review article U DC 574 MEIOBENTHiC HARPACTICOIDA (COPEPODA) FROM THE SOUTHERN PART OF THE GULF OF TRIESTE (NORTHERN ADRIATIC) II. ECOLOGY AND SPATIAL DISTRIBUTION Burul VRISER National institute of Biology, Marine Biologicai Station, Sl-6330 Piran, ForrtaCe 41 ABSTRACT The article presents a review of all ecological investigations of harpacticoid copepods in the southern part of the Culf of Trieste between 1970 and 2000, with emphasis on the spatial distribution of species, demonstrated with 90 charts. The diversity analysis of this group indicates that the greater part of the area is inhabited by a single community. Key words: Harpacticoida, Copepoda, Gutf of Trieste, spatial distribution ARPACTICOiDi (COPEPODA: HARPACTiCOIDA) MEiOBENTONIC I DELLA PARTE MERIDIONALE DEL GOLF O Di TRIESTE (NOR D ADRIATICO) IL ECOLOGÍ A E DISTRIBUZIONE SPAZIALE SINTESI L'articolo presenta i risultati deile ricerche etiettuate sulle caratteristiche ecologiche dei copepodi arpacticoidi délia parte méridionale del Golfo di Trieste, tra il 1970 ed il 2000. Particolare attenzione e' stata préstala alia distribuzione spaziale delle specie, che viene evidenziata in 90 cartíne. Nelía gran parte dell'area analizzata la diversité degli arpacticoidi e pressoché uguale, perianto l'autore parla di un 'única comunita costiera di arpacticoidi dei fondali molli. Parole chiave: Harpacticoida, Copepoda, Golfo di Trieste, distribuzione spaziale INTRODUCTIO N Gulf of Trieste: Marcotte & Coull (1974; near Piran), Vriser (1986; Says of Koper, Strunjan and Piran) and Like in almost every part of the marine environment, Vriser (1992; open waters of the Culf). The position of the harpacticoid copepods in the meiofauna of the Gulf sampling sites used during these investigations is shown of Trieste are ranked second as far as their abundance is in figure 1. concerned, immediately after Nematoda. This is why The more comprehensive studies of harpacticoid ecological knowledge of this group is often the key to a communities from larger areas in the Mediterranean in wider understanding of the entire meiobenthos. general are not particularly numerous, but there are Of the total six predominantly ecological studies of certainly none of this particular kind known to us from meiofauna, three have dealt in somewhat greater detail the wider North Adriatic, in the more important Mediter­with the biocenological aspects of harpacticoids and ranean studies, where names like Monard (1935, 1937), spatial distribution of species in the southern part of the Chappuis (1953), Raibaut (1962, 1965, 1967), Soyer Borut VRISER: MEIOBENTHI C HARPACTICOIDA (COPEPODA) FRO M THE SOUTHER N PART .... 39-54 Fig. 1: Investigated area with sampling stations. SI. 1: Raziskovano obmocje z vzorievalnimi postajami. (1964a, b, 1969, 1970a, b), Bodin (1964, 1968), Por (1964) and Castei (1985) stand out, systematics and ecology of this group are normally depicted. in the present article the ecological characteristics of Harpacticoida from the southern part of the Gulf of Tri­este are given as a summary of the three already men­tioned studies (Marcotte & Coull, 1974; Vriser 1986, 1992), with a scheme of spatial distributions of all spe­cies in the area, while a complete list of species and the group's systematics are presented in a separate article (Vriser, 2000). METHODS All samplings were carried out with the gravity core (Meischner & Rumohr, 1974), with 3 replicates (10 cm2 sediment surface, 10 cm deep). Meiofauna was ex­tracted with the sieving-decantation technique of Wieser (1960) on 1 mm, 0.125 mm and 0.050 mm sieves, fixed (4 % formalin with seawater), stained (rose - bengal), sorted out and counted. Homogeneity of the harpacti­coidal communities was examined with the similarity analysis according to Sanders (1960), while the species diversity was established with the Shanon-Wiener index (Pielou, 1969). ECOLOGICAL CONDITIONS Depth span of