Long-term perspective on the dynamics of brown tide blooms in Long Island coastal bays

Brown tide, a bloom of the picoplankter Aureococcus anophagefferens, first appeared in eastern Long Island (Suffolk County) waters in the late spring of 1985, at about the same time it emerged, although to a lesser degree, in Narraganset Bay, RI. Since then, it has recurred sporadically in Suffolk County, and blooms have been reported in New Jersey, Delaware, Maryland, and only one other area of the world, Saldanha Bay, South Africa. Bloom initiation and maintenance within Suffolk County appear to be related to A. anophagefferens’ ability to use dissolved organic nitrogen (DON) during periods of limited dissolved inorganic nitrogen (DIN) availability. Factors controlling DIN availability include groundwater influx related to meteorological conditions, introduction of septic leachate from on-site wastewater treatment systems, and biological removal. The complexity of bloom dynamics is illustrated by a cascade of events in Great South Bay involving shellfish clearing rates, a macroalgal bloom, and microbial decomposition.     

Pfiesteria piscicida and Pfiesteria shumwayae in coastal waters of Long Island, New York, USA

Water and sediment samples were collected during summer and early fall 1999–2004 from coastal waters of New York State, USA, to test for the presence of Pfiesteria piscicida and Pfiesteria shumwayae. Physical and chemical conditions were characterized, and real-time polymerase chain reaction assays were conducted. Both species were relatively common and found at most sites at least once, and the frequency of positive assays was higher in sediments than in the water column. In a subset of the data from Suffolk County, Long Island, the presence of Pfiesteria was related to high chlorophyll a and relatively high nutrient concentrations. Partial SSU rDNA sequences of four PCR amplicons generated using P. shumwayae primers indicated two sequences: three were identical to GenBank P. shumwayae entries, but one showed enough sequence difference (15 positions in a 454 bp amplicon) to suggest a possible new species. Three isolates were tested for toxicity, and one was found to kill fish in bioassays. Despite the widespread presence of both Pfiesteria species and demonstration of potential to harm fish, no blooms of these dinoflagellates have been observed, nor has there been evidence of Pfiesteria-related fish or human health problems in these waters, likely related to colder temperatures than optimal for Pfiesteria species.     

Genetic analyses of the Asian longhorned beetle (Coleoptera,Cerambycidae, <Emphasis Type="Italic">Anoplophora glabripennis</Emphasis>), in North America,Europe and Asia

The Asian longhorned beetle, (Coleoptera, Cerambycidae, Anoplophora glabripennis (Motschulsky)), is endemic to China and Korea and an important invasive insect in North America and Europe. We analyzed mitochondrial DNA sequence data of invasive populations of A. glabripennis in North America and Europe, and microsatellite allele frequency data of beetles from North America. We show that populations in New York City and Long Island NY; New Jersey, Chicago, IL, and Toronto, Canada have limited genetic diversity compared to populations in China. In addition, the data suggest that separate introduction events were responsible for many of the populations in North America and for European populations in Austria, France, Germany and Italy. Populations on Long Island, NY are suspected to have been initiated by the transport of cut wood from New York City. A. glabripennis beetles found in Jersey City, NJ appear to be derived from an expansion of the New York City, NY population, whereas beetles found in Linden, NJ are an expansion from the Carteret, NJ population. Limited genetic diversity did not stop this invasive insect from establishing damaging populations in North America. Founders of introduced A. glabripennis populations in North America and Europe are likely derived from populations in China that are themselves invasive, rendering difficult the identification of source populations. Invasiveness in an insect’s natural range could be an important predictor of potential pest status of introduced populations.     

An immunofluorescent survey of the brown tide chrysophyte Aureococcus anophagefferens along the northeast coast of the United States

