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.     

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.     

Locomotor Activity Rhythms in Cave Catfishes, Genus Taunayia, from Eastern Brazil (Teleostei: Siluriformes: Heptapterinae)

We studied the locomotor rhythmicity in heptapterine catfishes, genus Taunayia, under free-running conditions (DD) and LD cycles (12:12). Taunayia sp., anophthalmic and depigmented undescribed species from a cave in northeastern Brazil, is the fourth Brazilian troglobitic catfish studied with focus on circadian rhythms. Weak free-running rhythmicity, with absence of significant circadian components, was observed for this species when compared to the epigean, eyed relatives. On the other hand, the studied troglobitic catfishes in general presented significant circadian rhythms under LD cycles, with activity peaks in the night phase probably corresponding to nocturnal activity pattern inherited from their epigean ancestors. However, no residual oscillations were observed after transition from LD to DD. This indicates masking of activity by light-dark cycles. Regression of circadian rhythmicity in the stable, permanently dark subterranean habitat was also observed for other cave fishes. Such regression corroborates the notion that circadian rhythmicity is mainly selected in the epigean environment by ecological factors, namely daily cycles of light and/or temperature.     

Locomotor Activity Rhythms in Cave Catfishes,Genus Taunayia,from Eastern Brazil (Teleostei: Siluriformes: Heptapterinae)

We studied the locomotor rhythmicity in heptapterine catfishes, genus Taunayia, under free-running conditions (DD) and LD cycles (12:12). Taunayia sp., anophthalmic and depigmented undescribed species from a cave in northeastern Brazil, is the fourth Brazilian troglobitic catfish studied with focus on circadian rhythms. Weak free-running rhythmicity, with absence of significant circadian components, was observed for this species when compared to the epigean, eyed relatives. On the other hand, the studied troglobitic catfishes in general presented significant circadian rhythms under LD cycles, with activity peaks in the night phase probably corresponding to nocturnal activity pattern inherited from their epigean ancestors. However, no residual oscillations were observed after transition from LD to DD. This indicates masking of activity by light-dark cycles. Regression of circadian rhythmicity in the stable, permanently dark subterranean habitat was also observed for other cave fishes. Such regression corroborates the notion that circadian rhythmicity is mainly selected in the epigean environment by ecological factors, namely daily cycles of light and/or temperature.     

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.     

Cave adaptation in the spider genus Anthrobia (Araneae, Linyphiidae, Erigoninae)

The genus Anthrobia Tellkampf, 1844 is revised and expanded to include two epigean species, Anthrobia acuminata ( Emerton, 1913 ) comb. nov., and A. whiteleyae sp. nov., and two troglobites, the type species A. monmouthia Tellkampf, 1844 , and A. coylei sp. nov. The female of A. acuminata is described for the first time. All four Anthrobia species were added to a phylogenetic analysis of erigonine genera. Analysis of this matrix results in a single most parsimonious tree (length = 915, CI = 0.23, RI = 0.58; uninformative characters excluded: length = 911, CI = 0.23), which places Anthrobia sister to Diplocentria Hull, 1911. Troglobitic Anthrobia and epigean Anthrobia are both monophyletic. In comparison to the epigean species, troglobitic Anthrobia exhibit the following putative adaptations to cave life: loss of eyes, elongation of the legs, and reduction of the tracheal system. The reliability of phylogenetic results indicating monophyly of cave Anthrobia are discussed in light of the fact that potentially cave adaptive synapomorphies support this relationship.     

Developmental constraints in regressive evolution: studies of the expression of the γs-crystallin gene in the developing lens of cave-dwelling Astyanax fasciatus (Cuvier, 1819) (Teleostei, Characidae) by in situ hybridization

The early morphogenesis of the lens and the expression of the γs-crystallin gene was studied in epigean Astyanax fasciatus and its cave-dwelling derivative. At early stages, the lens of the cave fish develops in a way that is similar to the epigean form. Later, the developmental timing is delayed and growth ceases in the cave-fish lens. With the beginning of cytodifferentiation, the development of the lens breaks down. Crystallin lens fibres are not produced at any time and the γs-crystallin gene, which is transcribed during a limited period in the lens of epigean fishes, is not active in cave specimens. This study confirms earlier immunofluorescence observations that demonstrated the lack of crystallin proteins in the cave-fish lens, but is in contrast to results on the blind mole rat, which showed a persistence of functioning crystallins in the degenerated lens of this species. The significance of developmental constraints in regressive evolution is discussed.     

