carnivorous, feeding primarily on aquatic insects. Fishes with developed gonads recorded in caves were I. hollandi , Rhamdia quelen , Pseudocetopsis plumbeus , Hoplerythrinus unitaeniatus and Cichlasoma araguaiensis , indicating a potential for reproduction in the subterranean habitat.     

Hickmania Troglodytes, the Tasmanian Cave Spider, and its Potential Role in Cave Management

Cave faunas – which often contain a high representation of spiders – are subject to increasing pressure from the effects of epigean habitat degradation and recreational caving activities. Hickmania troglodytes is a prominent member of the Tasmanian cave fauna, a spider of phylogenetic, zoogeographic and ecological importance, but about which little has previously been known. Long-term monitoring has revealed many unusual life-cycle characteristics in this species, most of which occur over long periods of time and are dependent upon environmental stability. The species presents a potentially useful tool in the management and monitoring of cave fauna and karst, as it is large, conspicuous, numerous, ubiquitous, sedentary, functionally significant and potentially sensitive to various sources of disturbance. H. troglodytes may provide a visible and obvious measure of disturbance in and around cave entrances, and may also prove useful in detecting broader scale impacts affecting the entire cave. Many promising developments are being made in terms of cave management in Tasmania, but other issues are less well addressed and still need to be resolved. With further research, the use of indicator or sentinel species may prove to be well suited to the less complex and often sparsely populated subterranean environment, and may play an important role within larger management strategies for cave fauna and karst.     

贵州荔波玉屏洞穴陆生动物食物来源及营养级

洞穴是“极端地下生物多样性”的聚集地, 为了揭示洞穴陆生动物的食物来源以及营养级关系, 本文于2016年10月对贵州荔波玉屏镇的3个代表性洞穴内的陆生动物进行调查, 应用¹³C和¹⁵N稳定同位素方法, 分析了洞内陆生动物和基础碳源的δ¹³C、δ¹⁵N值变化特征, 探究了各动物类群的营养级位置以及不同的基础碳源对动物的食物源的贡献率。结果表明: 洞穴植物的δ¹³C值范围为-38.25‰ ± 0.95‰至-29.44‰ ± 0.49‰, 土壤有机质δ¹³C值范围为-32.73‰ ± 0.03‰至-24.68‰ ± 0.41‰, 大部分不在洞穴植物的δ¹³C变化范围内。洞穴动物δ¹³C、δ¹⁵N值范围分别为-34.22‰ ± 0.39‰至-14.83‰ ± 0.78‰、1.86‰ ± 0.20‰至16.43‰ ± 0.05‰, 表现出较大的种(或类群)间差异。IsoSource软件估算结果表明, 洞穴土壤有机质对78.3%动物类群的食物源的贡献率高于植物。荔波玉屏洞穴消费者的营养级范围为1.01-5.32, 其中, 巨蚓科、螺类、潮虫科、马陆、大蚊科、夜蛾科主要处于第2营养层次; 蚰蜒科、蚁蛉科(幼虫)、蝙蝠处于第3营养层次; 蜚蠊科处于第2-3营养层次; 驼螽科、蠼螋科处于第2-4营养层次; 蜘蛛处于第2-5营养层次, 跨3个营养级。综上认为, 洞穴土壤有机质是洞穴消费者的主要食物基础。大部分同一类群的动物在洞穴中所处的营养级位置较稳定, 少部分同属于一个类群的动物在不同洞穴或同一洞穴不同光带所处的营养级位置有差异。     