Surveys were conducted along the northeast coast of the USA.between Portsmouth, NH, and the Chesapeake Bay in 1988 and 1990,to determine the population distribution of Aureococcus anophagefferens,the chrysophyte responsible for massive and destructive ‘browntides’ in Long Island and Narragansett Bay beginning in1985. A species-specific immunofluorescent technique was usedto screen water samples, with positive identification possibleat cell concentrations as low as 10–20 cells ml^–1.Both years.A.anophagefferens was detected at numerous stationsin and around Long Island and Barnegat Bay, NJ, typically athigh cell concentrations. To the north and south of thus ‘center’,nearly half of the remaining stations were positive for A.anophagefferens,but the cells were always at very low cell concentrations. Manyof the positive identifications in areas distant from Long Islandwere in waters with no known history of harmful brown tides.The species was present in both open coastal and estuanne locations,in salinities between 18 and 32 practical salinity units (PSU).The observed population distributions apparently still reflectthe massive 1985 outbreak when this species first bloomed, giventhe number of positive locations and high abundance of A.anophagefferensin the immediate vicinity of Long Island. However, the frequentoccurrence of this species in waters far from this population‘center’ is disturbing. Aureococcus anophagefferensis more widely distributed than was previously thought. Numerousareas thus have the potential for destructive brown tides suchas those associated with the sudden appearance of the speciesin 1985.     

Zostera marina population genetics in Barnegat Bay, New Jersey, and implications for grass bed restoration

Within Barnegat Bay, New Jersey, Zostera marina populations have declined by 62% over the last 20 years, and restoration efforts have met with mixed success. We have completed a microsatellite-based genetic investigation of eight populations of Z. marina within Barnegat Bay to determine whether the genetic stock origins of the plants used in management projects may affect restoration success. Additionally, we assessed the genetic diversity of Z. marina in Barnegat Bay to better understand its population structure. Clonal diversity ranged from 0.70 to 0.95 for the populations studied. Individually, Barnegat Bay populations are not genetically diverse, and there is also little divergence among populations. The Atlantic populations had mean Hobs values (0.20–0.34) that were far lower than the Hexp values (0.69–0.83). Also, the F _IS values in all of the eastern populations indicate a surfeit of homozygotes over heterozygotes, suggesting a low degree of outcrossing in the Barnegat Bay populations. Six of the ten populations studied (Ham Island, Manahawkin Bay, Shelter Island, Marsh Elder, Harvey Cedar Sedge, and Long Island) show evidence of historical bottlenecks. Mean estimated F _ST values would suggest that most alleles are undergoing moderate genetic differentiation, with values that range from 0.06 to 0.13. Oyster Creek and Sedge Island demonstrate the largest estimated effective population sizes and may be the most appropriate populations for use in future eelgrass restoration projects.     

Pollution in Peter the Great Bay, Sea of Japan, and its biological consequences

This review summarizes information published in the 1980s–1990s about anthropogenic pollution in Peter the Great Bay, the largest of the bays in the northwestern part of the Sea of Japan. The coastal zone of the bay occupies about 12% of the area of Primorskii Province and is the most heavily populated. The bulk of the human settlements, the seaports of Vladivostok and Nakhodka, railways, industrial enterprises, and developed agriculture are located in the coastal zone. Sewage waters containing multicomponent mixtures of polluting agents of both mineral and organic origin are discharged into the coastal waters of the bay. This paper presents information about the concentration of major classes of polluting agents (oil hydrocarbons, polychlorinated hydrocarbons, surfactants, heavy metals, and radionuclids) in the water and bottom sediments of the bay. The results of physico-chemical and biogeochemical investigations performed in the 1970s–1990s justify considering Zolotoi Rog Bay, Bosfor Vostochnyi Strait, Nakhodka Bay (especially its innermost portion, around the harbor of Nakhodka), and Amurskii Bay to be the most polluted areas in Peter the Great Bay. The information about the biological consequences of pollution in these water areas is briefly reviewed.     

Environmental investigations of Vibrio parahaemolyticus in oysters after outbreaks in Washington, Texas, and New York (1997 and 1998)

Total Vibrio parahaemolyticus densities and the occurrence of pathogenic strains in shellfish were determined following outbreaks in Washington, Texas, and New York. Recently developed nonradioactive DNA probes were utilized for the first time for direct enumeration of V. parahaemolyticus in environmental shellfish samples. V. parahaemolyticus was prevalent in oysters from Puget Sound, Wash.; Galveston Bay, Tex.; and Long Island Sound, N.Y., in the weeks following shellfish-associated outbreaks linked to these areas. However, only two samples (one each from Washington and Texas) were found to harbor total V. parahaemolyticus densities exceeding the level of concern of 10,000 g(-1). Pathogenic strains, defined as those hybridizing with tdh and/or trh probes, were detected in a few samples, mostly Puget Sound oysters, and at low densities (usually <10 g(-1)). Intensive sampling in Galveston Bay demonstrated relatively constant water temperature (27.8 to 31.7 degrees C) and V. parahaemolyticus levels (100 to 1,000 g(-1)) during the summer. Salinity varied from 14.9 to 29.3 ppt. A slight but significant (P < 0.05) negative correlation (-0.25) was observed between V. parahaemolyticus density and salinity. Based on our data, findings of more than 10,000 g(-1) total V. parahaemolyticus or >10 g(-1) tdh- and/or trh-positive V. parahaemolyticus in environmental oysters should be considered extraordinary.     