Troglomorphic Sculpins of the Cottus Carolinae Species Group in Perry County, Missouri: Distribution, External Morphology, and Conservation Status

The existence of cavernicolous sculpin (here allocated to Cottus carolinae, banded sculpin, and referred to as grotto sculpin), in the karst regions of Perry County, Missouri, first came to our attention in 1991. Examination of 35 caves in Missouri, 96 in Illinois, 17 in Tennessee, two in Indiana, and 11 in Arkansas revealed that banded sculpin are common in cave habitats; however, grotto sculpin are limited to two karst areas of Perry County, Missouri, where they are known from only six cave systems. These caves and their streams are extensive and apparently provide a unique habitat compared to other karst systems; this may be a critical factor in the present restricted distribution of the grotto sculpin. Grotto sculpin occupy pools and riffles of cave streams, and occur over a variety of substrates, from sediment to breakdown. Density estimates in Mystery and Running Bull caves were 0.29 and 0.63 individuals m^-2, respectively. Grotto sculpin have small eyes (1–6% SL vs. 6–10% SL in epigean samples), significantly reduced pigmentation (including nearly complete loss of dorsal saddles), a reduction in pelvic fin ray number (from 4+4 elements to often 4+3 , or 3+3), and enlarged cephalic lateralis pores (e.g., mandibular pores of cavernicolous samples are 2–3 times those of epigean stream samples). Multivariate analyses of body shape revealed statistically significant separation of epigean and hypogean samples, with eye size highly variable, but smallest in the Running Bull Cave population. We interpret these results as representative of losses associated with long-term cave habitation. Caves of Perry County provide ample habitat for grotto sculpin, but because the caves are located downgradient of the city of Perryville and an intensively farmed landscape, point and non-point source pollution threaten their continued existence. Escape of farm-pond fishes through the extensive sinkhole network in Perry County has increased potential predation pressure on grotto sculpin by channel catfish, Ictalurus punctatus, and other species normally excluded from cave environments.     

Leptogenys khammouanensis sp. nov. (Hymenoptera: Formicidae). A possible troglobitic species of Laos,with a discussion on cave ants

The new species Leptogenys khammouanensis sp. nov. is described from two caves of the Khammouan karst (Laos). It is characterized by a set of striking morphological characters (reduced eyes, light pigmentation, slender body and very elongated legs and antennae), which recall the troglobiomorphic traits of cave arthropods. Relations between caves and ants are discussed at this occasion, in the light of the recent biological explorations of caves in Southeast Asia. The classical view that ants are rare and unimportant in caves is challenged. Ants are actually major and regular components of guano assemblages in many caves of the region, but none of these guano species exhibit cave-related adaptation in its external morphology. Conversely, ants are very rare in low-resources habitats, where only accidental occurrence of outside species are reported in Southeast Asia. Leptogenys khammouanensis has been found only in such an oligotrophic environment, very deep in the cave and far from any guano deposits. Its presence there, together with its troglobiomorphic traits, support the idea that Leptogenys khammouanensis might be the first truly troglobitic ant.     

One eye but no vision: cave fish with induced eyes do not respond to light

One of the most intriguing questions in evolutionary biology is the degree to which behavior is a necessary consequence of morphology. We explore this issue by examining phototactic behavior in epigean (eyed surface-dwelling) and troglomorphic (blind cave) forms of the teleost Astyanax fasciatus whose eyes were modified during embryogenesis by removing one or both lens vesicles from the epigean form or by transplanting the lens vesicle from an epigean fish into the optic cup of a blind cave form. Lens removal results in eye degeneration and blindness in adult epigean fish, whereas lens transplantation stimulates growth of the eye, inducing the development of optic tissues in the normally eyeless adult cave fish. Photoresponsiveness was examined by placing fish in an aquarium with one half illuminated and the other half dark and scoring their presence in the illuminated or dark half. Both the eyeless epigean fish and cave fish with induced eyes are indifferent to the illumination whereas the surface forms are scotophilic, suggesting that optic development and phototactic behavior are decoupled.     

Reaction to Alarm Substance in Cave Populations of Astyanax fasciatus (Characidae,Pisces)1

The effect of species-specific skin-extract containing alarm substance has been studied in two epigean and three hypogean populations of the Mexican-tetra Astyanax fasciatus. In all populations tested the feeding rate at the water surface decreased significantly after introducing the alarm substance and the fish avoided this surface area. Additionally epigean fish and some individuals from the Chica cave showed zigzag movements, rapid swimming and hiding. The fish did not respond to a skinextract of a species without alarm substance.     

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.     

Non-optical releasers for aggressive behavior in blind and blinded Astyanax (Teleostei, Characidae)

Prior to this study, it was believed that epigean and hypogean Astyanax differ markedly in their display of agonistic behavior. Research suggested that surface-dwelling individuals were extremely aggressive whereas their blind, cave-dwelling counterparts tended to show little or no aggressive behavior. Aggression in Astyanax was thought to be triggered by visual stimuli because surface fish in a dark environment or surface fish blinded late in life did not show aggression. Here, we demonstrate that surface fish blinded early on in their embryonic development are highly aggressive as adults. We also report the first case of a population of blind cave-dwelling Astyanax that is highly aggressive. We conclude that reduced aggression is not the only evolutionary pathway for troglobitic Astyanax and that there is some degree of developmental plasticity in the releaser of aggression and in the selection of its triggering stimuli.     