the sampling sites in the study of coastal barpactioids (Marcotte & Coull, 1974; VriSer, 1986) was between 1 to 15 m, while in the open waters of the Gulf (Vriser, 1992) it was between 19 and 25 m, exceptionally 30 m. Samplings in the coastal areas between 1 and 15 m were carried out in winter (February) and summer (August), all the rest in summer (August). Onl y a few of the shallowest sites (1 - 5 m) in the Bays of Koper and Piran were at. the time of sampling under the local influ­ence of sewage pollution (Marcotte & Coull, 1974; Vriser, 1 986), while all the others were not. The annual thermal conditions of the area's bottom layer vary between 9 and 21 °C, salinity between 35 and 37 PSU, while average oxygen saturation reaches 90% , except during hypoxia crisis when it can fall under 40% . The sediment of the investigated area is represented mostly by clayey silt (10-20% clay), which nearer to the coast and in the interior of the Bays gradually turns into silty clay (25% clay) and towards open waters of the Northern Adriatic into fine sands (Ogorelec etal., 1991). Borul VRLSER: ME íOSE nTHI C 3 EARPACTICO?D A (COPEPODA I FRO M TH E SOUTHER N PAR T .... 33-54 BIOCENOLOGiCA L ANALYSIS O f HARPACTICOIDA Abundance Summer harpacticoid abundances of the coastaf belt between the coast and the depth of 15 m are on average about 100 ind./1G crnz, while in summer they are by half smaller {Marcotte & Couil, 1974; Vriser, 1986). With the depth the abundance increases, so that out of the coastal bays but parallel to the main coastal line it reaches, at the depth of 20 m, 200-250 ind./10 cm2. Towards the centre of the Gulf of Trieste and towards Cape Savudrija the abundance is evenly reduced, at places even below 40 ind./1Q cm2. In the central part of the investigated area this decrease is less distinct. There were 25 harpacticoid species on average at separate localities. None of the species surpassed 9 % of total relative density, and only 22 occurred with more than 1%, which means that the great majority of the species are represented by only a few individuals (Vriser, 1992). The most common species in the area, in view of % of their relative abundance, are; Halo­ •schizopera pontarchis (8.47%), Typhlarnphiascus con­fusus (7.16%), Bulbamphiascus inermis (7.10%), Cle­todes pusillus (634%) , Stenhelia adriatica (4.96%), etc. Diversity and biocenoiogy of coastal harpacticoids At coastal stations the values of the Hs (Shanon ­Wiener) diversity index were in spite of some minor oscillations quite stable (Hs = 1.7-2.2). They drastically fell (Hs = 0.2) only in the direct vicinity of the two se­wage outflows in the Bays of Koper and Piran (Marcotte Couil, "1974; Vriser, 1986). W e beiieve that here w e ;Sr'e dealing with two communities - with due to pollu­tion impoverished community of stressful environment and with the community of the remaining clean, un­polluted coastal belt, where two associations are detected. The first, which is of somewhat higher diver­sity, coincides with the belt of sea meadows to the depth of 10 m, while the other, deeper association belongs to the bare sediment floor. Diversity and biocenoiogy oí harpacticoids of the outer areas ^ In further text the data presented are those from the study by Vriser (1992). The Shannon - Wiener diversity index (Hs) indicated a large diversity range (1.88-3.02) which, however, was in such diverse environment more or less expected. For the fact is that the geological structure of the sea floor changes from slimy clayey silts df the southeastern coastal margin to fine sands of the open sea in the west. Diversity increases in the direction towards the sandy floor and is here the greatest (3.02); it decreases towards the centre of the Gulf of Trieste (2.10) and in the immediate vicinity of Cape Piran. An estimate of the affinity indices of all available pairs of harpacticoid samples for the assessment of bio­cenologically