Research Article ISSN 2336-9744 (online) | ISSN 2337-0173 (print) The journal is available on line at www.ecol-mne.com

A checklist of halacarid and pontarachnid (: and ) associated with mangroves

TAPAS CHATTERJEE

Department of Biology, Indian School of Learning, I.S.M. Annexe, P.O. – I.S.M., Dhanbad – 826004, Jharkhand, India. E–mail: [email protected]

Received 14 June 2015 │ Accepted 23 June 2015 │ Published online 25 June 2015.

Abstract This paper is a compilation of the records for halacarid and pontarachnid species associated with mangroves. A total of 23 halacarid species (Acari: Halacaridae) belonging to the five genera Acarothrix, Agauopsis, , Isobactrus and Rhombognathus and six pontarachnid species (Acari: Pontarachnidae) belonging to the genus are associated with various microhabitats of mangroves. Mites are found mainly in the algae and sediment covering pneumatophores and aerial roots.

Key words: Checklist, Mangrove, Halacaridae, Pontarachnidae.

Introduction

Tidal mangrove forests cover a vast area of world’s coastlines and are precious resources for multiple economic and ecological reasons. As much as 39.3 million acres of mangrove forests are present along the warm-water coastlines of tropical oceans all over the world. However, mangroves are diminishing worldwide at a faster rate than other terrestrial forests, making them one of the most threatened ecosystems in the world. Mangroves are habitats for a diverse aerial, terrestrial and marine fauna (Nagelkerken et al. 2008). Vast amounts of intertidal small fauna and meiofauna are associated with mangroves, mainly on turf growing on mangrove aerial roots and pneumatophores (e.g. Farnsworth & Ellison 1996; Gwyther 2000; Procheş et al. 2001; Procheş 2002, 2004; Procheş & Marshall 2002a; Bartsch 2003a, b, c, 2006a, b; Chatterjee et al. 2012a, b, 2013), litter falls of mangrove leaves (eg. Alongi 1989, 1990; Gee & Somerfield 1997; Somerfield et al. 1998; Torres–Pratts & Schizas 2007), and mud flats in mangrove zones (e.g. Hodda & Nicholas 1985, 1986a, b, 1987, 1990; Nicholas et al. 1991; Sasekumar 1994; Ólafson et al. 2000; Netto & Gallucci 2003). Mites are found mainly in the algal or sediment cover on pneumatophores, stems and branches. The major groups of mites reported from mangroves are halacarids (Halacaridae, Prostigmata) (e.g. Bartsch 1990, 1991, 1997; Chatterjee et al. 2012a, b, 2013), oribatids (Oribatida, including the cohort Astigmatina) (e.g. Marshall & Pugh 2000, 2002; Ermilov et al. 2013) and pontarachnids (Pontarachnidae, Hydrachnidia, Prostigmata) (e.g. Pešić et al. 2008, 2011, 2012, 2013), alongwith some Mesostigmata. In the present paper, a checklist of halacarid and pontarachnid mites associated with mangroves is presented.

Ecol. Mont., 2 (4), 2015, 334-341

CHATTERJEE

Methods

This checklist was compiled using published records (up to June 13th, 2015). The genera and species are arranged in alphabetical sequence.

Results

Halacarid mites associated with mangroves: Family Halacaridae Murray, 1877

Acarothrix ampliumeris Bartsch, 2006 Record associated with mangroves: Cladophora mat, Chlorophyta, on muddy and sandy sediments in mangrove area, Singapore – Bartsch (2006a). Distribution: Singapore.

Acarothrix grandocularis Chatterjee et al., 2012 Record associated with mangroves: From algal turf growing on Rhizophora sp. pneumatophores at Batu Marang, Brunei Darussalam – Chatterjee et al. (2012a); from algal turf growing on Avicennia sp. pneumatophores at Chorao Island, North Goa, India – Chatterjee et al. (2012a). Remarks: Acarathrix grandocularis was also mentioned from Singapore along with Rhombognathus amplus in algae on mangroves (Bartsch 2013). Distribution: Brunei Darussalam; India; Singapore.