Fish Assemblages of Mediterranean Marine Caves

Fish assemblages associated with 14 marine caves and adjacent external rocky reefs were investigated at four Marine Protected Areas (MPAs) along the coasts of Italy. Within the caves sampling was carried out in different sub-habitats: walls, ceilings, bottoms and ends of caves. On the whole, 38 species were recorded inside the 14 caves investigated. Eighteen species were exclusively found inside the caves: they were mainly represented by speleophilic (i.e. species preferentially or exclusively inhabiting caves) gobids (e.g. Didogobius splechtnai) and nocturnal species (e.g. Conger conger). Forty-one species were censused outside, 20 of which were shared with cave habitats. Apogon imberbis was the most common fish found in all 14 caves investigated, followed by Thorogobius ephippiatus (recorded in 13 caves), and Diplodus vulgaris and Scorpaena notata (both censused in 12 caves). Distinct fish assemblages were found between external rocky reefs and the different cave sub-habitats. New data on the distribution of some speleophilic gobids were collected, showing the existence of a pool of species shared by marine caves on a large scale (i.e. hundreds of km). Considering the uniqueness of cave fishes (18 exclusive species and different assemblage structures), the inclusion of marine caves among the habitats routinely investigated for fish biodiversity monitoring could facilitate the achievement of more comprehensive inventories. Due to their contribution to local species diversity and the shelter they provide to species valuable for conservation, marine caves should be prioritized for their inclusion not only within future MPAs through the Mediterranean Sea, but also into larger management spatial planning.     

Free-Running Locomotor Activity Rhythms in Cave-Dwelling Catfishes,Trichomycterus sp., from Brazil (Teleostei,Siluriformes)

Locomotor activity of troglobitic (cave restricted) catfishes, Trichomycterus sp., was recorded in the laboratory under constant darkness for ten consecutive days. Infra-red photocell beam crossings were totalled every thirty minutes and stored for later analysis. Spectral analysis of the data followed by a statistical test designed for the detection of significant components was then performed. The results of 14 individuals, seven pigmented and seven albino fishes, are reported here. In eight individuals, four pigmented and four albinos, we were not able to detect significant components in the circadian range, although ultradian and/or infradian significant components were found in all cases. Former studies on other two Brazilian cave catfishes, Pimelodella kronei and Imparfinis sp., showed similar results, suggesting a picture of gradual loss of circadian rhythmicity and persistence of ultradian and infradian rhythms in the locomotor activity of troglobitic populations. The proportion of specimens without significant circadian components was higher in Trichomycterus sp., supposedly less specialized to cave life than the other two species. This is tentatively correlated with a higher environmental stability during the time of evolutionary isolation in the subterranean biotope. These results provide strong arguments for the importance of external selection over circadian rhythmicity.     

Cave dwelling cockroaches from Sarawak, with one new species

Abstract. The troglobitic cockroach, Trogoblattella chapmani , sp.n., is described. The genus was known previously from a single species found in limestone caves in Australia. Two of three other species found in the Sarawak caves, namely Symploce cavemicola (Shelford) and Pycnoscelus striata (Kirby) have previously been reported from cave habitats. The third, Pycnoscelus indicus (Fabricius), may have been reported previously from caves, under the name of Pycnoscelus surinamensis (Linnaeus). A single undetermined female of Blattella was present in the collection.     

Free-Running Locomotor Activity Rhythms in Cave-Dwelling Catfishes, Trichomycterus sp., from Brazil (Teleostei, Siluriformes)

Locomotor activity of troglobitic (cave restricted) catfishes, Trichomycterus sp., was recorded in the laboratory under constant darkness for ten consecutive days. Infra-red photocell beam crossings were totalled every thirty minutes and stored for later analysis. Spectral analysis of the data followed by a statistical test designed for the detection of significant components was then performed. The results of 14 individuals, seven pigmented and seven albino fishes, are reported here. In eight individuals, four pigmented and four albinos, we were not able to detect significant components in the circadian range, although ultradian and/or infradian significant components were found in all cases. Former studies on other two Brazilian cave catfishes, Pimelodella kronei and Imparfinis sp., showed similar results, suggesting a picture of gradual loss of circadian rhythmicity and persistence of ultradian and infradian rhythms in the locomotor activity of troglobitic populations. The proportion of specimens without significant circadian components was higher in Trichomycterus sp., supposedly less specialized to cave life than the other two species. This is tentatively correlated with a higher environmental stability during the time of evolutionary isolation in the subterranean biotope. These results provide strong arguments for the importance of external selection over circadian rhythmicity.     

Convergent Adaptations to Cave Life in the Rhamdia Laticauda Catfish Group (Pimelodidae, Teleostei)

The nocturnal Mexican catfish Rhamdia laticauda (Pimelodidae, Teleostei) is the surface sister species of a number of cave species. Comparison between two of them, R. zongolicensis and R. reddelli, reveals an intermediate state of reduction of the biologically functionless eyes, melanin pigmentation and the negative phototactic behavior. The surface species is perfectly adapted to life in darkness. Therefore only few constructive adaptations are developed in the cave species. For survival under conditions of low food supply in the caves the barbels are elongated to improve the senses of taste and touch and more fat can be deposited in the cave fish tissue. Due to convergent evolution the two cave species are morphologically much alike and show only minor diagnostic meristic differences. From geological data it can be concluded that the two cave species started troglobitic evolution at the end of Pleistocene.