Multiple cave invasions by species of the planthopper genus Oliarus in Hawaii (Homoptera: Fulgoroidea: Cixiidae)

Although geologically young, the Hawaiian Islands harbour a rich and remarkably diverse fauna of terrestrial troglobites: more than 70 cave species are known from Kauai, Oahu, Molokai, East Maui, and Hawaii Island. Among the more speciosc groups that have invaded the subterranean biome are the planthoppers (Homoptera Cixiidae) of the genus Oliarus. Five new obligately cavernicolous (troglobitic) Oliarus species which differ in their degree of troglomorphy and male genital structures, are described from lava tubes on the Hawaiian Islands: O. lorettae sp. nov. and O. makaiki sp. nov. from Hawaii Island, O. gagnei sp. nov. and O. ivaikau sp. nov. from Maui Island, and O. kalaupapae sp. nov. from Molokai Island. Short diagnoses of the two troglobitic species already known, 0. polyphemus Fennah, 1973 from Hawaii Island and 0. priola Fennah, 1973 from Maui are provided. Notes on the ecology and distribution of all cavernicolous species are given. Morphological evidence suggests that each of the seven cavernicolous Oliams species from Hawaii represents a separate, independent adaptive shift to underground environments. Potential relationships to the extant epigean species are discussed.     

Comparative Density of Hair Sensilla on the Legs of Cavernicolous and Epigean Harvestmen (Arachnida: Opiliones)

To allow an animal to behave appropriately, the location of sensorial structures is expected to be related to their function. As the different leg pairs of arachnids may have different functions (probing x supporting the body), one could expect them to have a different density of sensilla. Moreover, different regions of the same leg (dorsal, lateral, and ventral) would also be expected to have different densities of sensilla, according to the use of each region (e.g., the ventral part is often in contact with the substrate while the dorsal part is not). As cavernicolous animals are expected to be more sensitive than their epigean relatives, one could also expect a different density of sensilla when comparing cavernicolous and epigean animals. Using three epigean and three cavernicolous species of harvestmen (Arachnida, Opiliones), this study aimed at describing the morphology of hair sensilla on the legs and answering three questions: (1) Are there differences in the density of hair sensilla between the dorsal, lateral and ventral regions of each leg pair of the same individual? (2) Are there differences in the density of hair sensilla between the leg pairs of the same individual? (3) Are there differences in the density of hair sensilla when comparing the leg pairs of individuals of cavernicolous and non-cavernicolous species? The tarsi and metatarsi of all right legs of the six studied species were analyzed under a scanning electron microscope. The results (P < 0.05) showed that, in general: the ventral region of the tarsus was denser in sensilla trichodea than the lateral and dorsal regions, particularly on legs I and II; the density of sensilla chaetica did not differ on legs III and IV, but was greater on the dorsal region of legs I and II; the ventral part of legs I had the higher density of sensilla trichodea of the four pairs, whereas the second pair had the lower density; Holcobunus citrinus (Eupnoi) was the species with higher density of sensilla trichodea, on all legs; the cavernicolous species had a lower density of sensilla than the epigean species. The results are tentatively related to harvestmen behavior.     

Freshwater triclads (Turbellaria, Tricladida) from the oriental region

Two new freshwater triclad species of the genus Dugesia Girard, 1850 are described from the oriental region. The first is the only species of freshwater triclad described from Ceylon and the second, a cavernicolous form, is the only one known from Malaya. A synopsis, in tabular form, of the oriental Dugesia species is also given and some aspects of their distribution are discussed. It is intended that the tabular synopsis should provide diagnostic characters for the delimitation of the new and previously described species, and act as a basis for a future revision of the oriental representatives of the genus, many of which are poorly described or poorly known. Apreliminary dichotomous key to the valid species is also provided.     