中华菊头蝠的栖息生态特征及种群保护

中华菊头蝠(Rhinolophus sinicus)主要分布于我国境内,属于典型的洞栖食虫性蝙蝠,具有重要的研究意义与保护价值。近年,在湘西州境内的16个溶洞中对其栖息生态特征进行了观察,共记录到其个体12942只次,数据分析结果显示：该蝠是该区域溶洞中的居留型优势蝠种,具有集群冬眠的习性。每年的9月中下旬开始逐步聚群冬眠,翌年的3月底开始苏醒,其夏季的繁育场所非常隐蔽,通常位于人无法进入的洞段或人迹罕至的洞穴。总体上,其栖点主要集中于距洞口150—200 m(91.60%)的洞段,离地高度6—10 m(91.75%),多以“双足悬挂”的栖姿将身体悬空倒挂于洞道的顶壁。其体温与栖点温度之间不存在显著性差异(P>0.05),但具有明显的线性正相关关系(R~2=0.8886)。其栖点安全性春夏季高,秋冬季中等偏低。中华菊头蝠作为该区域洞栖性蝙蝠的优势种,在维持洞穴生态系统的稳定中发挥着至关重要的作用,加强对洞穴生态系统中蝙蝠伞护性的研究有助于洞穴生态保护教育及对蝙蝠种群的保护实践。     

Habitat and Population Data of Troglobitic Armored Cave Catfish, Ancistrus Cryptophthalmus Reis, 1987, from Central Brazil (Siluriformes: Loricariidae)

Population ecology of the troglobitic armored catfish, A. cryptophthalmus, from caves in São Domingos karst area of central Brazil, was studied in a two-census capture-mark-recapture study carried out during the dry season of 1999. Like many loricariids, including epigean Ancistrus catfishes from that area, A. cryptophthalmus is a typical bottom-dweller with a strong preference for rocky substrates in fast-flowing stream sectors. Relatively high population densities, when compared to other cavefishes, were recorded in the studied cave sections: 0.9 individuals m^–2 in Angélica Cave and 0.5 individuals m^–2 in Passa Três Cave. Passa Três Stream is a small tributary of the São Vicente Cave system, and the estimated population for Passa Três Cave (ca. 2000 m long) is around 1000 individuals. On the other hand, the large Angélica Cave Stream (8 km long) harbors a very large population, conservatively estimated at 20 000 individuals. No fish movements of distances longer than 100 m were recorded in the 40-day period. In Passa Três Cave, a few movements of distances up to 60 m were detected. Proportion of mature females in Angélica Cave was higher than in Passa Três Cave, possibly reflecting a greater food availability in the first cave. Mean individual growth rates in Angélica Cave (1.35 mm month^–1 for size class <41 mm SL, 0.17 mm month^–1 for size class >41 mm SL) point to relatively high longevities, possibly attaining 15 years.     

Nontroglobitic Fishes in Bruffey-Hills Creek Cave, West Virginia, and other Caves Worldwide

Six species of fishes were tagged and released in September and November 1995 and on five dates between November 1998 and October 1999 inside Bruffey-Hills Creek Cave. Most of the tagged fishes were creek chub, Semotilus atromaculatus, and green sunfish, Lepomis cyanellus. The overall recapture rate was 2.6% as only three of 117 fishes were recaptured. Forty-nine days was the longest time of residence by two L. cyanellus and one S. atromaculatus, and movement of 83.4 m was observed only for the creek chub. Tag loss was confirmed as one factor in the low recapture rate. Nine species of fishes were collected in the cave, including Phoxinus oreas and Pimephales promelas, two species never collected in a cave. Most of the fishes were pigmented normally, but many creek chubs were depigmented and appeared white or pigmentless when first observed in the cave stream. Four species, Phoxinus oreas, Pimephales promelas, Ameiurus nebulosus, and Lepomis macrochirus, were collected for the first time in Bruffey-Hills Creek Cave and in the Bruffey and Hills creeks drainage. Although data from this study did not shed light on residence time of fishes in the cave, the consistent occurrence of epigean fishes in this cave was shown. A list of epigean fishes from caves worldwide was included along with a discussion of aspects of the ecology of epigean fishes in caves and of evolution of troglobitic fishes.     

