Sandy beaches: state of the art of nematode ecology

MARIA, TATIANA F.; VANAVERBEKE, JAN; VANREUSEL, ANN; ESTEVES, ANDRÉ M.

doi:10.1590/0001-3765201620150282

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Biological Sciences • An. Acad. Bras. Ciênc. 88 (3 Suppl) • 2016 • https://doi.org/10.1590/0001-3765201620150282 linkcopy

Sandy beaches: state of the art of nematode ecology

Authorship SCIMAGO INSTITUTIONS RANKINGS

ABSTRACT

In this review, we summarize existing knowledge of the ecology of sandy-beach nematodes, in relation to spatial distribution, food webs, pollution and climate change. We attempt to discuss spatial scale patterns (macro-, meso- and microscale) according to their degree of importance in structuring sandy-beach nematode assemblages. This review will provide a substantial background on current knowledge of sandy-beach nematodes, and can be used as a starting point to delineate further investigations in this field. Over decades, sandy beaches have been the scene of studies focusing on community and population ecology, both related to morphodynamic models. The combination of physical factors (e.g. grain size, tidal exposure) and biological interactions (e.g. trophic relationships) is responsible for the spatial distribution of nematodes. In other words, the physical factors are more important in structuring nematodes communities over large scale of distribution while biological interactions are largely important in finer-scale distributions. It has been accepted that biological interactions are assumed to be of minor importance because physical factors overshadow the biological interactions in sandy beach sediments; however, the most recent results from in-situ and ex-situ experimental investigations on behavior and biological factors on a microscale have shown promise for understanding the mechanisms underlying larger-scale patterns and processes. Besides nematodes are very promising organisms used to understand the effects of pollution and climate changes although these subjects are less studied in sandy beaches than distribution patterns.

Key words:
biodiversity; benthos; distribution patterns; food webs; climatic changes