Acarothrix longiunguis Bartsch, 1997 Record associated with mangroves: In soft mud from mangrove area, Sadgroves Creek, near Darwin, Northern Australia – Bartsch (1997). Distribution: Northern Australia.

Acarothrix palustris Bartsch, 1990 Record associated with mangroves: Algal turf on salt marshes and mangrove flats, Hong Kong, southern China – Bartsch (1990); green algae and epibiota on Avicennia pneumatophores in a rockpool, Pandan River, southern coast of Singapore – Bartsch (2006a); northern coast of Singapore, end of Lim Chu Kang Road from Cladophora mat on muddy and sandy sediment in mangrove area – Bartsch (2006a); from algal turf growing on Avicennia pneumatophores at Chorao Island, North Goa, India – Chatterjee et al. (2013). Distribution: Southern China; Singapore; India.

Acarothrix umgenica Procheş, 2002 Record associated with mangroves: Beachwood and Bayhead Lagoon, near Durban, KwaZulu Natal, South Africa, sediment on Avicennia pneumatophores – Procheş et al. (2001). South Africa, KwaZulu–Natal, Beachwood mangroves in Durban, and Richards Bay (South Africa, KwaZulu–Natal) from sediment or algae covering the pneumatophores of the mangrove tree Avicennia marina – Procheş (2002). Distribution: South Africa; Mozambique.

Agauopsis arborea Bartsch, 2003 Record associated with mangroves: Western Australia, Dampier, on the Burrup Peninsula; in muddy algal turf (Bostrychia – Calloglossa algal association) on the lower trunks and pneumatophores of mangroves – Bartsch (2003b). Distribution: Western Australia.

Agauopsis dasyderma Bartsch, 2003 Record associated with mangroves: Western Australia, east coast of the Burrup Peninsula; turf on trunk of Avicennia marina, also among epibionts (barnacles, mussels, turf) on the underside branches of A. marina – Bartsch (2003a). Distribution: Western Australia.

Ecol. Mont., 2 (4), 2015, 334-341 335

HALACARID AND PONTARACHNID MITES ASSOCIATED WITH MANGROVES

Copidognathus balakrishnani Chatterjee, 2000 Record associated with mangroves: Among algal turf growing on Avicennia pneumatophores at Chorao Island, North Goa, western India – Chatterjee (2015). Remarks. Chatterjee (2000) described this species from intertidal algae in Cochin, Kerala, India. Distribution: India.

Copidognathus bruneiensis Chatterjee, Marshall & Pešić, 2012 Record associated with mangroves: On mud flat and in algae covering the pneumatophores of Avicennia marina trees; Brunei Darussalam, Kota Batu, Sungai Brunei Estuary and Bay – Chatterjee et al. (2012b). Distribution: Brunei Darussalam.

Copidognathus caloglossae Procheş, 2002 Record associated with mangroves: South Africa, Eastern Cape, Mtakatye Estuary; KwaZulu–Natal, Bayhead Lagoon in the Durban Harbour; Richards Bay, Isipingo and Beachwood mangrove forests in KwaZulu – Natal and Inhambane (Mozambique) from an algal association known as `Bostrychietum’, covering the pneumatophores of Avicennia marina – Procheş (2002); in sediment covering pneumatophore in Baywood mangrove forest – Procheş & Marshall (2002b). Beachwood and Bayhead Lagoon, near Durban, KwaZulu- Natal, South Africa, sediment on Avicennia pneumatophores – Procheş et al. (2001). Distribution: South Africa; Mozambique.

Copidognathus lutarius Bartsch, 2003 Record associated with mangroves: Western Australia, east coast of the Burrup Peninsula; algal turf on pneumatophores of Avicennia – Bartsch (2003a). Distribution: North western Australia.