Coliphage and indigenous phage in Mamala Bay, Oahu, Hawaii.

Public concern over the discharge of primarily treated sewage by two offshore outfalls in Mamala Bay, Oahu, prompted a multidisciplinary study to determine the impact of such activities on the water quality in the bay and at adjacent recreational beaches. As part of this study, we determined the abundance of coliphage as an indicator of fecal pollution along with total viral direct counts and phages infective for Vibrio parahaemoltyicus 16 at stations in Mamala Bay in four quarterly samplings over 13 months. Coliphage (< 1 to 1.2 x 10(3)/liter) were found during each quarterly sampling along an offshore transect to the Sand Island waste treatment facility outfall. The nonpoint coastal stations (Pearl Harbor, Ala Wai Canal, and Ke'ehi Lagoon) had high levels of coliphage during the storm event sampling in February 1994 but much lower levels or none when sampled during dry weather. Coliphage were absent at all samplings at Waikiki Beach and at the control station off Diamond Head. Viral direct counts in eutrophic coastal stations (Pearl Harbor, Ke'ehi Lagoon, Ala Moana Beach, and Ala Wai canal) averaged 10(9)/liter, while counts at offshore stations ranged from 9 x 10(7) to 1 x 10(9) viruses/liter, values similar to those for other marine environments. Vibriophage were found mainly in eutrophic coastal environments (Ala Wai Canal, Pearl Harbor, and Ke'ehi Lagoon) and at the Sand Island Transect stations D1 and D2. The greatest abundance was found during the storm event (February 1994) sampling. These results suggest that the Sand Island outfall influenced the water quality of the immediate surrounding waters but had little effect on the quality of the recreational beaches. Nonpoint discharge sources appeared to be more important in the distribution of fecal indicators in the coastal zone.     

Microelement composition of the green alga Ulva fenestrata from Peter the Great Bay, Sea of Japan

Concentrations of heavy metals Fe, Mn, Cu, Zn, Pb, Cd, and Ni were determined in the thalluses of the green alga Ulva fenestrata sampled from different locations in Peter the Great Bay (Sea of Japan). According to the metal concentrations in Ulva, the degree of pollution of the surveyed areas in Peter the Great Bay decreases in the following series: Amur Bay > Ussuri Bay > Nakhodka Bay > Vostok Bay > the water area of Far Eastern State Marine Nature Biosphere Reserve. The microelement composition of Ulva from open-shore stations reflects the heavy metal pollution level of water areas as a whole. The concentrations of trace elements in U. fenestrata from closed coastal areas are indicative of marine coastal water pollution from local sources. Generally, metal concentrations in U. fenestrata from Peter the Great Bay are similar to heavy metal levels in non-polluted or weakly polluted coastal areas of the world.     

Use of CUSUM and Shewhart charts to monitor regional trends of birth defect reports in New York State

BACKGROUND: Cumulative sum (CUSUM) charts were originally developed for industrial quality control, but may be adapted for the surveillance of health outcome data, such as birth defects. The reported prevalence of birth defects can vary due to differences in case ascertainment, surveillance practices, or true changes in prevalence. We examined the utility of CUSUM and Shewhart charts for detect-ing changes in prevalence of two different birth defect groups. We chose obstructive renal defects because we expected an increase in reporting due to improved diagnosis. We chose oral clefts for comparison because we expected reporting to be unaffected by changes in diagnostic technologies. METHODS: Data from the New York State Congenital Malformations Registry from 1992-1999 were analyzed using self-starting binomial CUSUM and Shewhart charts for four regions of New York State. RESULTS: CUSUM charts show that reports of obstructive urinary defects have increased from 1992-1999 in all regions of New York State. Reports of oral clefts increased only on Long Island. CONCLUSIONS: The CUSUM method proved useful for identifying changes in birth defect reporting and was able to detect the expected increases in obstructive renal defects. The apparent increase is likely due to improvements in diagnostic imaging techniques. In contrast, we only detected an increase in oral clefts on Long Island, which may be related to under report-ing of cases in the earlier years. CUSUM charts are useful in detecting small, sustained increases in prevalences over time while Shewhart charts are easier to interpret and can detect large sharp increases.     