Danielopolina kornickeri sp. n. (Ostracoda, Thaumatocypridoidea) from a western Australian anchialine cave: morphology and evolution

Danielopolina kornickeri sp. n . is described from an anchialine cave in Western Australia. This is the first Danielopolina species occurring outside of the Western Hemisphere (Atlantic and Eastern Pacific Oceans and Islands). A cladistic analysis indicates that this species is a primitive taxon of the anchialine cavernicolous clade; the deep-sea dwelling species Danielopolina carolynae appears more primitive and does not cluster with the other Danielopolina species. The ecological environment of D. kornickeri is described. Detailed discussions consider: (1) the shell ornamentation and its relevance to the phylogeny of the Thaumatocyprididae; (2) the antero-dorsal node of the Permian Thaumatomma (hypothesized here to be an ocular lens which regressed in the post-Palaeozoic thaumatocypridids); (3) the distal chaetotaxy of the antennae (hypothesized to perform the chemosensorial function known in the aesthetascs of other myodocopids or podocopids); (4) palaeobiogeographical arguments for a shallow marine origin of cavernicolous species Danielopolina .     

EVOLUTION OF ADULT MORPHOLOGY AND LIFE-HISTORY CHARACTERS IN CAVERNICOLOUS PTOMAPHAGUS BEETLES

Morphological and life-history characters were determined for a series of six increasingly cavernicolous species of eastern United States Ptomaphagus beetles. Contrary to expectations, dimensions of only some adult structural characters uniformly covary as an overall measure of evolutionary adaptation for cave life. Adult reproductive characters that show adaptation for cave life are loss of reproductive seasonality and production of fewer and larger eggs. Significant change in pre-imaginal life-cycle stages was not found. This is in contrast to cavernicolous bathysciine beetles of Europe which show remarkable adaptive trends in pre-imaginal stages, but larval adaptations are not strongly coevolved with adult cave-adaptive characters. This suggests that evolution of cave adaptation in adult endopterygote insects occurs before and independently from that in larvae.     

Troglobitic scorpions: a new genus and species from Brazil

A new genus and species of troglobitic buthid scorpion are described on the basis of a single specimen collected in Brazil. This is the first cavernicolous scorpion ever found in Brazil, and only the second to be found in South America. Some considerations on troglobitic scorpions are proposed.     

First record of troglobitic Hemiptera (Fulgoromorpha: Cixiidae) from La Réunion Island

A new species of troglobitic Cixiidae (Hemiptera: Fulgoromorpha), Brixia briali n. sp. is described from a lava tube on La Réunion Island. Information on its ecology, distribution, and comments on its tentative systematic placement into the genus Brixia is provided. The new discovery from La Réunion is placed into the context of cavernicolous Fulgoromorpha of the world, their zoogeography, ecology and biology.     

Time, area and isolation: factors driving the diversification of Azorean arthropods

Aim R. J. Whittaker et al. recently proposed a ‘general dynamic model of oceanic island biogeography’ (GDM), providing a general explanation of island biodiversity patterns by relating fundamental biogeographical processes – speciation, immigration, extinction – to area (A) and time (T; maximum island geological age). We adapt their model, which predicts a positive relationship with area combined with a humped relationship to time (designated the ATT² model), to study the factors promoting diversification on the Azores for several arthropod groups. Location The Azorean archipelago (North Atlantic; 37–40° N, 25–31° W). Methods We use the number of single‐island endemics (SIEs) as a measure of diversification, to evaluate four different predictions for the variation in SIEs between different islands, derived from the GDM theory and our knowledge of the fauna and history of the Azores. We calculated the number of SIEs for seven out of the nine Azorean islands and six groups of species (all arthropods, beetles, cavernicolous and non‐cavernicolous species, and taxa with high and low dispersal abilities). Several variables accounting for island characteristics (area, geological age, habitat diversity and isolation) and generalized linear models were used to evaluate the reliability of each prediction. Results A linear and positive relationship between SIEs and an AT (area + time) model was the most parsimonious explanation for overall arthropod diversification. However, cavernicolous species showed the opposite pattern (more SIEs inhabiting the youngest islands). Also, isolation was an important predictor of diversification for all groups except for the species with high dispersal ability; while the former were negatively related to the distance from the main source of colonizing lineages (Santa Maria island in most cases), the latter were related to area. Dispersal ability was also a key factor affecting the diversification of most groups of species. Main conclusions In general, the diversification of Azorean arthropods is affected by age, area and isolation. However, different groups are affected by these factors in different ways, showing radically different patterns. Although the ATT² model fails to predict the diversification pattern of several groups, it provides a framework for integrating these deviations into a general theory. Further improvements of the GDM theory need to take into account the particular traits of each group and the role of isolation in shaping island diversity.     