Copidognathus mangrovorum Chatterjee, Marshall & Pešić, 2012 Record associated with mangroves: On mud flat and in algae covering the pneumatophores of Avicennia marina; Brunei Darussalam, Kota Batu, Sungai Brunei Estuary and Bay – Chatterjee et al. (2012b). Distribution: Brunei Darussalam.

Copidognathus paluster Bartsch, 1991 Record associated with mangroves: Hong Kong, southern China; algae (Cladophora) on mangrove flats – Bartsch (1991). Distribution: Southern China.

Copidognathus piger Bartsch, 2003 Record associated with mangroves: Western Australia, east coast of the Burrup Peninsula; turf on pneumatophores of Avicennia marina – Bartsch (2003a). Distribution: Western Australia.

Copidognathus rhombognathoideus Bartsch, 2006 Record associated with mangroves: Singapore, among mangrove areas – Bartsch (2006a); among algal turf growing on Rhizophora pneumatophores at Batu Marang, Brunei Darussalam – Chatterjee et al. (2012b). Distribution: Singapore; Brunei Darussalam.

Isobactrus australiensis Bartsch, 2003 Record associated with mangroves: Western Australia, east coast of the Burrup Peninsula; underside of Avicennia branches among barnacles – Bartsch (2003c). Distribution: Western Australia.

Isobactrus obesus Bartsch, 1992 Record associated with mangroves: Western Australia, Dampier, east coast of the Burrup Peninsula; underside of Avicennia marina branches among barnacles; also found in coarse sediments in small mangrove area – Bartsch (2003c).

336

CHATTERJEE

Remarks: Bartsch (1992) described this species among shells of rock oysters (Saccostrea cucullata), sediment and pebbles with green algae from Hong Kong, southern China. Distribution: Southern China; Western Australia.

Rhombognathus abirus Pepato & Da Silveira, 2015 Record associated with mangroves: Algal turfs on aerial roots, trunks and pneumatophores of mangroves Rhizophora mangle and Avicenia schaueriana from Ubatuba, São Paulo, Brazil along the Fazenda River – Pepato and Da Silveira (2015). Distribution: Brazil.

Rhombognathus amplus Bartsch, 2013 Record associated with mangroves: Singapore, among green and red algae on trunks of Rhizophora sp. – Bartsch (2013). Distribution: Singapore.

Rhombognathus aspidotus Bartsch, 2006 Record associated with mangroves: Singapore, among Bostrychia mats (Rhodophyta) on Avicennia pneumatophores – Bartsch (2006b). Distribution: Singapore

Rhombognathus bulbosus Bartsch, 2005 Record associated with mangroves: Found among unbranched thread-like green algae (Cladophorales, Chlorophyta) and debris from stem of mangrove Rhizophora sp., in a small bay of the Pandan River, Singapore and from algae (Chlorophyta) on Avicennia pneumatophores growing in adjacent tidal field – Bartsch (2005) Distribution: Singapore.

Rhombognathus ocularis Bartsch, 2003 Record associated with mangroves: Western Australia, east coast of the Burrup Peninsula; in algal turf (mainly red algae Caloglossa and Bostrychia) present on branches, stem and pneumatophores of Avicennia marina – Bartsch (2003a, c). Distribution: Western Australia.

Rhombognathus picinguabensis Pepato & Da Silveira, 2015 Record associated with mangroves: Algal turfs on aerial roots, trunks and pneumatophores of mangroves Rhizophora mangle and Avicenia schaueriana from Ubatuba, São Paulo, Brazil along the Fazenda River – Pepato & Da Silveira (2015). Distribution: Brazil.

Pontarachnid mites associated with mangroves: Family Pontarachnidae Koenike, 1910

Litarachna bartschae Smit, 2003 Record associated with mangroves: Brunei Darussalam, Brunei Bay, Pulau Bedukang; among epiphytal algae from mangrove pneumatophores – Pešić et al. (2013). Remarks: Smit (2003) described this species without providing information as to the exact microhabitat in Watering Cove, Burrup Peninsula, Western Australia. Distribution: Western Australia; Brunei Darussalam.