The Occurrence of Fungi in the Recently Discovered Jarkowicka Cave in the Karkonosze Mts. (Poland)

Our study is one of the very few cases of speleomycological research in recently discovered caves. The aim of this research was to assess the population size of fungal colonies and their species composition in the Jarkowicka cave, discovered in 2012. The air samples were taken from one location outside the cave and from two locations inside of it. Mycological evaluation of the rocks inside the cave was performed usingswab sampling procedure. In the Jarkowicka cave we found 22 species of fungi, including 13 isolated from air at the entrance and from the walls, and 8 species from air inside the cave. Cladosporium spp. were the fungi most frequently isolated from internal atmosphere of the Jarkowicka cave, and from the external air. On the other hand, the fungi most frequently isolated from the rocks were Mucor spp. We found several species not yet described as cave inhabitants: Hypocrea pachybasioides, Cladosporium uredinicola, and Embellisia abundans. Our study may provide a basis for comparison to other similar studies conducted in frequently visited caves by tourists.     

Ecological characteristics of a cave‐dwelling moth species,Triphosa dubitata (Lepidoptera: Geometridae), in Baram and Ssang caves in Gangwon Province,Korea

This study was performed to clarify the ecological characteristics of a cave‐dwelling moth species, Triphosa dubitata Linnaeus (Lepidoptera: Geometridae), at Baram cave and at Ssang cave within Pyeongchang‐gun of Gangwon province in south Korea. We investigated T. dubitata population for 3 days from March 2013 to October 2013 (except September) and in January 2014. We used a section investigation method in the caves every month to determine their ecological characteristic. A nocturnal investigation using a high intensity discharge lamp adjacent to the entrance of the cave was conducted in parallel with a diurnal investigation using butterfly traps to investigate the movement of T. dubitata. In order to investigate the range of habitat temperature of moths, we used a dual laser targeting infrared thermometer to measure the cave temperatures. We caught individuals of T. dubitata L.: 8,574 individuals in Baram cave and 828 individuals in Ssang cave. As 9,095 individuals (97 %) appeared at a temperature of ?3°C ~ 8°C, it was verified that the adults were living at specific temperatures. In addition, T. dubitata adults were not active outside the caves; they entered the cave as soon as they developed from pupa into adults. These results showed that T. dubitata depended on food plants outside of the cave during the egg–larva–pupa stages, and they used the cave as a habitat for long‐term survival when they develop into adults.     

中国喀斯特洞穴维管植物多样性

洞穴是喀斯特景观中普遍存在的地质地貌结构, 拥有丰富的植物资源。仅2001-2020年间, 就发表了中国喀斯特洞穴维管植物新属1个; 新种(含种下等级) 170个, 隶属于16科31属, 占近20年发表的中国维管植物新种的3.9%, 说明洞穴植物具有较高的物种多样性潜力。据统计, 中国喀斯特洞穴维管植物有110科339属871种, 主要分布于广西、广东、贵州、云南、重庆等地。物种数排名前5的科包括苦苣苔科、荨麻科、鳞毛蕨科、秋海棠科和茜草科, 其所含物种数共占整个洞穴植物种类的39.4%; 排名前5的属包括报春苣苔属(Primulina)、楼梯草属(Elatostema)、耳蕨属(Polystichum)、秋海棠属(Begonia)和凤尾蕨属(Pteris), 其所含物种数共占整个洞穴植物种类的25.1%。本名录的所有洞穴植物中, 草本习性的种类占一半以上。从洞内分布来看, 仅有10.7%的种类分布于弱光带, 其中石松类和蕨类植物占比远高于被子植物, 说明前者更能适应弱光带的极端环境。不仅如此, 洞穴维管植物也具有较高的特有性成分, 其中省级特有种占比25.8%, 分别包括非洞穴特有种(11.0%)、多洞穴特有种(3.3%)和单洞穴特有种(11.5%)。依据文献显示受威胁种类有115种(占13.0%)。受威胁种类占比较高的类群均是近年来发表新种较多的科, 包括鳞毛蕨科、苦苣苔科、荨麻科、秋海棠科、茜草科等。针对这些类群, 有待更详细地调查来完善对它们的评估。此外, 本名录中至少有36%的种类没有任何分子数据, 亟待补充。     