A New PCR-Based Method Shows That Blue Crabs (Callinectes sapidus (Rathbun)) Consume Winter Flounder (Pseudopleuronectes americanus (Walbaum))

Winter flounder (Pseudopleuronectes americanus) once supported robust commercial and recreational fisheries in the New York (USA) region, but since the 1990s populations have been in decline. Available data show that settlement of young-of-the-year winter flounder has not declined as sharply as adult abundance, suggesting that juveniles are experiencing higher mortality following settlement. The recent increase of blue crab (Callinectes sapidus) abundance in the New York region raises the possibility that new sources of predation may be contributing to juvenile winter flounder mortality. To investigate this possibility we developed and validated a method to specifically detect winter flounder mitochondrial control region DNA sequences in the gut contents of blue crabs. A survey of 55 crabs collected from Shinnecock Bay (along the south shore of Long Island, New York) in July, August, and September of 2011 showed that 12 of 42 blue crabs (28.6%) from which PCR-amplifiable DNA was recovered had consumed winter flounder in the wild, empirically supporting the trophic link between these species that has been widely speculated to exist. This technique overcomes difficulties with visual identification of the often unrecognizable gut contents of decapod crustaceans, and modifications of this approach offer valuable tools to more broadly address their feeding habits on a wide variety of species.     

Sequencing and Amplified Restriction Fragment Length Polymorphism Analysis of Ribonucleotide Reductase Large Subunit Gene of the White Spot Syndrome Virus in Blue Crab (Callinectes sapidus) from American Coastal Waters

In the present study, the existence of white spot syndrome virus (WSSV) in blue crab (Callinectes sapidus) collected from 3 different American coastal waters (New York, New Jersey, and Texas) was confirmed by 2-step diagnostic polymerase chain reaction and in situ hybridization analysis. When geographic isolates were also compared using a gene that encodes the WSSV ribonucleotide reductase large subunit RR1 (WSSV rr1), a C¹⁶⁶¹-to-T point mutation was found in the New Jersey WSSV isolated. This point mutation, which resulted in the creation of an additional RsaI endonuclease recognition site, was not found in the WSSV from the New York and Texas blue crab samples, or in the WSSV Taiwan isolate, or in any of the other WSSV geographical isolates for which data are available. WSSV rr1-specific RsaI amplified restriction fragment length polymorphism of an amplified 1156-bp fragment thus distinguished the New Jersey blue crab samples from the other WSSV isolates. Received June 29, 2000; accepted October 11, 2000     

Temporal variations in phytoplankton, particulates, and colored dissolved organic material based on optical properties during a Long Island brown tide compared to an adjacent embayment

Since 1985, the coastal embayments of Long Island, New York, have been plagued with recurrent blooms, aptly called brown tides, of the pelagophyte Aureococcus anophagefferens. The distinct ocean color observed during these blooms suggests that optical methods can be used as a tool to study, detect, and track brown tides. Thus, the goal of our project was to compare the optical properties and pigment composition during bloom and non-bloom conditions and assess temporal variations in the phytoplankton and other constituents in the seawater associated with bloom development. From 17 May to 8 June 2000, we measured a time series of particle size distributions and concentrations as well as size-fractioned algal pigments and optical properties in two Long Island embayments where brown tides are known to occur. During our study, A. anophagefferens represented an insignificant contribution to the algal community in West Neck Bay (WNB), whereas a bloom developed in Quantuck Bay (QB). Initially, temperature and salinity were similar at the two locations; however, bulk optical properties, chlorophyll, and particle concentrations were nearly a factor of 2 greater at QB. Bulk optical properties remained constant at WNB, yet increased exponentially at QB as the bloom developed. The composition of particulates, including phytoplankton, varied little at QB, and the optical properties suggested the dominance of A. anophagefferens (confirmed by microscopy). The largest temporal variations were observed in the colored dissolved organic material (CDOM); the colloidal (0.2–0.7 μm) fraction, exhibiting a strong protein-like signal, increased dramatically at the height of the bloom. At WNB particle sizes and algal composition varied despite the invariant bulk optical properties; CDOM variations were minimal. Overall, the optical properties in the two bays demonstrated that at QB temporal variations were dominated by biomass and colloidal protein changes, whereas shifts in the algal community occurred at WNB. This study demonstrates the utility of in situ optical observations to resolve temporal changes in the ecological conditions associated with algal bloom development.     

