Metrics details
Insecticide resistance is a serious problem for vector control programmes worldwide
Resistance is commonly attributed to mutations at the insecticide’s target site or increased activity of detoxification enzymes
We determined the knockdown concentration (KC50) and lethal concentration (LC50) of deltamethrin in six natural populations of adult Aedes aegypti from southeastern Mexico
These populations were then selected over five generations using the LC50 from the preceding generation that underwent selection
and the heritability of deltamethrin resistance was quantified
we also determined the frequency of the kdr alleles L410
and the levels of activity of three enzyme families (α- and β-esterases
mixed-function oxidases and glutathione S-transferases (GST)) associated with insecticide detoxification
There was an increase in KC50 and LC50 values in the subsequent generations of selection with deltamethrin (FS5vs FS0)
we detected increases in LC50 ranging from 1.5 to 5.6 times the values of the parental generation and in KC50 ranging from 1.3–3.8 times the values of the parental generation
Triple homozygous mutant individuals (tri-locus
LL/II/CC) were present in the parental generations and increased in frequency after selection
The frequency of L410 increased from 1.18-fold to 2.63-fold after selection with deltamethrin (FS5vs FS0) in the populations analyzed; for I1016 an increase between 1.19-fold to 2.79-fold was observed
and C1534 was fixed in all populations after deltamethrin selection
Enzymatic activity varied significantly over the generations of selection
only α- esterase activity remained elevated in multiple populations after five generations of deltamethrin selection
We observed an increase in the mean activity levels of GSTs in two of the six populations analyzed
The high levels of resistance and their association with high frequencies of kdr mutations (V410L
V1016I and F1534C) obtained through artificial selection
suggest an important role of these mutations in conferring resistance to deltamethrin
We highlight the need to implement strategies that involve the monitoring of kdr frequencies in insecticide resistance monitoring and management programmes
Given the background levels of pyrethroid and DDT resistance in many parts of Mexico
the extent to which ongoing insecticide pressure selects for specific mechanisms of resistance is not well understood
To better understand how insecticide exposure selects for resistance we evaluate the effect of laboratory selection with deltamethrin on the underlying resistance mechanisms in populations of Ae
Collection sites of Aedes aegypti from Veracruz and Yucatan states from Mexico
The biological material obtained from the field was reared under insectary conditions at 26 ± 2 °C and 70–80% relative humidity with a 12h:12h (light:dark) photoperiod
The adults obtained from field-collected material were allowed to intermate
and the eggs resulting were designated FS0 (without previous selection)
The New Orleans strain (NO) was used as a susceptible reference in the study
this strain was originally obtained from the CDC (Atlanta
Each bioassay consisted of bottles containing different concentrations of deltamethrin resulting in mortalities between 9–90%
with at least three replicates for each concentration
The numbers of knocked-down mosquitoes were recorded at 1 h
all mosquitoes were gently transferred to a recovery container without insecticide and were offered a cotton ball soaked in a sugar solution
Both the bottles and recovery containers were kept at 24 ± 2 °C and 70% RH
The selection was carried out exposing 650–1000 FS1 males and females (1:4) for 1 h to the LC50 obtained for the previous generation (FS0) by the method described above
The survivors at 24 h were transferred to mosquito cages for breeding the subsequent generation (FS2)
The dose response parameters were calculated again for the FS1 cohort and were used to select the next generation (FS2)
This procedure was repeated for each generation of selection until FS5
The New Orleans strain was tested at the same time as the FS0 and was not tested in each generation of selection
KC50 and LC50 values were compared between generations to monitor changes in resistance