similar areas indicated a few nuclei. The most distinct is the area of the seven stations with high fauna! affinity (over 70% ) on clayey silt, of the outer margin of Koper Say (Fig. 2, area A). Other nuclei were also observed and represent a gradual transition from clayey-sandy silt to sandy floor of the open sea (Fig. 2., areas B, C, D) and in the direction of poorer community (in terms of its species composition and abundance) of the centre of the Gulf and the vicinity of Cape Piran (Fig. 2., areas E and F). By considering the degree of associability estab­lished with the above mentioned similarity analysis, the species diversity and arrangement of the specific species occurring only on certain types of floor, and the prevail­ing, dominating species, w e can divide the investigated area into six ecological units (Fig. 2): Zone A - province of clayey silty floor. Seven stations of high associability. Average diversity Hs = 2.57. Spe­cific species: Pontocletodes ponticus, Stenhelia (Dela­valia) sp.1, sp.2, Harpacticus sp.5, Laophonte sp.4, Typhlarnphiascus sp.1.; Dominant species (succession of prevalence): Haloschizopera sp.1, Typhlarnphiascus confusus, Cletodes pusillus, Stenhelia (Oetavalia) adri­atica, Proameira sp. 1. Zone B - province of sandy-silty floor. Ten stations of medium associability of samples. Average diversity Hs = 2.53. Specific species: Nitocra fcagMs, Ameira sp.1; Dominant species {succession of prevalence)'. Suibam­ phiascus inermis, Haloschizopera pontarchis, Halo­schizopera sp. 7, Cletodes pusillus. Zone C - province of sandy floor. Ten stations of medium associability of samples. Average diversity Hs = 2.77. Specific species: Cletodes limicola, Paradactylo­podia brevicornis, Paralaophonte sp. 1, Laophonte sp.3, Harpacticus sp3., sp4., Enhydrosomelia staufferi, Am­phiascopsis sp.1, Ectinosoma sp.2, Pseudobradya sp.1, Harpacticus sp.1, Laophonte longicaudata; Dominant species: Haloschizopera pontarchis, Heteropsyllus sp.1, Bulbamphiascus inermis, Heterolaophonte stroemi, Stenhelia (Delavalia) adriatica. Zone D - sandy floor of deeper open waters. This probably transitional area is represented by two stations with iow associability without a clear affinity for other stations in the area. Average diversity Hs - 2.59. Spe­cific species: D'Arcythompsonia sp.1, Ameira sp.1; Dominant species: Typhlarnphiascus confusus, Halo­schizopera pontarchis, Bulbamphiascus inermis. Robert­sonia knoxt, Cletodes pusillus. BOFUI VRIŠER: MEIOBENTHI C H ARPACTI C O ID A (COPEPODA) FRO M THE SOUTHER N PART ..., 39-54 Fig. 2: Biocenological distribution of ecological provincesD, E, F transition areas. Si. 2: Biocenološka razmejitev ekoloških provinc A, B,področij D, F, F. Zone E - clayey-sandy silt in the central part of the Gulf. On e station only. Hs = 2.32. No specific species. Dominant species: Enhydrosoma sordidum, Cletodes pusillus, Longipedia coronata. Zone F - with only one station off Cape Piran, very low affinities for the stations in zones B and C, and low diversity (Hs - 1.88). No specific species. Dominant species: Bulbamphiascus inermis, Typhlamphiascus confusus, Cietodes pusillus. The presented spatial distribution of harpacticoids is similar to the distribution of the entire meiofauna and its remaining leading groups (Nematoda, Polychaeta; Vriger, 1989, 1991). However, the reduced abundance of copepods in the direction of the Gulf's centre (known for its frequent hypoxia) and in the direction of the open sea is more distinct than in other meiofauna. A partial reason for this may lie in the smaller resistance of har­pacticoids to periodica! hypoxic conditions, at least in comparison with nematods, as reported in a number of studies (e.g. Josefson & Widbom, 1988; Murell & Fieeger, 1989; Travizi, 1990). A, B, C in the coastaI community of Harpacticoida with C obalne združbe harpaktikoidov in njenih prehodnih Among the dominant species of the