Litarachna bruneiensis Pešić, Chatterjee, Marshall & Pavićević, 2011 Record associated with mangroves: Collected from dense Avicennia marina pneumatophore mats, depth about 2 m, Brunei Darussalam, Kota Batu, Sangai Brunei Estuary and Bay – Pešić et al. (2011); among epiphytal algae from mangrove pneumatophores, Pulau Bedukang, Brunei Darussalam, Brunei Bay – Pešić et al. (2013).

Ecol. Mont., 2 (4), 2015, 334-341 337

HALACARID AND PONTARACHNID MITES ASSOCIATED WITH MANGROVES

Distribution: Brunei Darussalam.

Litarachna caribica Pešić, Chatterjee & Schizas, 2008 Record associated with mangroves: Among epiphytal algae from red mangrove Rhizophora mangle roots, Caribbean Sea, Netherland Antilles, Curaçao, Niewport – Pešić et al. (2008). Remarks: Pešić et al. (2015) recorded this species in algae and rubble from Taboguilla Island, Panama (Pacific coast of Panama). Distribution: Caribbean Sea; Pacific coast of Panama.

Litarachna degiustii Cook, 1958 Record associated with mangroves: Among epiphytal algae from red mangrove Rhizophora mangle roots, Caribbean Sea: Netherland Antilles, Curaçao, Niewport and also from Bahamas, near Lee Stocking Island, Norman's Pond Cay – Pešić et al. (2008) Remarks: Cook (1958) described this species from Bimini, Bahamas. Distribution: Caribbean Sea.

Litarachna minuta Pešić, Chatterjee & Marshall 2013 Record associated with mangroves: Brunei Darussalam, Brunei Bay, Pulau Bedukang; among epiphytal algae from mangrove pneumatophores – Pešić et al. (2013). Distribution: Brunei Darussalam.

Litarachna indica Pešić, Chatterjee & Ingole, 2012 Record associated with mangroves: India, Goa (North Goa), Virnoda Pernem, Arabian Sea, from algae associated with Rhizophora pneumatophores – Pešić et al. (2012). Distribution: India.

Discussion

Halacarid mites live in a variety of substrata including within tufts of algae, on large fronds, in and on colonies of sponges, hydrozoans, bryozoans, barnacles, mussels and polychaetes, in interstices of coarse sediment or flocculent ooze, and in intertidal sand and turf growing on mangrove pneumatophores. Halacarids are also known to dwell on the rough and hairy body surfaces of crustaceans and gastropods, adjacent to gill chambers of crustaceans and mollusks and between the spines and in the gut of echinoderms. More than 1000 species of halacarids are currently recognized of which only 23 species have thus far been found to be associated with mangroves. Among the diverse mangrove microhabitats of mangroves (turf growing on pneumatophores, turf growing on small lower branches touching the sediment or water level, mud flats, litter falls), halacarids are mostly found on turf growing on mangrove aerial roots and pneumatophores and on turf growing on the small lower branches which touch the sediment or the water surface. While individual halacarid species have been recorded only from mangrove habitats, no halacarid genera have thus far been found to be exclusively associated with mangroves (although majority of Acarothrix seems to occur mainly here). Extensive sampling in other habitats may prove that some apparent mangrove specialists may be found elsewhere, as well. The Pontarachnidae Koenike, 1910 is the only family of water mites (Hydrachnidia) that is adapted to marine waters. Pontarachnids have been reported from both tropical and temperate littoral habitats. Most species live in the marine littoral zone; a few species have been described from freshwater, but these species have been found to occur only in estuaries or in locations near the sea (Smit 2009). The family is represented by two genera, Pontarachna Philippi, 1840 and Litarachna Walter, 1925 and 45 described species. Six species of Litarachna are known to be associated with mangroves, while none of the 24 known species of Pontarachna have been found in mangrove habitats. Our current knowledge of the diversity of mangrove-associated halacarid and pontarachnid mites is far from complete. Future investigations of the mangrove milieu are expected to reveal many more species.