Code for fig. 1	Reference	Fauna	Focus	N of studied beach	Main Findings
01	Boaden and Platt 1971	M and N	DT	1	Daily tidal variations and biotic interactions affect the vertical distribution of meiofauna
02	Gray and Rieger 1971	M and N	DT	1	Highest meiofauna richness and diversity in less exposed (finer grain) beach
03	Harris 1972	M	DT	1	High temperatures in the upper beach is a limiting factor for meiofauna zonation
04	Ott 1972	N	DT	1	Presence of a typical nematofauna in sulfide systems
05	Hulings and Gray 1976	M	DT	89	Physical factors control the meiofauna abundance (e.g. wave, tide and currents) in tidal beaches while biological interactions (e.g. predation and competition) may control abundance in atidal beaches
06	McLachlan et al. 1977	M	DT and P	1	Meiofauna can occur down to 35 cm. Sewage can cause an increase of the meiofauna abundance, specially the nematode one
07	Platt 1977	N	DT	1	Horizontal distribution is driven by physical factors (e.g. grain size) and availability of food while vertical distribution is limited by oxygen availability
08	Munro et al. 1978	M	DT	2	Tropical beach shows higher respiration rates, less microbial production and meiofauna biomass when compared to a temperate one
09	Fricke et al. 1981	M and N	P	2	Harpacticoid copepods are more sensitive to oil pollution than nematodes; meiofauna can be recovered six months after an oil spill
10	McLachlan and Harty 1982	M	P	1	Oligochaetes are more sensitive to oil pollution than nematodes; meiofauna can be recovered five months after an oil spill
11	Ansari and Ingole 1983	M	DT	6	Meiofauna is more abundant in the upper 4 centimeters
12	Blome 1983	N	DT	1	Nematodes are more abundant in the oxic layers and shows a horizontal zonation; vertical migration is attributed to temperature variations
13	Sharma and Webster 1983	N	DT	2	Highest nematode density in well sorted medium grain size: most nematodes occur in the top 4 cm, but those that occur deeper seems to be tolerant to anoxic conditions
14	Ansari et al. 1984	M	DT	2	Highest density in the upper tide level for sheltered beach and highest density in the midtide level for exposed beach
15	Ansari et al. 1990	M	DT	6	Meiofauna abundance is reduced in zones of the beach exposed to desiccation
16	Pattnaik and Rao 1990	M	DT	1	Harpacticoid copepods are the dominant group; meiofauna is abundant in the midtide level occuring up to 20 cm deeper; their spatial distribution is controlled by physical factors.
17	Alongi 1990	N	TP	1	Highest nematode densities in austral autumn and winter and lowest densities in spring and summer
18	Ólafsson 1991	M	DT	4	Meiofauna is more abundant in the top 10 cm and nematodes show the most uniform vertical distribution in 95 cm deeper
19	Szymelfenig 1995	M	DT	10	The occurrence of meiofauna taxa is highly correlated to climatic characteristics of the habitats
20	Gourbault et al. 1995	M and N	DT	13	Harpacticoid copepods are more abundant than nematodes; this latter taxa shows higher diversity in coarse sediment
21	Long and Ross 1999	N	DT	1	Nematodes occur at 30cm deep and their vertical distribution is related to food availability
22	Nicholas and Hodda 1999	N	DT	1	Higher nematode abundance in the mid and high tide level of the beach; persistence of nematode zonation during a short period (24h and under calm conditions
23	Armonies and Reise 2000	M	DT	1	Turbellarian is the dominant taxa; highest meiofauna richness at midtide level
24	Nicholas 2001	N	DT	1	Nematodes occur at 60 cm deep; highest densities in the warmer months of the year
25	Rodriguez et al. 2001	M	DT	3	Highest density of the meiofauna in reflective beach and at upper and midtide level
26	Gheskiere et al. 2002	M and N	DT	1	Increase in meiobenthic density towards low tide level; presence of nematode horizontal zonation
27	Menn 2002	M	DT and FW	2	Nematodes are an important food sources for higher trophic level in intermediate beaches while macrofauna plus nematodes are the food sources in dissipative beaches
28	De Oliveira and Soares-Gomes 2003	M	P	1	No relationship between sewage disposal and meiofauna
29	Moellmann and Corbisier 2003	M	P	2	Nematodes migrate to the deeper layers of the sediment in intense trampling areas
30	Souza-Santos et al. 2003	M	DT	1	Physical factors (e.g. grain size, chla, phaeopigments, sediment skewness) are the main factors structuring the meiofauna community seasonally
31	Rodríguez et al. 2003	M	DT	10	Richness and biomass of meiofauna increases toward very exposed and coarse beaches
32	Gheskiere et al. 2004	N	DT	1	Higher nematode density in the low tide level of the beach; high richness in the midtide level; presence of nematode zonation
33	Rodriguez 2004	M	DT	1	Highest meiofauna density in the midtide level
34	Urban-Malinga et al. 2004	M and N	DT	1	Turbellarian is the dominant meiofauna taxa; sheltered beach showed a higher meiofauna density and respiration rates than exposed beach
35	Gheskiere et al. 2005a	N	DT	2	Highest nematode density and diversity in coarse-grained sandy beach; presence of isocommunities only in the upper beach
36	Kotwicki et al. 2005a	M	DT	5	Meiofauna show high density in the upper 5 cm; high densities in the midtide level or in the lower tide level; presence of horizontal zonation
37	Calles et al. 2005	M and N	DT	2	Turbellarian is the dominant taxa in the sheltered beach; high meiofauna density in the exposed beach; no difference between meiofauna diversity of sheltered and exposed beaches
38	Kotwicki et al. 2005b	M	DT	13	Nematodes are dominant in warmer regions of the globe while turbellarian are dominant in cold water regions; no latitudinal trend was found for meiofauna density or diversity
39	Lee and Correa 2005	M	P	12	Reduction of density and diversity in sites impacted by mine tailing.