entire investi­ gated area w e came across three species (Halo­ schizopera pontarchis, Typhlamphiascus confusus, Cle­ todes pusillus), which had been as dominant species referred to also by Soyer (1970b) after carrying out a re­ search in Catalonia on a similar floor. All the leading species are estimated as predominantly eurivalent, dis­ tributed in places from clayey silt to sand. Although the total diversity span of the entire area was relatively large (Hs = 1.88-3.02), it was only be­ tween 2.40 in 2.60 in the greater part of the investigated area, the only exception being the extreme N W part, where the index was higher than 2.80. With the exception of Cape Piran (Hs = 1.88) and the centre of the Gulf (Hs = 2.26), the greater part of the area, including the coastal belt, is still more or less similar and it therefore belongs to the same harpacticoid community. The northernmost part of the area belongs to other, in terms of species and abundance truncated harpacticoid community of the central part of the Gulf known for its frequent hypoxia. The low diversity around Piran Cape is perhaps caused by specific hydrodynamics, but is for the time being still unexplainable. ANNALES • Ser. hist. nat. • 10 - 2000 • 1 (19) Borut VRI ŠE R, MEiOBENTHI C HARPACTICOIDA (COREPODA) FRO M TH€ SOUTHER N PART -., 39-54 The harpacticoid community of the greater part of area is in terms of its diversity more or less uniform, w e the investigated area is not homogeneous but is, accord-believe that w e are dealing with the very same coastal ing to the associabiiity criterion, composed of three community of soft bottom harpacticoids composed of ecological provinces, the most compact of which is the three subunits, which reflect the floor's transition from one by the entrance to the sifty Koper Bay, while the clayey silt to fine sand. other two are much less distinct. The harpacticoid in­vestigations have in fact been confirmed by similar as-SPATIAL DISTRIBUTION OF SEPARATE SPECIES sessments (Vriser, 1989, 1991) about the meiofauna's division into coastai community (although in copepods Spatial distribution of ail to date registered harpacti­with more distinct marks of ecologically transitional area coid species of the southern part of the Gulf of Trieste is than in meiofauna) and hypoxic, poorer meiofaunal presented on figures 3-12. The number in front of the community of the Gulf s centre. name of each species is a successive number from the Considering that the greater part of the investigated collective list of species (see Vriser, 2000). Figs, 3-12: Spatial distribution of harpacticoid species. Si. 3-12: Prostorska razporeditev harpaktikoidnih vrst. Chart K shows the distribution of the following species: Karta K prikazuje razporeditev sledečih vrst: 5. Arnonardia similis, 7. Amphiasceila proxima, 9. Amphiascopsis thaiestroides, 10. Amphiascopsis cinctus, 14. Amphiascus caudaespinosus, 15, Amphiascus congener, 16. Amphiascus minutus, 23. Buibamphiascus minutus, 25. Brianoia stebieri, 31. Cietodes longicaudatus, 32. Cletodes tenuipes, 35. Dactylopodeiia fiava, 37. Danieissenia perezi, 45, Ectinosoma melaniceps, 50. Enhydrosoma longifurcatum, 53. Enhydrosorna tunisensis, 56. Eurycietodes (Oligocletodes) iatus, 59. Halectinosoma angulifrons, 60. Harpacticus obscurus, 61. Harpacticus tenellus, 67. Haloschizopera bulbifera, 68. Haloschizopera junodi, 73. Heterolaophonte quinquispinosa, 79. Itunelia muelieri, 88. Laophontopsis lamellifera, 90. iongipedia pontica, 92. Mesopsyilus atargatis, 94. Nitocra divaricata, 98. Pa ra da n ielssen i a kunzi, 100. Paralaophonte brev'srostris, 111. Robertgurneya rostrata, 112. Robertgurneya ecaudata, 114, Robertson i a propinqua, 118. Stenhelia (Delavaiia) intermedia, 119. Stenheiia (Oelavaiia) refiexa, 125. Tisbe gracilis, 126. Tisbe clodiens, 127. Tisbe reluctans, 130. Zosime atlantica Bofu! VRISEB: MFSOBENTHIC HARPACTÍCOSDA (COPEPODA} FROM THE SOUTHERN PART ..., 39-S4 1. Acrenhydrosoma perplexum & 2. Ameira parvula . 