The KC50 and LC50 values were considered significantly different if their 95% confidence intervals did not overlap
A low h2 predicts no additive genetic variance for a trait and a poor or very slow response to artificial selection
while a high h2 predicts a large additive genetic variance at one or a few loci that govern a trait and predicts a rapid response to artificial selection
and the DNA pellet was resuspended in 30 μl of ultrapure molecular grade water (Corning CellgroTM
The quantity and quality of DNA were determined using a NanoDrop spectrophotometer 2000 (Thermo Fisher Scientific
a master mix was prepared as follows: 12.50 μl of GoTaq (Promega
USA) was mixed with 11.35 μl of nuclease-free H2O (NFW
The master mix (24 μl) was then placed in 0.2 ml stoppered tubes followed by 1 μl of mosquito genomic DNA (~25 ng)
The tubes were centrifuged for 1 min at 3300× rpm
then placed in a T100 thermocycler (Bio-Rad) with the following temperature program: 3 min at 95 °C; 30 cycles of 1 min at 95 °C
20 s at 56 °C and 20 s at 72 °C; followed by 5 min at 72 °C
DNA from the New Orleans susceptible strain was used as a negative control for the kdr assays
and previously genotyped individuals were used as positive controls
V1016I and F1534C assays were visualized on 2.5%
To measure the activity levels of α- and β-esterases
100 μl of either α- or β-naphthyl acetate (Sigma-Aldrich
MO,USA) dissolved in acetone (CTR Scientific
pH 7.2) was added to each well and incubated for 20 min at room temperature
100 μl of fast blue (tetrazotized O-dianisidine (Sigma-Aldrich) dissolved in distilled water) was added to each well and the microplates incubated for another 4 min
before absorbance was read at 540 nm using a spectrophotometer (ASYS Hitech GmbH
200 μl of TMBZ (3,3,5,5-tetramethyl-benzidine dihydrochloride (Sigma-Aldrich) previously dissolved in methanol (Jalmek
Mexico) and 0.25 M sodium acetate buffer pH 5.0) were added to each well
Subsequently 25 μl of 3% hydrogen peroxide (H2O2) (Jalmek) was added to each well
After 10 min of incubation at room temperature the microplate was read at a wavelength of 620 nm
100 μl of reduced glutathione (GSH; Sigma-Aldrich) and 100 μl 1-chloro-2,4 dinitrobenzene (CDNB; Sigma-Aldrich) previously diluted in acetone and KPO4 buffer were added to each well
Absorbance was read at 340 nm immediately (T0) and after 10 min of incubation (T10)
The values obtained after subtracting the initial reading (T0) from the reading at 10 min (T10) were used for the statistical analyses
Positive and negative controls were included for MFOs and esterases
The same volume of homogenate which was used in the respective assays was used in running the controls
α- and β -naphthyl acetate solution was used as positive controls
Germany) solution was the positive control for the MFO assay
KPO4 buffer was used as a negative control for each biochemical assay
The calibration curves were generated using control solutions for each enzyme family
α- or β-naphthol (Sigma-Aldrich) was used for α-esterases and β-esterases
with a concentration range of 0.3 to 5 μg/μl and 0.5 to 4 μg/μl
Cytochrome C concentrations from 0.002 to 0.65 μg/μl were used for MFOs
and 0.1 to 6.5 μg/μl BSA solutions were used for total protein determination
The mean activity values underwent an analysis of variance (ANOVA, P < 0.05) and Tukey’s multiple comparison of means (P < 0.05). Normality of the variance was verified by the Bartlett test. Statistical analyses were carried out with GraphPad Prism v.7 (GraphPad Software, Inc, Version 6.01, La Jolla, CA, USA; https://www.graphpad.com)
Activity levels for all enzymes were determined in the parental population and in each deltamethrin-selected generation (FS0–FS5)
the LC50 values and mean enzyme activity values of the parental generation and each selected generation (FS1–FS5) underwent linear regression analysis
Correlation coefficients were calculated to determine the degree of association between the two variables
When comparing the LC50 values obtained from the parental generations to the NO strain, five of the populations (Merida, Progreso, Hoctun, Agua Dulce and Jose Cardel) were highly resistant to deltamethrin with an RRLC50 of 18–134-fold, and only one population (Hunucma) showed moderate resistance (RRLC50 of 6-fold) (Table 3).