338

CHATTERJEE

Acknowledgements Thanks are due to Dr. Nikolaos V. Schizas, University of Puerto Rico, Mayagüez, USA for linguistic help. The author is also grateful to Dr. G. W. Krantz, Oregon State University, USA and Dr. Şerban Procheş, University of KwaZulu-Natal, South Africa for their valuable remarks.

References

Alongi, D.M. (1989) The role of soft – bottom benthic communities in tropical mangrove and coral reef ecosystems. Reviews in Aquatic Sciences, 1, 243–280. Alongi, D.M. (1990) Intertidal zonation and seasonality of meiobenthos in tropical mangrove estuaries. Marine Biology, 95, 447–458. Bartsch, I. (1990) Acarothrix palustris gen. et spec. nov. (Halacaroidea, Acari), ein Bewohnerder Salzwiesen Südchinas. Zoologischer Anzeiger, 224, 204–210. Bartsch, I. (1991) Halacariden (Acari) von Hong Kong. Beschreibung von drei Arten der Gattung Copidognathus. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 88, 175– 184. Bartsch, I. (1992) Hong Kong rhombognathine mites (Acari: Halacaridae). In: Morton, B. (Ed.), The Marine Flora and Fauna of Hong Kong and Southern China III, 1. University Press, Hong Kong, pp. 251– 276. Bartsch, I. (1997) Copidognathinae (Halacaridae, Acari) from northern Australia; description of four new species. In: Hanley, J.R., Caswell, G., Megirian, D. & Larson, H.K. (Eds), Proceedings of the Sixth International Biological Workshop. Museums & Art Galleries of Northern Territory, Darwin, pp. 231–243. Bartsch, I. (2003a) Mangrove halacarid fauna (Halacaridae, Acari) of the Dampier region, Western Australia, with description of five new species. Journal of Natural History, 37, 1855–1877. Bartsch, I. (2003b) A new species of the brevipalpus group, genus Agauopsis (Halacaridae: Acari) from Dampier, Western Australia: description and key to related species. In: Wells, F.E., Walker, D.I. & Jones, D.S. (Eds), The Marine Flora and Fauna of Dampier, Western Australia. Western Australian Museum, Perth, pp. 241–254. Bartsch, I. (2003c) Rhombognathine mites (Halacaridae: Acari) from Dampier, Western Australia: and biogeography. In: Wells, F.E., Walker, D.I. & Jones, D.S. (Eds), The Marine Flora and Fauna of Dampier, Western Australia. Western Australian Museum, Perth, pp. 255–280. Bartsch, I. (2005) Upper littoral rhombognathines (Acari: Halacaridae) of Singapore: description of three new species. Cahiers de Biologie Marine, 46, 273–287. Bartsch, I. (2006a) Copidognathines (Acari: Halacaridae) in mangroves of Singapore. I. Description of three species. The Raffles Bulletin of Zoology, 54, 83–92. Bartsch, I. (2006b) A new species and new record of Rhombognathus from Singapore (Acari:Halacaridae). Zootaxa, 1120, 41–49. Bartsch, I. (2013) Upper littorals Rhombognathines (Acari: Halacaridae) from Singapore: Description of a new species. Acarologia, 53 (3), 305–313. Chatterjee, T. (2000) Two new species of Copidognathus (Halacaridae, Acari) from Kerala. Journal of the Bombay Natural History Society, 97, 253–259. Chatterjee, T. (2015) Scanning electron microscopic observations of Copidognathus balakrishnani Chatterjee (Acari: Halacaridae) with notes on species of the Copidognathus balakrishnani group. Acta Biologica. (in Press). Chatterjee, T., Marshall, D.J., Guru, B.C., Ingole, B. & Pešić, V. (2012a) A new species of the genus Acarothrix (Acari, Halacaridae) from Brunei Darussalam and India. Cahiers De Biologie Marine, 53 (4), 541–546 Chatterjee, T., Marshall, D.J. & Pešić, V. (2012b) New records of Copidognathus mites (Acari: Halacaridae) from mangroves in Brunei Darussalam with descriptions of two new species. Zootaxa, 3269, 18–30. Chatterjee, T., Guru, B.C. & Sørenson, M.V. (2013) Report of Acarothrix palustris Bartsch (Acari: Halacaridae) from the Indian Ocean. Acta Biologica, 20, 17–26. Cook, D.R. (1958) A new species of Litarachna from the British West Indies (Acarina: Pontarachnidae). Proceedings of the Entomological Society of Washington, 60, 19–22.