A new Stygonitocrella Petkovski (Copepoda: Harpacticoida) from a cave in Northern Mexico with comments on the taxonomy of the genus

The cave-dwelling harpacticoid copepod fauna of Mexico is still yet to be known. There are only a few records related to karstic caves, but most refer to epigean forms. The analysis of samples collected in a cave system in Northern Mexico yielded two female specimens of an unknown harpacticoid copepod species which is described herein. It was assigned to the genus Stygonitocrella Petkovski, 1976 mainly because of the presence of a single endopodal segment on legs 1–4; the new species, S. mexicana n. sp. differs from its congeners by a combination of characters related to the segmentation and armature of the swimming legs 1–4, the fifth leg of the female, and the strcture of the antennae. Based on the current limits of Stygonitocrella, two new subgenera are proposed to recognize different morphological patterns based on the presence or absence of an inner baseoendopodal lobe in the female fifth leg, a character not evaluated before. This lobe is absent in the new species and in other five species now contained in the subgenus Eustygonitocrella. A lobe is present in the new subgenus Fiersiella, also with six species. This is the second record of Stygonitocrella in North America, after S. (Fiersiella) sequoyahi Reid, Hunt & Stanley, 2003; it is the only North American cave-dwelling species of the genus. It is suggested that the influence of marine transgressions in Northern Mexico was related to the origin of the new species as part of a group of troglobitic forms originated from marine interstitial ancestors that colonized the freshwater caves. A key for the identification of all the known species of Stygonitocrellais also provided.     

Six new cavernicolous cixiid planthoppers in the genus Solonaima from Australia (Homoptera: Fulgoroidea)

Abstract The monophyletic genus Solonaima , endemic to northern Queensland, is reviewed and six new species are described: S.sullivani sp.n., S.pholetor sp.n., S.stonei sp.n., S.halos sp.n., S.irvini sp.n. and S. baylissa sp.n. The previously known epigean species, S.solonaima Kirkaldy and S.pallesens (Distant) are illustrated and redescribed. The six new species represent four separate independent cave invasions and nearly double the number of known cavernicolous cixiids in the world. This is the first report of a closely related group of cave-dwelling Homoptera showing varying degrees of cave adaptation. One facultative cave species, S.sullivani , shows virtually no modification from surface relatives. Two species, S.pholetor and S.stonei , show some eye, colour, and wing-reduction, but they are still flighted and are considered facultative cave species. S.halos and S. irvini are nearly blind and have reduced, non-functional wings and are considered obligate cave species. S. baylissa , the most remarkable obligate cave species, is highly modified: eyeless, nearly wingless and colourless. The evolutionary trends displayed in cave adaptation are illustrated and discussed. These include reduction in body colour, eyes and wings, alteration of head and thorax shape, and possibly simplification of the male genitalia. The geology of the caves is complex and ranges from greater than 5-million-year-old caves in Silurian-aged metamorphosed limestone to 190,000-year-old lava tubes. The facultative species are generally found in the most open caves, the two intermediate obligate species are found in deeper caves, and the most highly modified species is restricted to damp cave passages with high CO₂ levels. There is no correlation between the ages of the caves and the degree of cave adaptation.     

Predation of a cave fish (Poecilia mexicana, Poeciliidae) by a giant water-bug (Belostoma, Belostomatidae) in a Mexican sulphur cave

Abstract.  1. Caves are often assumed to be predator-free environments for cave fishes. This has been proposed to be a potential benefit of colonising these otherwise harsh environments. In order to test this hypothesis, the predator–prey interaction of a belostomatid (predator) and a cave fish (prey) occurring in the Cueva del Azufre (Tabasco, Mexico) was investigated with two separate experiments.
2. In one experiment, individual Belostoma were given a chance to prey on a cave fish, the cave form of the Atlantic molly ( Poecilia mexicana ), to estimate feeding rates and size-specific prey preferences of the predator. In the other experiment, population density of Belostoma was estimated using a mark–recapture analysis in one of the cave chambers.
3. Belostomatids were found to heavily prey on cave mollies and to exhibit a prey preference for large fish. The mark–recapture analysis revealed a high population density of the heteropterans in the cave.
4. The absence of predators in caves is not a general habitat feature for cave fishes. None the less predation regimes differ strikingly between epigean and hypogean habitats. The prey preference of Belostoma indicates that cave-dwelling P. mexicana experience size-specific predation pressure comparable with surface populations, which may have implications for life-history evolution in this cave fish.     