A new cave-dwelling and endemic species of the genus Pholeuonopsis (Coleoptera, Leiodidae) from Serbia

In the present study, a new species of a cavernicolous beetle is described from Serbia: Pholeuonopsis (Pholeuonopsis) zlatiborensis sp. n. (Leptodirini, Cholevinae, Leiodidae). All important taxonomic features have been thoroughly analyzed and illustrated. The new species probably belongs to an old phyletic lineage of mesogeid origin, like other known Pholeuonopsis species inhabiting cave and endogean habitats in the western part of the Balkan Peninsula. Therefore, the species is both relict and endemic inhabitant of cave habitats in Serbia.     

西双版纳地区犬蝠和棕果蝠食性的初步研究

犬蝠和棕果蝠是西双版纳地区较为常见的两个果蝠物种,大多数时间内它们同域分布,共同利用当地许多野生果实。从2004年6月至12月,我们采用拾遗法、粪便分析法以及种子萌发鉴定法并结合雾网采样对西双版纳地区这两种果蝠的食性进行了初步研究。结果发现犬蝠利用11科18种植物的果实,2科2种植物的叶片;而棕果蝠利用9科12种植物的果实,1科1种植物的叶片。研究发现雨季(6－10月)两种果蝠食物类型在本地区重叠程度较高,它们共同利用的植物类型占记录植物类型总数的65%。在干旱季节(11－12月),棕果蝠避开与犬蝠在食物方面的竞争而去别的地力开拓食物资源。     

Phylogenetic evidence for multiple invasions and speciation in caves: the Australian planthopper genus Solonaima (Hemiptera: Fulgoromorpha: Cixiidae)

Abstract. The Australian genus Solonaima comprises thirteen described plus two undescribed species. Six are cavernicolous, obligate or not, and are found in different caves. The phylogeny presented here confirms the monophyly of the genus. This phylogeny was compared with the estimate obtained using the method of Marques and Gnaspini, who recommend coding characters susceptible to parallelism differently from the others. Further comparison was made with a cladogram derived from the matrix from which such characters susceptible to parallelism were withdrawn. Scenarios concerning historical invasions of caves were tested using phylogenetic inference. The most-parsimonious hypothesis proposed four invasions of the caves, within two of which a diversification of species took place.     

Test for Bioenergetic Progress and Specific Energy Metabolism in Isopod Crustaceans (Isopoda) of Various Ecology

We studied energy metabolism of terrestrial and cavernicolous isopods and demonstrated much lower standard metabolism in the troglobionts as compared to other Isopoda representatives. The test for bioenergetic progress proved to be applicable for both aromorphosis and katamorphosis. Different patterns of the relationship between energy metabolism and temperature in stenothermal and eurythermal species have been proposed.     

Morphometric analysis of interspecific and microgeographic variation of crayfish from a Mexican cave

Two undescribed species of Procambarus crayfish (Crustacea, Decapoda) inhabit the subterranean stream running in the recently explored Cueva de Los Camarones, Chiapas, Mexico. Individuals ranging from dark, robust, large-eyed forms with an epigean mien to pale, comparatively slender, microphthalmic ones resembling related troglobites were found, but only the extremes could be distinguished visually. Evolutionary relationships among individuals were investigated morphometrically by means of multivariate analysis'on 14 body dimensions. These analyses revealed a significant discontinuity between the two species, mainly determined by the shape of the rostrum, chelae and telson. The occurrence of a few aberrant phenotypes may indicate the existence of some gene flow between the two species.
Microgeographic variation was demonstrated within the less cavernicolous species. The characters involved were the same as those that discriminate between the two species. These results are compared with allozymic data presented in a parallel paper and the role of genie introgression and environmental induction are discussed.     

A new species of Thetispelecaris (Crustacea: Peracarida) from submarine cave on Grand Cayman Island

A new species of the peracaridan order Bochusacea, Thetispelecaris yurikago, is described from a submarine cave on Grand Cayman Island, the Caribbean Sea. The new species is the fourth species of the order and family, and the second of the genus. Recent studies have strongly suggested a close phylogenetic affinity between cave-dwelling and deep-sea taxa in the Bochusacea as recognized in other cavernicolous/deep-sea crustaceans such as amphipods and copepods. The morphology of the gut and female reproductive system is observed for the first time in the Bochusacea: the stomach is complex with structures such as ridges, processes, spinules, and hairs in the lumen; paired gonopores are located near the base of the fifth pereiopods on the sternite.     