大亚湾水质遥感监测指标分析

我国沿海经济高速发展,对近海岸水质环境影响极大。传统的近海岸水质监测手段不仅成本大,而且空间信息有限,难以对相关海域进行全面、准确的监测与评价;利用遥感的手段进行近海岸水质监测可克服这些局限。利用遥感进行水质评价,需要建立相应的指标体系。论文以大亚湾为例,分析了大亚湾的水质环境现状,讨论了具有光学敏感性或热敏感性的水质因子特性,指出水温、悬浮物及叶绿素是大亚湾水质遥感监测与评价的首选指标。讨论了悬浮物、叶绿素的污染表征物特征,以及在近海岸、湖泊及河流水质研究中的重要指示意义,同时建议将叶绿素作为常规监测指标纳入我国海水水质评价体系。     

Elevated organic nutrient ratios associated with brown tide algal blooms of Aureococcus anophagefferens (Pelagophyceae)

Natural assemblages of Aureococcus anophagefferens collectedfrom the coastal bays of Maryland and Long Island, New Yorkoccurred under water quality conditions higher in dissolvedorganic carbon and phosphorus (relative to nitrogen) than thosesame stations when Aureococcus was not present in significantnumbers. Additional work, both in culture and in the field,confirmed the importance of a heterotrophic nutritional modein this brown tide organism.     

Mixed‐stock analysis of wintertime aggregations of striped bass along the Mid‐Atlantic coast

Most larger individuals of migratory striped bass Morone saxatilis from the two major Atlantic coast stocks, the Chesapeake Bay and Hudson River, appear to winter in mid‐Atlantic coastal waters. But it is not known whether they exhibit differential wintertime distributions in accordance with the latitudinal differences in locations of these two estuaries. Mixed‐stock analyses were conducted based on mitochondrial DNA and nuclear DNA genotypic frequencies on wintertime collections of striped bass from coastal waters. No significant differences (P > 0.05) were seen in the proportions of striped bass from the two stocks between collections made from the Delaware Bay mouth and Cape Hatteras in 1997. However, there was a substantially higher Hudson contribution to a 1995 collection from coastal New Jersey (0.349, SD = 0.136) than to the combined 1997 Delaware Bay mouth and Cape Hatteras collection (0.157, SD = 0.072), suggesting this question deserves further study. Additionally, use of the original four reference samples from Chesapeake Bay tributaries (Choptank, Potomac, Rappahannock, Upper Bay) proved adequate alone in characterizing the Chesapeake Bay stock in simulations in which additional tributary collections (Nanticoke, Patuxent, Pocomoke) were added.     

Assessment of brown tide blooms, caused by Aureococcus anophagefferens, and contributing factors in New Jersey coastal bays: 2000–2002

A 3 year study (2000–2002) in Barnegat Bay-Little Egg Harbor (BB/LEH), New Jersey (USA), was conducted by the New Jersey Department of Environmental Protection, Division of Science Research and Technology (DSRT) in cooperation with several partners to assess brown tide blooms in coastal waters in NJ. Water samples were collected by boat and helicopter at coastal stations from 2000 to 2002 along with field measurements. Aureococcus anophagefferens were enumerated and associated environmental factors were analyzed. A. anophagefferens abundances were classified using the Brown Tide Bloom Index and mapped, along with salinity and temperature parameters, to their geo-referenced location using the ArcView GIS. The highest A. anophagefferens abundances (>10⁶ cells ml⁻¹), including category 3 blooms (≥200,000 cells ml⁻¹) and category 2 blooms (≥35,000 to ≤200,000 cells ml⁻¹), recurred during each of the 3 years of sampling and covered significant geographic areas of the estuary, especially in Little Egg Harbor. While category 3 blooms were generally associated with warmer water temperatures (>16 °C) and higher salinity (>25–26 ppt), these factors were not sufficient alone to explain the timing or distribution of A. anophagefferens blooms. There was no significant relationship between brown tide abundances and dissolved organic nitrogen measured in 2002 but this was consistent with other studies. Extended drought conditions, with corresponding low freshwater inputs and elevated bay water salinities, occurring during this time were conducive to blooms. A. anophagefferens abundances were well above the reported levels that have been reported to cause negative impacts on shellfish. It was shown that over 50% of the submerged aquatic vegetation (SAV) habitat located in Barnegat Bay/Little Egg Harbor was categorized as having a high frequency of category 2 or 3 blooms for all 3 years.     