Ecol. Mont., 2 (4), 2015, 334-341 339

HALACARID AND PONTARACHNID MITES ASSOCIATED WITH MANGROVES

Ermilov, S.G., Chatterjee, T. & Marshall, D.J. (2013) Two new species of oribatid mites of Oripodoidea (Acari: Oribatida) from Brunei. Annales Zoologici, 63 (3), 393–400. Farnsworth, E.J. & Ellison, A.M. (1996) Scale-dependent spatial and temporal variability in biogeography of mangrove root epibiont communities. Ecological Monographs, 66(1), 45–66. Gee, M.J. & Somerfield, P.J. (1997) Do mangrove diversity and leaf litter decay promote meiofaunal diversity? Journal of Experimental Marine Biology and Ecology, 218, 13–33. Gwyther, J. (2000) Meiofauna in phytal-based and sedimentary habitats of a temperate mangrove ecosystem— a preliminary survey. Proceedings of the Royal Society of Victoria, 112, 137–151. Hodda, M. & Nicholas, W.L. (1985) Meiofauna associated with mangroves in the Hunter river estuary and Fullerton Cove, southern Australia. Australian Journal of Marine and Freshwater Research, 36, 41– 50. Hodda, M. & Nicholas, W.L. (1986a) Temporal changes in littoral meiofauna from the Hunter River Estuary. Australian Journal of Marine and Freshwater Research, 37, 729–741. Hodda, M. & Nicholas, W.L. (1986b) Nematode diversity and industrial pollution in the Hunter river estuary, NSW, Australia. Marine pollution Bulletin, 17, 351–355. Hodda, M. & Nicholas, W.L. (1987) Free living marine nematodes from Darwin mangroves. The Beagle, 4, 3–10. Hodda, M. & Nicholas, W.L. (1990) Production of meiofauna in an Australian Estuary. Wetlands, 9(2), 41– 48. Marshall, D.J. & Pugh, P.J.A. (2000) Two new species of Schusteria (Acari: Oribatida: Ameronothroidea) from marine shores in southern Africa. African Zoology, 35(2), 201–206. Marshall, D.J. & Pugh, P.J.A. (2002) Fortuynia (Acari: Oribatida: Ameronothroidea) from the marine littoral of southern Africa. Journal of Natural History, 36, 173–183. Nagelkerken, I., Blaber, S.J.M., Bouillon, S., Green, P., Haywood, M., Kirton, L.G., Meynecke, J.O., Pawlik, J., Penrose, H.M., Sasekumar, A. & Somerfield, P.J. (2008) The habitat function of mangroves for terrestrial and marine fauna: a review. Aquatic Botany, 89, 155–185. Netto, S.A. & Gallucci, F. (2003) Meiofauna and macrofauna communities in a mangrove from the Island of Santa Catarina, South Brazil. Hydrobiologia, 505, 159–170. Nicholas, W.L., Elek, J.A., Stewart, A.C. & Marples, P.G. (1991) The nematode fauna of a temperate Autralian mangrove mudflat- its population density, diversity and distribution. Hydrobiologia, 209, 13–28. Ólafson, E., Carlstrm, S. & Ndaro, S.G.M. (2000) Meiobenthos of hypersaline tropical mangrove sediment in relation to spring inundation. Hydrobiologia, 426, 57–64. Pepato, A.R. & Da Selveira, P.S.A. (2015) Two new species of Rhombognathus (Halacaridae, ) from a mangrove in the northern littoral zone of São Paulo State (Brazil). Zootaxa, 3905(4), 500–510. Pešić, V., Chatterjee, T. & Schizas, N.V. (2008) Marine water mites (Acari: Hydrachnidia: Pontarachnidae) from the Caribbean Sea, with description of one new species. Cahiers de Biologie Marine, 49(3), 253–259. Pešić, V., Chatterjee, T., Marshall, D.J. & Pavićević, A. (2011) New records of water mites (Acari: Hydrachnidia) from Brunei Darussalam, Borneo, with descriptions of two new species. Zootaxa, 3018, 50–58. Pešić, V., Chatterjee, T., & Ingole, B., Velip, V. & Pavićević, A. (2012) New species of Litarachna Walter, 1925 (Acari, Hydrachnidia) from the west Indian coast, with a discussion on the biodiversity of the family Pontarachnidae Koenike, 1910. Cahiers De Biologie Marine, 53(4), 547–553. Pešić, V., Chatterjee, T. & Marshall, D.J. (2013) Marine water mites (Acari: Hydrachnidia, Pontarachnidae) from the Brunei Bay, with description of one new species. Cahiers de Biologie Marine, 54(3), 405– 410. Pešić, V., Chatterjee, T. & Schizas, N.V. (2015) First record of Litarachna caribica (Acari, Pontarachnidae) from the Pacific coast of Panama. Marine Biodivetrsity Records, 8 (e85), 1–3. Procheş, Ş. (2002) New species of Copidognathinae (Acari: Halacaridae) from southern Africa. Journal of Natural History, 36, 999–1007. Procheş, Ş. (2004) Ecological associations between organisms of different evolutionary history: mangrove pneumatophore as a case study. Journal of the Marine Biological Association, U.K. 84, 341–344.

