V. Sbordoni

GENETIC VARIABILITY AND DIVERGENCE IN CAVE POPULATIONS OF TROGLOPHILUS CAVICOLA AND T. ANDREINII (ORTHOPTERA, RHAPHIDOPHORIDAE)

This paper reports initial data on genetic variability and genetic divergence in five populations of two cave dwelling orthopterans, genus Troglophilus. Most of the present literature concerning electrophoretic variation of cavernicoles reports low levels of genetic variability, e.g., cave populations of fishes (Avise and Selander, 1972), millipedes (Laing et al., 1976a), leiodid beetles (Laing et al., 1976b) and spiders (Johnston and Carmody, 1977). However several recent reports (Sbordoni et al., 1976a, 1978, 1979, 1980a, 1980b; Giuseffi et al., 1978; Dickson et al., 1979; Sbordoni, 1980) and additional unpublished data from our own genetical analyses on different cavernicolous animals, indicate that troglophilic and troglobitic populations of aquatic and terrestrial invertebrates subjected to long term cave isolation may display levels of genetic variability comparable to, or even higher than, the levels observed in non-cave invertebrates (Powell, 1975; Selander, 1976). As far as cave crickets (Rhaphidophoridae) are concerned, Cockley et al. (1977) report low variability in the camelback cave cricket Ceuthophilus gracilipes from Pennsylvania, with heterozygosity values ranging, in different populations, from 0.020 to 0.033. In contrast to these results our preliminary data on Dolichopoda geniculata (Cobolli Sbordoni et al., 1974; Sbordoni et al., 1976b) and a further extensive study of 23 populations belonging to five species of Dolichopoda from Central Italy indicate a fairly high degree of variability, with levels of heterozygosity ranging from 0.113 to 0.280 (Sbordoni et al., unpubl.). Mediterranean Troglophilus cave crickets are similar to American Ceuthophilus. Both are Rhaphidophoridae and even if not closely related (Hubbell and Norton, 1978), occupy parallel niches in the two continents, most of the species being adapted to the twilight zones of caves, feeding on living arthropods, guano and chlorophyllan plants, sometimes outside caves (Remy, 1931; Vandel, 1964). The range of the genus Troglophilus extends from Anatolia to Lombardy including Greece, Yugoslavia and Austria. In Italy this genus shows a disjunct distribution: two species, T. cavicola (Kollar) and T. neglectus Krauss, are limited to limestone areas along the foothills of the Alps in northern Italy, from Friuli west to pre-alpine Lombardy, namely the Lake Como region, where marginal parthenogenetic populations of T. cavicola occur (Capra, 1951; Baccetti, 1961). A third species, T. andreinii Capra, is present in Apulia where two subspecies have been identified: T. andreinii andreinii, which is widespread in caves of northwestern Apulia, and T. andreinii hydruntinus La Greca, confined to the southern edge of the Salentine Peninsula (Baccetti and Capra, 1969). We investigated the patterns of genetic variability in five Troglophilus populations, two belonging to T. cavicola from northern Italy, and three belonging to T. andreinii from Apulia. We attempted to evaluate 1) levels of heterozygosity in Troglophilus to compare them with both Ceuthophilus and Dolichopoda and 2) the degree of genetic divergence between populations and species. The populations

Genetic variability and divergence between populations and species of Nesticus cave spiders

Genetic variability and divergence at 21 enzyme loci were studied in and between Italian populations of the cave spiders Nesticus eremita (13 populations), N. menozzii and N. sbordonii (one population each). The three species differ with respect to the degree of specialization to cave life, dispersion ability, isolation of populations, abundancy, extent of the distribution area, and range from the troglophilic and widespread N. eremita to the troglobitic N. sbordonii, endemic to a single cave in the Central Appennines.Heterozygosity ranges from 0.05 to 0.15 in N. eremita populations and appears to be largely controlled by the occurrence and the extent of gene flow among populations. The relatively low polymorphism levels of N. menozzii (H=0.081) and N. sbordonii (H=0.106) are also associated with reduced gene flow and small population sizes.Genetic distances between N. eremita populations vary considerably and are strictly related to the geographical distances involved, again indicating a major role of gene flow in determining the patterns of genetic differentiation between populations. This view is strongly supported by the results of a principal component analysis applied to the gene-frequency data. Estimates of genetic divergence between species suggest that the major cladogenetic events leading to complete separation of these three Nesticus species occurred in the Middle-late Pliocene.

Biochemical divergence between cavernicolous and marine Sphaeromidae and the Mediterranean salinity crisis

Allozymic variation in proteins encoded by 14 loci was analyzed in 3 cavernicolousMonolistra and in 2 marineSphaeroma species. Genetic distance data, high levels of heterozygosity and the divergence time calculations support the hypothesis thatMonolistra diverged from its Sphaeroma-like marine ancestor during the Messinian, in connection with the Mediterranean salinity critis.