40	Gheskiere et al. 2005b	M and N	DT and P	4	Lowest species diversity in the upper tide level, especially in touristic beaches
41	Hourston et al. 2005	N	DT	3	Highest density in the sheltered habitat, but declining during winter; weak zonation reflecting the presence of a small tidal range
42	Nicholas and Trueman 2005	N	DT	3	Richness increases with decreasing latitude
43	Urban-Malinga et al. 2005	N	DT	2	Oligochaete is the dominant taxa; highest diversity in sheltered beaches; density increases towards the low water level
44	Moreno et al. 2006	M and N	DT	1	Highest density and diversity in the swash level
45	Harriague et al. 2006	M	DT	1	Higher density in the surfzone than in the swash level
46	Gheskiere et al. 2006	M	CM	1	There is no reduction in density or diversity of mechanical cleaning areas.
47	Urban-Malinga and Moens 2006	M	FW	2	Fastest carbon remineralization in exposed coarse sandy beach
48	Nicholas 2006	M and N	DT	1	Nematodes occur at 60 cm deep
49	Albuquerque et al. 2007	M	DT	1	Tardigrades are the most dominant taxa; highest density in the midtide level
50	Papageorgiou et al. 2007	M	FW	1	Bacteria are a potential food source for meiofaunal organisms
51	De Jesús Navarette 2007	N	DT	2	Highest richness in coarse-grained beach
52	Mundo Ocampo 2007	N	DT	2	Highest density and diversity in the coarse-grained beach
53	Rodríguez et al. 2008	M	DT	10	Eh and water content are the best environmental variables that explains meiofaunal densities
54	Harriague et al. 2008	M	DT	6	Food availability is a key role in structuring meiofauna community
55	Liu et al. 2008	N	DT	1	Physical factors (e.g. temperature, chla and grain size are responsible for the nematode community structure
56	Maria et al. 2008	N	DT	3	Highest richness in very coarse sandy beach
57	Urban-Malinga et al. 2008	N	DT	1	Fastest organic matter weight loss in the highest diverse habitat of the beach - midtide level
58	Delgado et al. 2009	M	DT	23	Gravel and organic matter are best correlated to meiofauna composition and abundance; absence of pattern in meiofaunal abundance along a degree of exposure
59	Grzelak et al. 2009	M	NI	3	Fast recover of meiofauna community after a Tsunami event
60	Gomes and Rosa Filho 2009	M	DT	1	Highest density and diversity in the midtide level; presence of zonation
61	Urban-Malinga and Burska 2009	M	DT	1	Wrack-associated meiofauna community and density from upper and lower shores are similar
62	Veiga et al. 2010a	M	P	9	Low densities in exposed sandy beaches contaminated by polycyclic aromatic hydrocarbons
63	Veiga et al. 2010b	M	DT	2	Highest density in the top 5 cm; oxygen is the limiting factor especially for copepods and ostracods
64	Yamanaka et al. 2010	M	DT	3	Positive correlation between meiofauna abundance and exposure index
65	Gingold et al. 2010	N	DT	1	Highest diversity in the midtide level; presence of similar zonation patterns in topographical heterogeneous habitat
66	Gingold et al. 2011	N	DT	1	Patch distribution is more prone to occur in the runnel habitat than in the sandbar
67	Nanajkar and Ingole 2010	N	P	1	Highest density in organically polluted sites
68	Barnes et al. 2011	M	DT	6	Highest density in the mid and lower tide level; diversity decreases towards the tropics
69	Maria et al. 2011a	M and N	FW	1	Benthic diatoms are the preferential food source of meiofaunal organisms
70	Maria et al. 2011b	N	Co	1	Polychaetes seems to facilitate the colonization of azoic sediments by nematodes
71	Riera et al. 2011	M	TV	1	Total abundance and community structure are similar along a year of study
72	Lee and Riveros 2012	N	DT	66	Richness decreases with increasing latitude
73	Mantha et al. 2012	M	DT	5	Harpacticoid copepods are thedominanttaxa
74	Maria et al. 2012	M and N	DT and FW	1	Biological factors are important in controlling the vertical distribution of nematodes; interstitial and macrofaunal food webs seems do not compete for food sources
75	Riera et al. 2012	M and N	DT	2	Copepods are dominant in medium sand while nematodes are dominant in fine sand, i.e. grain size influences in the meiofauna composition
76	Gingold et al. 2013	N	CC	1	Temperature increase shows a negative impact on dominant predator species
77	Maria et al. 2013a	N	Co	1	Polychaetes did not affect the colonization of azoic sediments by nematodes
78	Maria et al. 2013b	N	DT	1	Runnels and sandbar harbors different communities showing different zonation patterns
79	Maria et al. 2013c	N	DT	2	Highest density in the upper tide level; gravel is the most important variable that explains the nematode community
80	Schlacher and Hartwig 2013	M	FW	1	Meiofauna community is strongly influenced by sediment grain size; the density of nematodes and ostracod is controlled by bottom up processes
81	Kang et al. 2014	M and N	P	1	Reduction of density in oil polluted sites; meiofauna can be recovered one month after an oil spill
82	Kotwicki et al. 2014	M	DT	7	Mean grain size and sorting coefficient showed great influence on meiofauna variation
83	Cooke et al. 2014	M	DT	3	Patch distribution of the meiofauna in the upper tide level, especially harpacticoid copepods and gastrotrichs
84	Netto and Meneghel 2014	M	FW	1	Surfzone diatoms are an important food source for meiofaunal organism of high-energy sandy beaches
85	Sun et al. 2014	M	P	5	Anthropogenic disturbed beaches showed high meiofauna abundance reflecting high chl a and dissolved oxygen levels
86	Venekey et al. 2014a	N	DT	1	Lowest density in dry months; no influence of the tidal cycle in the nematode density
87	Venekey et al. 2014b	N	DT	1	Seasonal and spatial variation of nematode communities in a tropical sandy beach

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