340

CHATTERJEE

Procheş, Ş., Marshall, D.J., Ugrasen, K. & Ramcharan, A. (2001) Mangrove pneumatophore assemblages. Journal of the Marine Biological Association, U.K. 81, 545–552. Procheş, Ş. & Marshall, D.J. (2002a) Epiphytic algal cover and sediment deposition as determinants of arthropod distribution and abundance on mangrove pneumatophores. Journal of the Marine Biological Association, U.K., 82, 937–942. Procheş, Ş. & Marshall, D.J. (2002b) Diversity and biogeography of southern African intertidal Acari. Journal of Biogeography, 29, 1201–1215. Sasekumar, A. (1994) Meiofauna of a mangrove shore on the west coast of peninsular Malaysia. Raffles Bulletin of Zoology, 42(4), 901–915. Smit, H. (2003) Five new species of the water mite family Pontarachnidae from western Australia (Acari: Hydrachnidia). In: Wells, F.E., Walker, D.I. & Jones, D.S. (Eds), The Marine Flora and Fauna of Dampier, Western Australia. Western Australian Museum, Perth, pp. 547–562. Smit, H. (2009) Water mites of the family Pontarachnidae from Singapore, with a description of one new species (Acari: Hydrachndia). Raffles Bulletin of Zoology, Supplement, 22, 203–205. Somerfield, P.J., Gee, M. & Aryuthaka, C. (1998) Meiofaunal communities in a Malaysian mangrove forest. Journal of the Marine Biological Association of the United Kingdom. 78, 717–732. Torres–Pratts, H. & Schizas, N.V. (2007) Meiofauna colonization of decaying leaves of the red mangrove, Rhizophora mangle in south western Puerto Rico, Caribbean Journal of Science, 43(1), 127–137.

Ecol. Mont., 2 (4), 2015, 334-341 341