Genetic variability and divergence between cave dwelling populations of Typhlocirolana from majorca and sicily

The genus Typhlocirolana includes six species discontinuously distributed in underground freshwater systems throughout southern Mediterranean coasts. The recent discovery of a Sicilian population of Typhlocirolana morphologically close to T. moraguesi from the Balearic Islands raised two issues: the taxonomic status of the Sicilian population and the evolutionary history leading to the present distribution of Typhlocirolana species which are thought to descend from a Cirolana-like marine ancestor. Electrophoretic analysis of eighteen gene loci encoding for enzymes and other proteins revealed a consistently high value of genetic distance (D=0.815) between the Sicilian sample and the two Majorcan population samples, strongly suggesting their status as separate species. Both genetic distance data and the low levels of heterozygosity detected in all samples suggest a relatively recent age for the Typhlocirolana species so far examined. Results from this study support the hypothesis by Stock according to which Typhlocirolana, as well as other ‘thalassoid’ hypogean Crustaceans, colonized ground-waters in connection with the sedimentary cycles of the Mediterranean coastal lines during the Upper Miocene or Lower Pliocene.

Divergenza genetica tra popolazioni e specie ipogee ed epigee di Niphargus (Crustacea, Amphipoda)

VALERIO SBORDONI, MARINA COBOLLI SBORDONI e ELVIRA DE MATTHAEIS Istituto di Zoologia dell’Universitz‘1 di Roma Dive-rgenza genetiea tra popolazioni e specie ipogee ed epigee di Niphargus (Crustacea, Amphipoda) * INTRODUZIONE Ne11’u1timo decennio la biologia di popolazioni ha ricevuto un notevole impulso da11’uti1izzazione della tecnica elettroforetica, Che, opportunarnente diretta allo studio della mobilite di singole varianti enzimatiche, haq precisato e in gran parte rivoluzionato 1e nostre conoscenze sulla variabilite genetica delle popolazioni animali e vegetali. L’e1ettroforesi permette di riconoscere differenze tra individui negli enzimi e in altre proteine dovute alla 101:0 mobilite in un campo elettrico. Le differenze di carica -elettrostatica che con- dizionano la mobilitzl sono il risultato di sostituzioni, delezioni 0 aggiunte nelle sequenze arninoa-cidiche dei polipeptidi, costituenti Ie proteine, che a loro Volta sono l’effetto di rnutazioni nelle se- quenze nucleotidiche del gene strutturale. Prima de~11’introduzione di questa tecnica da parte di Harris (1966) e Hubby e Lewontin (1966), lo studio della variazione genetica di popolazioni naturali era insoddisfacente per diverse ra- gioni. Per la maggior parte dipendeva soprattutto dalla identifica~ zione di rari mutanti recessivi Che si rnanifestavano soltanto allo stadio omozigote 0 di polimorfismi morfologici e crornosornici che privilegiavano deterrninati organisrni rispetto ad altri come Drow- pbilzz, varie specie di Lepidotteri e Culicidi, Cepaezz etc. (V. Ford, 1975). Oggi grazie a queste tecniche e possibile individuare, stu- diare e misurare differenze genetiche a Iivello molecolare tra in- dividui, popolazioni e specie nei pi1‘1 svariati gruppi animali e vegetali. * Lavoro eseguito con un contribute del CN.R.

Natural and experimental interspecific hybridization between populations ofDolichopoda cave crickets

Evidence of hybridization between 2 pairs of allopatricDolichopoda species is provided by electrophoretic analysis. Occurrence of hybrids was revealed by laboratory crosses and in nature both by occasional co-existence due to passive dispersal and by transplantation experiments.

Divergenza genetica tra popolazioni di Spelaeomysis bottazzii Caroli (Crustacea, Mysidacea) della regione pugliese

13. DE MATTHAEXS, R. COLOGNOLA, M. COBOLLI SBORDONI, L. PESCE q‘, V. SBORDONI Istituto di Zoologia de1l’Universitz‘1 di Roma 7q‘ Istituto di Zoologia de]l’Universit51 di L’Aqui1a Divergenza genetica tra popolazioni di Spelaeomysis bottazzii Caroli (Crustacea, Mysidacea) della regione puglieseq“q‘ INTRODUZIONE Spelczeomysis bottczzzii Caroli e una specie ipogea dulcacquicola di Misidacei, limitata geograficamente alla Puglia a sud del Gargano e conosciuta per Varie popolazioni adattate alle acque cavernicole e freatiche (Ruffo, 1958; Pesce, 1978). Al genere Spelae0myszs ap- partengono anche altre specie, tutte cavernicole, con una geonemia peculiarer si rinvengono, infatti. nelle acque sotterranee della zona centro-americana (Messico, Cuba), (Bacescu e Orghidan, 1971; Bow- man, 1973) e in quella indo-pacifica (Zanzibar), (Page, 1925; Clarke, 1961; Ingle, 1972). Questa distribuzione e Paffinite deile varie specie hanno suggerito 1’ipotesi (Ruffo, 1955) di una colonizzazione antica da parte di queste forrne deH’ambiente ipogeo. ‘ In questo studio vengono presentati dati relativi aila variabilite e grado di divergenza genetica tra quattro popolazioni di Spe- laeonzysis bottazzii indagate attraverso elettroforesi di sistemi gene- enzima. MATERIALE E METODI I carnpioni esaminati provengono da tre pozzi artificiali (Moia di Bari: MOL, Gallipolis GAL, Porto Badisco: PBA) e una grotta naturale (L’Abisso: ABI), situati tutti in term d’Otranto. Si tratta di Iocalitzi costiere delle quali tre si affacciano su11’Adriatico, mentre la quarta, Galiipoli, suilo Ionio. ** Lavoro eseguito con un contribute del CNR.