Local genetic structure in the critically endangered,cave‐associated perennial herb Primulina tabacum (Gesneriaceae)

The local spatial genetic structures of cave‐associated plants are seldom studied. Given that these plants are mainly confined to small areas in and around the entrances of caves, we hypothesized that they might lack genetic structures at local scales. To test this hypothesis, we sampled two large populations (named D and T) of a critically endangered perennial herb, Primulina tabacum, which is endemic to karst caves in southern China. We analysed nine microsatellite loci and sequenced four chloroplast DNA (cpDNA) intergenic spacer regions to study the genetic diversity and structure within and between both populations. Both populations have distinct genetic characteristics. Samples from two subpopulations in population D showed considerable genetic divergence. This is not consistent with the hypothesis that P. tabacum has a weak genetic structure at a local scale. However, 94% of the individuals in population T shared the same multilocus genotype, which indicates little genetic structure within this population. The contributions of seed flow, pollen flow and (sub)population history to the genetic diversity and structure in each and both populations are discussed. Our study is the first to investigate local genetic diversity and structure in a cave‐associated plant, and provides valuable information for the sustainable conservation of such species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 747–756.     

贵州部分洞穴裸灶螽的鉴定及其遗传多样性分析

贵州是中国具有丰富喀斯特洞穴生态系统的省份之一,裸灶螽作为洞穴中的优势物种又为生态环境指示种,其遗传多样性能够反应洞穴生物多样性.为评估洞穴开发对洞穴生物多样性的影响,本研究通过形态鉴定和分子辅助鉴定的方式对来自贵州省毕节、黔西南、铜仁和黔南4个地区8个洞穴97号裸灶螽标本进行鉴定,并对其遗传多样性进行分析.结果共鉴定...     

应用稳定同位素研究广西东方洞食物网结构和营养级关系

稳定性同位素技术已经成为研究生态系统食物网结构和营养级关系及其动态变化的重要手段。应用稳定同位素技术(δ13C和δ15N)研究了东风洞的食物网结构和营养级关系。研究结果表明,东风洞主要为腐食食物链,由于在洞穴内黑暗带中无光照也无植物生长,所以碳源主要为东风洞中的土壤有机质,作为第一营养级;主要摄食关系为马陆与土壤有机物质,螺类与土壤有机物质等;在该洞生态系统中,将3种马陆直接作为初级消费者,以隅蛛和其它马陆平均值之间的差值(2.04‰)作为东风洞食物网稳定氮同位素的富集因子。根据营养级模型可知,马陆类群、细长钻螺等土壤动物,裸灶螽和涂闪夜蛾形成第二营养级,即初级消费者;第三营养级包括蜘蛛类、盲蛛、地蜈蚣以及黑眶蟾蜍,为次级消费者。     

Origins and Relationship of Cave Populations of the Blind Mexican Tetra, Astyanax Fasciatus, in the Sierra De El Abra

The blind morph of Astyanax fasciatus (Pisces: Characidae) has been more thoroughly studied than any other cave inhabiting organism. Most studies of A. fasciatus have used individuals from different caves of the Sierra de El Abra, Mexico, and have assumed that each population independently evolved to live in the cave environment. We analyzed the relationships among several cave populations that delineate the Sierra de El Abra using RAPD markers. The results indicate that all cave populations are more closely related to one another than they are to the surface populations. This suggests that present day cave populations derived from a common ancestral stock, most likely due to a single colonization event, or alternatively, that strong gene flow among cave populations has occurred, raising precaution against considering each cave population as independent.     

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.