3. Ameira sp. 1. 4. Ameira sp.2 6. A mph iascoides debilis 8 . Amphiascoides sp, 1 r 11. Amphiascopsis sp. 1 12. Amphiascopsis sp.2 13. Amphiascopsis sp.3 17. Amphiascus varians ANNALES • Ser. hist. nat. 10 • 2000 -1 (19) Boro! VRIŠER: MFlORENTHC HARPACTCOÍDA COPEPOPA) FROM THE SOUTHLRN PART 39-S4 18. Amphiascus sp.l 19. Asellopsis sp. 1 20, Bradya ('Bradya) typica R^I 21. Bulbamphiascus imus 22, Bulbamphiascus inermis 24. Bulbamphiascus sp. 2 € 26. Canuellafurcigera JT"* 27. Canuellaperplexa j f 29. Cletodespusillus C -> 30. Cletodes limicola f L1 Borit! VRíSER: MEIOSE NTH IC HARÍ'ACTJCOIDA (COPITOÜA) FROM THE SOUTHERN PART .... 39-M 33. D Arcythompsonia scotti t jT^ 36. Dactylopodia tisboides 39. Diarthrodes minut us 41. Diosaccus sp.l J i 43. Ectinosoma obtmum 34. DArcythompsonia sp 1 38. Diagoniceps menaiensis ITs*­ J r r 40. Diosaccus tenuicornis ( eptr** * w. € 42. Ectinosoma normani 44. Ectinosoma dentatum BoruS VRlSER: MEfOBENTHiC HARPACT1COIDA (COPEPODA) FRO M THE SOUTHER N PART .. 39-54 46. Ectinosoma sp. 1 47. Ectinosoma sp.2 i 48. Enhydrosoma buchholtzi V 1 .^y-' f ^ ^ t 51. Enhydrosoma sordidum 54. Enhydrosomella staufferi 57. Eurycletodes (Oligocletodes) spAr­ 1 •V­ f / 49. Enhydrosoma caeni 52. Enhydrosoma propinqum 55. Esola longicauda j T I 58. Euterpina acutifrons Boru! VRíSER: MEiOSENTHtC HARPACTiCCHOA (COPfPODAI FRO M THE SOUTHERN PART ..., 33-51 62. Harpacticus sp. 1 63. Harpacticiis sp.2 ^-Jfry. 64. Harpacticus sp. 3 65. Harpacticus spA € 66. Harpacticus sp. S 69. Haloschizopera pontarchis é­ -a c . 70. Haloschizopera sp.i 71. Hemimesochra nixe r (% f ^ . V. c ­ 72. Hemimesochra sp. 1 74, Heterolaophonte stroemi € p or aim n u1 a' v * t : c . 80. Laophonte longicaudata 81. Laophonte sima 82. Laophonte inornata 83. Laophonte cornuta " t L 84. Laophonte sp. 1 f 85. Laophonte sp.2 J% i Boru! VRiŠK : MtlOSENTHfC HARPACTICOIDA (COPEPODA) FROM THE SOUTHERN PART .59-S4 Bonn VRlSER: ML!OBENTHIC HARP ACTiCOlOA {COPEPODA; FROM THE SOUTHERN PART 39-54 86» Laophonte sp. 3 87. Laophonte spA « S f > 88. Laophontopsis lamelifera L r " c 102. Paramphiascella coulli «3 * 104. Parathalestris sp. 1 J ­ . T W' " I . 106. Phyllopodopsyllus sp, 1 I 108. Proameirq sp. 1 i 110. Rhynchothalestris rufocinct S ' ii (i ' t 4 B^TvRiŠER : MEIOBENTHIC HARI'ACTICOIDA (CO PEPO D A) FROM THE SOUTHERN PART ^ 113. Robertsonia knoxi . % 115. Stenhelia (Delavalia) normami— " . * . ' . • * > . * • f C '""-JS : . .. 116. Stenhelia (Delavalia) mimit^ ^' 117. Stenhelia (Delavalia) adriati^a ­. * * N-... ••••••' > • ? ! /• ' -v. ° y 1 1 %.. ' 120. Stenhelia (Delavalia) sp. 1 ^ ^ 121. Stenhelia (Delavalia) sp. 2 ^ ^ • « < • Jtv" • J?'­ ' -• • '. • * > 122. Stylicletodes longicaudatus ^^ 123. Thalestris rufoviolascens — if^"--. i ' 124. Tisbe lancii ^^ 128. Typhlamphiascus confusus • ' ^ • . r S •. • • -•. • X . ' • ••'••' • * 'I t ^ ' — Somi VRÉER : MEIOBENTHf C HASPACÎICOID A (COPEPODA) FRO M THE SOUTHER N PART 39-54 K çr' ' ' r > ^ fu. ij % •. ' J . ... MEIOBENTOŠKI HARPAKTIKOIDi (COPEPODA: HARPACT1COIDA) JUŽNEGA DELA TRŽAŠKEGA ZALIVA. II. EKOLOGIJA IN RAZŠIRJENOST VRST Borut VRIŠER Nacionalni inštitut za biologijo, Morska biološka postaja, Sl-6330 Piran, Fornače 41 POVZETEK Prispevek podaja pregled dosedanjih raziskav ekoloških značilnosti harpaktikoidov južnega dela Tržaškega za­liva s posebnim poudarkom na prostorski razširjenosti posameznih vrst; ki jih ponazarja 90 kart. Diverziteta harpa­ktikoidov pretežnega dela raziskovanega področja je dokaj enotna, zato lahko govorimo o eni, skupni obalni združbi harpaktikoidov mehkega dna. To združbo sestavljajo tri izrazitejše ekološke province, ki odsevajo prehod morskega dna od glinastih meljev obalnega pasu do finih peskov odprtega dela zaliva. Dve drugi opaženi združbi harpaktikoidov sta obrobnega pomena. 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