Exoskeletal cuticle of cavernicolous and epigean terrestrial isopods: A review and perspectives

Comparative ultrastructural studies of the integument in terrestrial isopod crustaceans show that specific environmental adaptations of different eco-morphotypes are reflected in cuticle structure. The biphasic molting in isopods is a valuable experimental model for studies of cuticular matrix secretion and degradation in the same animal. The aim of this review is to show structural and functional adaptations of the tergal cuticle in terrestrial isopods inhabiting cave habitats. Exoskeletal cuticle thickness, the number of cuticular layers, epicuticle structure, mineralization, pigmentation and complexity of sensory structures are compared, with greater focus on the well-studied cave trichoniscid Titanethes albus. A large number of thinner cuticular layers in cave isopods compared to fewer thicker cuticular layers in related epigean species of similar body-sizes is explained as a specific adaptation to the cavernicolous life style. The epicuticle structure and composition are compared in relation to their potential waterproofing capacity in different environments. Cuticle mineralization is described from the functional point of view as well as from the aspect of different calcium storage sites and calcium dynamics during the molt cycle. We also discuss the nature and reduction of pigmentation in the cave environment and outline perspectives for future research.     

龙眼荔枝卷叶蛾类寄主植物种类的初步研究

对广西龙眼荔枝卷叶蛾类寄主植物种类的初步考查结果表明,目前所采到的卷叶蛾种类中,它们的寄主范围不尽相同。柑桔褐带卷蛾Adoxophyes cyrtosema达15科31种;柑桔长卷蛾Homona coffearia和棉褐带卷蛾A.orana的寄主分别有14科21种和13科20种;后黄卷蛾Cacoecia asiatica和柑桔黄卷蛾Archips seminubibis的寄主有10科14种和8科13种;三角新小卷蛾Olethreutes leucaspis、圆角卷蛾Ebodocellerigera和斜斑小卷蛾Andrioplecta oxystaura,目前仅发现危害龙眼及荔枝。新发现柑桔长卷蛾可危害翠蜜枣、指天椒和皱果苋,柑桔黄卷蛾危害粪箕笃、苦丁茶、指天椒,柑桔褐带卷蛾可危害黑面神(Breyniafruticosa)。     

Rhabditophanes schneideri (Rhabditida) phoretic on a cave pseudoscorpion

Information of phoretic nematode-pseudoscorpion associations and cases of parasitism on five European species of pseudoscorpions was summarized by Cur?ic et al. [Curci?, B.P.M., Dimitrijevi?, R.N., Makarov, S.E., Luci?, L.R., Curci?, S.B., 1996. Further report on nematode-pseudoscorpion associations. Acta arachnol. 45, 43-46; Curci?, B.P.M., Sudhaus, W., Dimitrijevi?, R.N., Tomi?, V.T., Curci?, S.B., 2004. Phoresy of Rhabditophanes schneideri (Bütschli) (Rhabditida: Alloionematidae) on pseudoscorpions (Arachnida: Pseudoscorpiones). Nematology 6 (3), 313-317]. An examination of a sample of the cavernicolous pseudoscorpion Neobisium rajkodimitrijevici (Curci? and Tomi?, 2006) (comprising a holotype male and a paratype tritonymph) from a cave in Eastern Serbia revealed this false scorpion to be a nematode carrier; the present paper reports this finding and extends our knowledge of phoresy of Rhabditophanes on pseudoscorpions. This is the first time that the rhabditid R. schneideri (Bütschli, 1873) has been noted in association with a cavernicolous pseudoscorpion. There must be some patchily distributed micro-habitats in caves where saprobiotic nematodes and small arthropods can complete their life-cycles, for example something like deposits of bat guano. The transportation of Rhabditophanes J3 by pseudoscorpions indicate that Neobisium specimens often visit these micro-habitats to find their prey.     

Zur genetischen interpretation morphologischer gesetzmäßigkeiten der degenerativen evolution

Complex organs such as the eyes of vertebrates are based upon highly polygenic systems. So, remarkable differences in the eyes of cave fishes and their overground ancestors are caused by several mutations. Although in cavernicolous populations natural selection seems to play no part in eye degeneration, there is evidence for morphologic regularities in the degenerative process. These morphologic regularities are due to pleiotropic effects of the mutating genes.