a Knockdown concentrations values (KC50) as response to the selection with deltamethrin
b Lethal concentrations values (LC50) as response to the selection with deltamethrin in six populations of Aedes aegypti through five generations
With respect to knockdown in the parental generation, RRKC50 was high in two populations, Merida with RRKC50 of 20-fold and Jose Cardel with RRKC50 of 14-fold. Three populations (Hoctun, Progreso and Agua Dulce) showed moderate resistance with RRKC50 ranging between 5–9-fold, and one population (Hunucma) had a low RRKC50 of 3-fold (Table 2)
When comparing the LC50 and KC50 values for the last deltamethrin-selected generation (FS5) vs the parental generation (FS0), we observed an increase of ~1.5–5.5 times for LC50 and ~1.4–2.7 times for KC50 in all populations (Fig. 2)
The knockdown resistance ratio (RRKC50) value increased from 20-fold to 38-fold from FS0-FS5 in the Merida population
and slight increases were observed in the Agua Dulce and Jose Cardel populations with 9–12-fold and 14–22-fold differences
Lethal concentrations also increased between FS0 and FS5
with the RRLC50 increasing in the Hunucma populations from 6-fold to 34-fold
doubling in the Agua Dulce and Progreso populations (25–49-fold and 18–36-fold
respectively) and with lower increases detected in the Merida
Hoctun and Jose Cardel populations (134–197-fold
When analyzing the relative increases in RRKC50 and RRLC50 during the selection
the Hunucma population showed the highest single-generation increase (> 0.3-fold) from FS1vs FS2 for KC50; Hunucma showed also the highest increase in RRLC50 (2.83) in the first selected generation relative to the parental generation (FS1vs FS0)
Tables 6, 7, 8 summarize the frequency of L410
I1016 and C1534 alleles across all populations and generations (FS0–FS5)
The three kdr mutations were present in all the parental populations
with frequencies ranging between 0.36–078 for the L410 allele and 0.34–0.77 for I1016
All populations were in Hardy-Weinberg equilibrium at the parental generation (FS0)
with the exception of Jose Cardel for the mutation F1534C
Frequencies of the tri-locus genotypes in the parental generation (FS0) of each population of Aedes aegypti. Genotypes order: 410/1016/1534
Frequencies of the resistant trilocus genotype LL410/II1,016/CC1,534 across selection with deltamethrin (FS0-FS1) from six populations of Aedes aegypti from Mexico
the triple heterozygous genotype (VL410/VI1016/FC1534) was only found in the populations of Hunucma
in which the triple wild-type genotype (VV410/VI1016/FC1534) was also found
the frequencies of the three kdr alleles increased over the selected generations
The frequencies of L410 from the parental generation to the last selected generation increased from 0.78 to 0.92 in the Merida population
0.55 to 0.98 in Agua Dulce and 0.66 to 0.80 in Jose Cardel
increases of 0.77–0.92 were observed in the Merida population
0.56–0.96 in Agua Dulce and 0.63–0.92 in Jose Cardel
The frequencies of the C1534 allele increased from 0.59 to 1 in the Hoctun population
0.89 to 1 in Jose Cardel and 0.97 to 1 in Progreso; the C1534 allele was fixed at 1.0 in the Merida and Agua Dulce populations in all generations
This allele reached fixation after the first selection with deltamethrin (FS1) in the population from Progreso and Jose Cardel
in the third selected generation for Hoctun and in the fourth selected generation for Hunucma
Allele frequencies of L410 (a), I1016 (b) and C1534 (c) and their maximum and minimum frequencies in six Aedes aegypti populations across selection with deltamethrin (FS0-FS5)
Regression analysis of allele frequencies of two para mutations: I1016 vs L410 across the selection with deltamethrin (FS0-FS5) in Aedes aegypti from Merida (a), Hunucma (b), Hoctun (c), Agua dulce (d)
Regression analysis of allele frequencies of C1534 vs I016 (a
c) across the selection with deltamethrin (FS0-FS5) in Aedes aegypti from Hunucma and Hoctun
The frequency of the mutant alleles for each mutation was correlated with the KC50 and LC50 values across the generations of selection (Table 9)
KC50 values were significantly correlated with L410 frequencies in the Merida and Hoctun populations (P < 0.05); and with the frequencies of I1016 in the Merida
the frequencies of L410 and I1016 were significantly correlated with LC50 values in the Merida and Hunucma populations as well as C1534 correlated with LC50 values in Hunucma and Jose Cardel (P < 0.05)
The presence and frequency of triple homozygous mutant individuals showed a significant correlation with KC50 only in the Merida population and LC50 in the Hunucma population (P < 0.05)
Table 10 shows the mean enzyme activity levels detected in the biochemical assays in the parental generation and all deltamethrin-selected generations
where at least 30 individuals per population were analyzed
When comparing values from the parental generations with the 99th percentile of the New Orleans reference strain
five of the six populations analyzed (Merida
Jose Cardel and Agua Dulce) showed altered enzyme activity for α-esterases (˃ 50% of the individuals exceeded the threshold of resistance)