Microplankton and its functional activity in zones of shallow hydrotherms in the Western Pacific

Water above the active underwater volcanoes on the slopes ofEpi Island, New Hebrides, Raul Island, Kermadek Island and the‘Calypso’ gas anomaly area, Bay of Plenty, New Zealand,being enriched with methane and hydrogen, contains abundantbacterioplankton with a biomass several times greater than thesurrounding oceanic waters. Dark microbial CO₂ uptake in thesesites was 10–30 times greater than in adjacent oceanicwaters. In the shallow underwater volcanic hydrotherm areasaround Whale and White Islands (New Zealand), as well as inMatupi Harbour Bay (New Britain Island), significant rates ofmicrobial chemosynthesis were recorded in the water column (5–10mg C m^–3 day^–1. This was reflected in high microbialbiomass: 20–60 mg C m^–3. Over the field of algal-bacterialmats, it exceeded 100 mg C m^–3. The cell volume of bacteriagrowing in waters influenced by hydrothermal activity was 2–4times greater than that of those growing in sea areas remotefrom hydrotherms. Primary phytoplankton production in thesewaters was significantly greater than in surrounding oceanicwaters. A high activity of hydrogen- and methane-oxidizing bacteria,as well as thiobacilli, was also recorded there. The microzooplanktonin areas of the sea influenced by hydrotherms (Whale Island)contained a large percentage of planktonic amoebae, up to 30x 10³ I^–1. The probable ways in which shallow hydrothermaldischarges influence the biota of coastal waters are discussed.     

Cryptic Diversity in Metropolis: Confirmation of a New Leopard Frog Species (Anura: Ranidae) from New York City and Surrounding Atlantic Coast Regions

We describe a new cryptic species of leopard frog from the New York City metropolitan area and surrounding coastal regions. This species is morphologically similar to two largely parapatric eastern congeners, Rana sphenocephala and R. pipiens. We primarily use bioacoustic and molecular data to characterize the new species, but also examine other lines of evidence. This discovery is unexpected in one of the largest and most densely populated urban parts of the world. It also demonstrates that new vertebrate species can still be found periodically even in well-studied locales rarely associated with undocumented biodiversity. The new species typically occurs in expansive open-canopied wetlands interspersed with upland patches, but centuries of loss and impact to these habitats give some cause for conservation concern. Other concerns include regional extirpations, fragmented extant populations, and a restricted overall geographic distribution. We assign a type locality within New York City and report a narrow and largely coastal lowland distribution from central Connecticut to northern New Jersey (based on genetic data) and south to North Carolina (based on call data).     

Genetic structure in a coastal dune spider (Geolycosa pikei) on Long Island, New York Barrier Islands

In this study, we analyse genetic structure among ten isolated populations of a sedentary burrowing wolf spider, Geolycosa pikei, collected from Long Island, New York barrier islands. Using allozyme electrophoresis to estimate gene frequencies at 12 loci, only four loci were polymorphic (i.e. Aat, Apk, Gpi, Idh-1), and populations showed little differentiation (mean F_sr=0.020). Contrary to stepping-stone model predictions, the matrix of genetic distances among sites was not significantly associated with the physical distance matrix, which suggests that G. pikei on Long Island barriers does not adhere to a stepping-stone model. Geolycosa pikei may be better dispersers than previously estimated and/or the ephemeral quality of the inlets may not allow for the development of significant inter-populational genetic differences. In addition, geologic evidence suggests that these barriers became relatively stable only 8000 BP. Therefore, these habitats have been only recendy colonized, which may result in low genetic variability and inter-populational differentiation possibly due to genetic drift from repeated and prolonged bottlenecks during recolonization (i.e. founder events).