and Hoctun showed incipient altered enzyme activity (39% of the individuals exceeded the threshold)
Agua Dulce and Jose Cardel showed altered enzyme activity for β-esterases (˃ 50%)
Hunucma and Hoctun showed incipient enzyme alteration with percentages ranging between 16–43%
Altered activity of MFOs was detected in Progreso
Agua Dulce and José Cardel in 69–95% of the specimens analyzed
Only the Agua Dulce population showed altered enzyme activity for GSTs (71%) in the parental generation
GSTs were the only enzyme family for which a significant correlation was detected between level of activity and LC50 values
five of the populations analyzed were found to be highly resistant to the pyrethroid deltamethrin in the parental generation according to the LC50 (RRLC50 > 10-fold)
and only one population showed moderate resistance (RRCL50 5–10-fold)
high RRKC50 values were detected in two populations
These results suggest that deltamethrin resistance is well-established in these populations
who reported increased levels of MFOs in response to deltamethrin selection
we found that MFO activity decreased with successive selection
Given that the frequencies of the kdr mutations increased significantly with deltamethrin selection
the rapid selection of these mutations could have conditioned the activity of the enzymes in relation to deltamethrin resistance
One criticism in our methodology is that we compare the enzymatic activity of each generation of deltamethrin in each of the populations with respect to the enzymatic activity of the susceptible New Orleans strain
when the most appropriate comparison should be with respect to the same population
where the significant participation of an esterase marker discovered by QTL
and its relationship to resistance development in conjunction with mutations in the voltage-gated sodium channel was demonstrated
as for this mechanism to be in itself the one that confers insecticide resistance is questionable since other mechanisms were described for these populations
All data generated or analyzed during this study are included in the published article
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We would like to thank Kevin Ontiveros-Zapata (Universidad Autonoma de Nuevo Leon
Mexico) for processing mosquitoes in the laboratory
The findings and conclusions in this paper are those of the authors and do not necessarily represent the official position of the Centers for Disease Control and Prevention
This study was financially supported by CONACYT Mexico through the fund Problemas Nacionales
Division of Parasitic Diseases and Malaria
Unidad Colaborativa para Bioensayos Entomologicos
Campus de Ciencias Biologicas y Agropecuarias
AEF conceived the study and obtained funding
PMS and GPG carried out collections of the populations included in the study
All authors read and approved the final manuscript
The authors declare that they have no competing interests
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations
unless otherwise stated in a credit line to the data
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Jose Maria Barril returns to the Islanders for his third season
after joining the team on a transfer from Real Madrid in 2014
2015 (City Islanders): Barril led the team in assists with seven
The midfielder made 27 appearances for the club and recorded three goals off of eight shots
2014 (City Islanders): In his first season with the club
Barril appeared in 21 matches and scored two goals while notching one assist
spending over 14 years at their prestigious academy
participating in both reserve and first team matches
Cardel Benbow’s returns for his second season with the Islanders after a successful rookie campaign in 2015
Benbow scored two goals and had two assists while playing a total of 936 minutes
midfielder from Jamaica played three seasons for St
Georges College where he helped the team to a double championship
winning both the Manning Cup and Olivier Shield in 2012
He then started his club career with Waterhouse F.C
competing in the Jamaican National Premier League
Benbow has made a total of ten appearances for the Jamaican National Team in both Senior and Youth competitions
In 2014 he went 90 minutes on his debut vs
MLS club DC United in the CONCACAF Champions League
Cardel represented the Jamaica U-21 Team at the CAC games in Veracruz Mexico in 2014 scoring his first international goal against Honduras
Benbow made his senior international debut with the Reggae Boyz in October 2014 vs
Cardel represented his home nation at the U20 CONCACAF Championship held in Kingston
Mathew Bolduc will be playing his first season with the Islanders in 2016
At the University of Massachusetts in 2013
Bolduc started 12 out of the 16 matches during regular season
While playing he had a total of 18 shots seven of which were on goal
Bolduc was a four-year letter winner at The Winchendon School and at The Dublin School
During his junior and senior season he scored 36 goals
He was then named All-New England by the Boston Globe in 2011 and 2012
while earning NEPSAC All-Star honors as well
Throughout his entire high school career he scored 48 goals and 38 assists during regular season play
joins the Islanders this year following a season with the New York Cosmos in 2015
Broome was selected in the 3rd Round of the 2015 MLS SuperDraft by the San Jose Earthquakes but was ultimately not signed
Broome signed with NASL club New York Cosmos
That same season he played every minute of every game for the Providence Friars
securing a 2–1 win over Xavier in the Big East title game and subsequent NCAA triumphs over Dartmouth
Campbell joins the Islanders for his rookie season following a collegiate stint at Penn State University and University of Louisville
Campbell recorded his first career goal in victory over West Virginia as a sophomore in 2013
He was part of defense’s 275-minute shutout streak over four matches in 2013 and also scored in 2-1 win over Penn as a sophomore
Shane transferred to the University of Louisville where he started in 11 matches and appeared in 18 matches during first season with the Cardinals
He scored one goal and one assist during the season
In high school he was named first team All-State in 2011 at Dallastown Area HS
He also was a three- time YAIAA All-County Selection
Additionally he was a member of Philadelphia Union U-17/U-18 Academy for two season
He scored the game-winning goal in U-19 youth national championship game for Lehigh valley in the summer of 2016
Mouhamed Dabo makes his USL debut with the Islanders following stint with Inter Milan
Dabo was on trail with Arsenal for two months before he was offered a contract to Inter Milan B team when he was 16 years old
He went on to play for the inter Milan B team for three years
He was a member of the Inter B team that won the Italian youth cup and spent most of his time there training with the first team
Dabo made five appearances for the Senegal U-17 national team
After suffering a season-ending injury in 2015
Danny DiPrima will return for his third season with the Islanders in 2016
2015 (City Islanders): DiPrima scored one goal and played a total of 881 minutes
before suffering a season-ending knee injury
2014 (City Islanders): Danny helped lead the Islanders to the USL Championship Final
tallying 465 minutes on the pitch and netting one goal and one assist
John’s University before transferring to NC State University
During his three seasons with the Wolfpack
netting four goals and dishing out five assists
DiPrima had an outstanding youth career with Northern High School and the City Islanders Youth Academy
Danny was a four-year varsity starter and earned First-Team All-Conference honors as a junior and senior
He was also voted Conference Player of the Year as a junior
DiPrima captained the Polar Bears during both his junior and senior seasons
2016 MLS SuperDraft pick and Ohio State alumni
joins the City Islanders for the 2016 season
2015 was a big year for Doyle at Ohio State Univeristy
During the season he was named Big Ten Defensive Player of the Year
Preseason All-American by CollegeSoccerNews
he was third on the team with a total of 2 goals and 6 assists as well as starting every game
During the 2014 season with Ohio State he was named 2014 First Team All-Big Ten selection and second team NSCAA All-Midwest
He started all 22 games and had 5 goals and 2 assists for the season
he collected 13 goals and 13 assists where he was named MVP and Athlete of the Year as well as additional NCSCAA and NJCAA awards
His first season with Cincinnati State he scored 8 goals and had 10 assists
Darvin will make his debut appearance for the City Islanders this 2016 season
Ebanks attended Mount de Sales High School where he was a 5-star recruit by Top Drawer Soccer
and also the second best player in the South Atlantic Region according to Top Drawer Soccer
he attended University of North Carolina where he had a redshirt season
In 2015 he joined the National Premier Soccer League (NPSL) where he played in the South Region division for Georgia Revolution
Craig Foster returns for his second season with the Islanders in 2016
after join the squad midway through the season
and helping lead the team with 8 goals and two assists
2015 (City Islanders) Foster exceeded expectations last season and helped lead the team with 8 goals throughout the season and two assists
Foster scored 15 league goals for Reno FC and voted Jamaica Premier League Player of the Year in 2010/11
He spent the following two season on loan with Swedish club Motala AIF
where he scored 11 goals in 23 league appearances
Craig was named league MVP and also won the Golden Boot with 20 league goals in the Jamaica Premier League in the 2014/15 season
The 23-year-old striker went on to become a crucial member of the Jamaican National Team that competed at the 2011 CONCACAF U20 Championship held in Guatemala
Foster earned his first senior call-up in 2014 and was selected in the Jamaican squad that won the Caribbean Cup in 2014
Chris Hill makes his USL debut with the Islanders following a successful collegiate career at Villanova University
Hill appeared in 15 matches during his first season at Villanova in 2013 season
That season he led the team with three goals and three assists
In the 2014 season he started all 19 games and contributed a pair of assists and had a total of three shots on goal
Hill was also named to the BIG EAST All-Academic Team
Hughes joins the Islanders for his rookie USL season after making his professional debut with NASL side Atlanta Silverbacks
Hughes signed a professional contract with NASL side Atlanta Silverbacks
2015 in a 2–1 defeat to the Tampa Bay Rowdies
Hughes spent his collegiate career at Berry College
He made a total of 69 appearances for the Vikings and tallied 14 goals and six assists
Hughes was picked to the Southern Athletic Association all-conference team
He led Berry with nine goals and 19 points
Hughes also became Berry’s first student-athlete to earn Capital One Division III Academic All-America honors
He held a perfect 4.0 GPA and majored in biochemistry
Brett Jankouskas returnS to the club in 2016 for his second season
Jankouskas was one of the top scoring players
scoring 5 goals and recording one assist for the season
2015 (City Islanders): Jankouskas was one of the top scoring players
scoring 5 goals and notching one assist for the season
The former Palmyra High School All-American and Pennsylvania player of the year continued his collegiate career at Syracuse University
Shane Johnson returns for his third season with the Islanders
Johnson first joined the club on loan from the Richmond Kickers midway through the 2014 season and returned on loan in 2015
when he became a starter for the Islanders
2015 (City Islanders): Last season Johnson appeared in 22 games while starting in 17
2014 (City Islanders): While on loan from the Kickers
tallying a total of 180 minutes on the pitch before returning to Richmond
2013 (Richmond Kickers): Johnson logged 996 minutes of play in 17 matches
recording a goal and an assist in the process
2012 (Richmond Kickers): Appeared in all 22 matches
scoring his lone goal of the season against the City Islanders
Johnson played collegiate soccer at Longwood University
where he contributed to 17 shutouts in 71 appearances and two Atlantic Soccer Conference Championships
The 23-year old Bermudian international is back for his sophomore season with the Isles
2015 (City Islanders): Leverock appeared in 22 games for the Islanders
scoring three goals and contributing to 9 shutouts (4th in the USL)
Dante spent time with the Baltimore Blast before signing with the Islanders professional team
Leverock played for the City Islanders Academy under Denis Clark
assistant coach for the Harrisburg City Islanders
Dante has 6 caps and 1 goal since he was first brought onto the Bermudian National Team in March 2015
Leverock is currently representing Bermuda in the 2016 Caribbean Cup Qualifying Tournament
The Islanders welcome Shawn McLaws after a season with the New York Red Bulls and NYRB II
PROFESSIONAL EXPERIENCE 2015 (New York Red Bulls): McLaws was selected by the New York Red Bulls with the 17th pick of the third round
making him 59th overall in the 2015 MLS SuperDraft
As an unsigned draft pick in the preseason
he participated in camps and played 58 minutes in his debut versus USL side Oklahoma City Energy FC
McLaws played four seasons at Coastal Carolina University
ending his collegiate career with a 73-18-3 record
including three Big South regular season titles and two tournament championships
McLaws was selected by the New York Red Bulls with the 17th pick of the third round
making him 59th overall in the 2015 MLS SuperDraft
In 2014 he was named to College Soccer News Third Team All-American
NSCAA First Team All-South Atlantic Region
as well as Big South Preseason Defensive Player of the Year
This will be Youssef Naciri’s first season with the City Islanders Football Club
Youssef Naciri attended Christian Brothers University where he played midfielder and forward on the 2013 squad
During that 2013 season he scored one goal and had three assists for the team
Before college he attended LAK High School
The veteran goalkeeper returns to Harrisburg for his fifth season in the net for the Islanders
Noble comes off of a strong 2015 season in which he made 28 appearances
tallying a total of 2,475 minutes while recording 9 shutouts
Noble has made 104 appearances for the Islanders
Noble notched 28 appearances and nine shutouts
helping him surpass his 100th career appearance with the Islanders
2014 (City Islanders): He made 26 appearances and tallied 2486 minutes on the pitch
Noble helped lead his 8 seed squad to the USL Championship match against Sacramento
2013 (City Islanders): Noble's successful season moved him into the lead spot for multiple City Islanders all-time records
Noble spent a portion of his career competing in the MLS and in Europe
Noble was a first-round draft pick by the Chicago Fire in the 2007 MLS Supplemental Draft
and debuted for the Fire during the 2008 US Open Cup
After spending time with both the Chicago Fire and Austin Aztec for his first three professional seasons
Noble joined Ljungskile SK in Sweden for the 2010 and 2011 seasons
before he returned to the US and signing a contract with the LA Galaxy at the end of 2011
Noble is a graduate of West Virginia University
where he was a four-year standout net-minder
and was named the Big East Goalkeeper of the Year in 2006
Pettis returns to Harrisburg for his fourth year and is poised to produce a productive season on the pitch in 2016
Pettis dished-out one assist and scored 2 goals in 24 appearances
he appeared in 23 matches for the Islanders
tallying 671 minutes on the pitch while netting three goals and dishing-out two assists
2012 (City Islanders): Pettis appeared in 22 matches in his rookie campaign with the Islanders
scoring three goals and notching one assist before suffering a season-ending injury that would not allow him to return to the pitch until 2014
Pettis had a standout collegiate career at Lipscomb University in Nashville
He boasts the all-time goals record at Lipscomb
and earned the Atlantic Sun Freshman of the Year award
Pettis is a graduate of Lower Dauphin High School
and was a member of the City Islanders Youth Academy
Thomas joins the Islanders after spending the 2015 season with Eastern Conference rival
2015 (NYRB II): Thomas signed with the New York Red Bulls II for the 2015 season and made his debut as a starter for the team in its first ever match on March 28
He made 23 appearances for the USL club last season
Thomas played all four years of college ball at Saint Johns
During the 2011 season he started in all 23 matches and had one goal
Thomas started in 18 of the 19 matches and was named 2nd team All-Big East
a former MLS Super Draft first round pick and Mechanicsburg Area High School standout
will join the Islanders’ midfield after spending the last three years in Europe as a part of several different leagues
2015 (Hønefoss): Warshaw signed with First Division side Hønefoss in August 2015
2014 (Bærum): Warshaw moved to Norwegian First Division club Bærum in August 2014
2013 (FC Dallas): Warshaw scored eight goals in nine appearances while on loan with Swedish Superettan Club Angelholms before signing a one-year contract with GAIS of the same league
He went on to play 16 matches with GAIS and tallied two goals
he logged 450 minutes with the FC Dallas Reserves
2012 (FC Dallas): Bobby made his debut for FC Dallas in 2012
2011 (FC Dallas): Warshaw was selected in the first round (17th overall) by FC Dallas in the 2011 MLS SuperDraft
coming on as a substitute in a 0-0 draw with D.C
Warshaw was named Gatorade Player of the Year twice and was named the Pennsylvania Soccer Coaches Association’s Player of the Year in 2006
He started his college soccer career at Stanford University in 2007
starting in 17 matches and scoring five goals
he led Stanford to the Round of 16 in the NCAA tournament with wins over Saint Mary’s and UC Irvine in the first two rounds
he was a Hermann Trophy (USA College Player of the Year) semifinalist
despite being moved to defense for his final two season
The midfielder debuted for the USA U-17 National Team in 2005 against Honduras
He scored his first goal for the National Team in 2007 against Venezuela in the Pan-American Games held in Rio de Janeiro
USA won the match 2-1 thanks to Warshaw’s game-winner
Wheeler joins the club after two seasons with Major League Soccer’s Philadelphia Union and a successful 2015 campaign with the Wilmington Hammerheads
fourinch forward appeared in 15 matches during the Philadelphia Union’s 2014 MLS regular season
starting in eight matches for a total of 759 minutes
2015 (Wilmington Hammerheads): Appeared in 10 matches and recorded 2 goals for the USL squad
2014 (Philadelphia Union): Appeared in 15 games and started eight for a total of 759 minutes
Wheeler was a regular for much of the first half of the season
starting eight games in a stretch of nine from March 15 to May 3
only missing a midweek game against Real Salt Lake
He was named to the MLS Team of the Week by MLSSoccer.com in Week 4 after anchoring the backline to help the Union draw with Montreal
2013 (Philadelphia Union): Wheeler appeared in 10 matches for the Union
That same season he was loaned to the City Islanders for three matches where he scored two goals
Wheeler played for the Reading Rage in the USL Premier Development League during his collegiate career at Lenoir-Rhyne University and Towson University
He was the 2009 PDL Most Valuable Player and led the league in scoring with 17 goals
He later signed with the Vancouver Whitecaps before signing with FC Tampa Bay
The Islanders welcome rookie Paul Wilson to the squad for the 2016 season
Wilson began his career with Portmore United FC
He then went on to represent the club at the U17
He assisted in winning of multiple championships for the club
The graduate of Glenmuir High School represented Jamaica on the U17
where he claimed the ‘Most Outstanding Player’ award on the Jamaica team for the 2015 CAC Games in Mexico
He was also selected to participate at the Sunderland Academy in the United Kingdom
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